FLORA OF THE USSR

Volume XXVII

COMPOSITAE

Tribes Echinopsideae and Cynareae

bs
x een

FLORA OF THE USSR

Initiated under the supervision and chief editorship
of Academician V.L. Komarov

VOLUME XXVII
COMPOSITAE

Tribes Echinopsideae and Cynareae

Volume Editors

B.K. SCHISCHKIN AND F.G. BOBROV

General Scientific Editors
Stanwyn G. Shetler
Galina N. Fet

Translated from Russian

Smithsonian Institution Libraries
Washington, D.C.
1998

AS |
E 1% 6A,
z OF

SMIN B86 - SIL A001

Flora SSSR, Tom. XXVII
Akademiya Nauk SSSR Publishers,
Moscow-Leningrad, 1962

© 1998 Amerind Publishing Co. Pvt. Ltd., New Delhi

Translator: Dr. B.R. Sharma
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. —27: 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.

I. Botany—Soviet Union—Classification.
2. Plants—Identification. I. Komarov, V. L.
(Vladimir Leont evich), 1869-1945. II. Botanicheskii
institut im. V.L. Komarova. III. Title: Flora SSSR.
III. Title: Flora U.S.S.R. V. Title.
QK321.F69613 1985 581.947 85—904577
ISBN 3-87429-222-3 (v. 2)

Translated and published under an agreement for the Smithsonian
Institution Libraries, Washington, D.C., by Amerind Publishing Co. Pvt.
Ltd., 66 Janpath, New Delhi 110001

Typeset at Indira Printers, New Delhi and printed at Rekha Printers Pvt. Ltd.,
New Delhi 110 020.

V.L. KOMAROV BOTANICAL INSTITUTE
ACADEMY OF SCIENCES OF THE USSR

Contributors

E.G. Bobrov, V.P. Botschantzev, M.M. Iljin,
I.A. Linczevsky, S.Yu. Lipschitz,
E.V. Sergievskaja, O.V. Czerneva,
S.K. Czerepanov, S.V. Juzepczuk

INSTITUTUM BOTANICUM nomine V. KOMAROVII ACADEMIAE SCIENTIARUM URSS

FLORA
URSS

(FLORA UNIONIS RERUMPUBLICARUM
SOCIALISTICARUM SOVIETICARUM)

XXVII

EDITIO ACADEMIAE SCIENTIARUM URSS
MOSQUA . 1962. . LENINGRAD

SCIENTIFIC EDITOR’S PREFACE TO VOLUME XXVII

This is the third volume of the six (Vols. XXV—XXX) that treat the
large family Compositae (Asteraceae). I have, as a practical matter,
concentrated my editorial review of this volume on the discussions and
the habitat and distribution statements. It is in these parts of the text
that the specific rendering in English is the most critical. We are
confident of the general accuracy of the translation but also recognize
that there will be imperfections.

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
secondary inflorescence, i.e., an arrangement of capitula.

Galina N. Fet also has reviewed this volume in its entirety, paying
particular attention to the geographic and place-name terminology in
the distribution statements following the morphological descriptions. Her
corrections of the geographical information and other editorial suggestions
have added substantially to the accuracy of the translated work.

STANWYN G. SHETLER

Curator of Botany Emeritus
National Museum of Natural History
Smithsonian Institution

Washington, DC 20560

October 1997

Assisted by Galina N. Fet
Huntington, West Virginia 25701

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CONTENTS

SCIENTIFIC EDITOR’S PREFACE TO VOLUME XXVII__.... Vil

PREFACE = xi

SYSTEMATIC INDEX OF THE SPECIES OF THE on xili

TWENTY-SEVENTH VOLUME OF THE “FLORA OF

THE USSR”

TRIBE 11. Echinopsideae Cass. i 1
Genus 1567. Echinops L. ae 2
Genus 1568. Acantholepis Less. ae 55

TRIBE 12. Cynareae Less. he 5!)

Subtribe 1. Carlininae O. Hoffm. ne 58
Key to Genera of Subtribe Carlininae ae 58
Genus 1569. Cousiniopsis Nevski ut. 32)
Genus 1570. Xeranthemum L. ne 61
Genus 1571. Siebera J. Gay ne 69
Genus 1572. Chardinia Desf. ae qk
Genus 1573. Amphoricarpus Vis. ae 74
Genus 1574. Carlina L. bee 79
Genus 1574a. Thevenotia DC. sai 90
Genus 1575. Atractylodes DC. ane 90
Subtribe 2. Carduinae O. Hoffm. ue 92
Key to Genera of Subtribe Carduinae mes 93
Genus 1576. Arctium L. sae 96
Genus 1577. Anura (Juz.) Tschern. iN 109
Genus 1578. Cousinia Cass. ie 110

Genus 1579. Lipskyella Juz. at 352

X

Genus 1580. Hypacanthium Juz.
Genus 1581. Schmalhausenia Winkl.
Genus 1582. Saussurea DC.

Subgenus
Subgenus

Subgenus

1. Stephanodontos Lipsch.

2. Amphilaena (Stschegl.) Lipsch.

3. Frolovia (DC.) Lipsch.

Subgenus 4. Eriocoryne Wall. ex DC.

Subgenus

Subgenus

Genus 1583.
Genus 1584.
Genus 1585.

Genus 1586.
Genus 1587.
Genus 1588.

5. Saussurea

6. Theodorea (Cass.) Lipsch.
Polytaxis Bge.

Jurinea Cass.

Jurinella Jaub. and Spach
Perplexia in

Pilostemon I\jin

Nikitinia jin

ADDENDA XXVI
INDEX ALPHABETICUS

354
3955
356
a75
376
378
382
385
514
530
535
697
701
702
705
707
123

NG PREFACE

The twenty-seventh volume contains descriptions of 22 genera comprising
636 species of groups of Compositae according to the accepted
classification.

This volume includes results of analysis of materials of the small
tribe Echinopsideae and a large part of the tribe Cynareae. The analysis
of the latter tribe is concluded in the next volume (Volume XXVIII). A
large part of that volume is devoted to the tribe Cynareae.

The present volume provides an analysis of some of the largest
Compositae genera of the indigenous flora: Cousinia, Jurinea, Saussurea,
and Echinops. The analysis of these genera, of course, required much
time and effort, and provided new information for a more complete
understanding. It suffices to say that this volume lists 272 of the total
350 Cousinia species, 146 of 300 Jurinea species, 115 of 400 Saussurea
species, and 58 of 120 Echinops species.

Dozens of new taxa were formally described in various local
publications during the course of analysis of the material and only about
30 descriptions are included in the Latin appendix to the present volume.

Editors.

‘Refers to Russian original page number—General Editor.

hy eich sth Gi saboaaas m

Ahem

SYSTEMATIC INDEX OF THE SPECIES IN THE TWENTY-

SEVENTH VOLUME OF THE “FLORA OF THE USSR”*

Tribe 11. Echinopsideae Cass.
Genus 1567. Echinops L.

Section 1. Oligolepis Bge.

15052. 1. E. transcapicus Bornm.
2. E. maracandicus Bge.
3. E. polygraphus Tschern.
4. E. brevipenicillatus Tschern.
5. E. leucographus Bge.
6. E. lelopolyceras Bornm.
7. E. praetermissus Nevski
8. E. lipskyi Ijin
Section 2. Ritrodes Bge.
15060. 9. E. ritrodes Bge.
10. E. multicaulis Nevski
11. E. albicaulis Kar. and Kir.
12. E. kafirniganus Bobr.
13. E. dubjanskyi \ljin
14. E. subglaber Schrenk
15. E. pubisquameus {in
16. E. dasyanthus Rgl. and Schm.
17. E. tjanschanicus Bobr.
18. E. chantavicus Trautv.
15070. 19. E. karatavicus Rgl. and Schm.
20. E. obliquilobus Ijin
21. E. babatagensis Tschern.
22. E. fedtschenkoi (jin

10
11
1]
12
14
14
lis)
16

16
17
18
18
19
20
20
2a
22
22
23
23
24
2S

“Reproduced from the Russian original. Russian page numbers appear in the left-hand

margin of the text—General Editor.

VI

X1V

15080.

15090.

15100.

Ze
24.
2S:
26.

27:
28.
WS).
30.
Sie
32:
33.
34.
Sy
36.
a7.
38.

3).

40.
Al.
42.
43.
44.
45.
46.
ai,
48.
49.
50.

ot

32
53:

orientalis Trautv.
opacifolius Yin
connatus C. Koch

macrophyllus Boiss. and Hausskn.

Section 3. Ritro Endl.

ritro L.

. meyeri (DC.) Ijin
. Spiniger \ljin ex Bobr.

pungens Trautv.
polyacanthus {jin
conrathi Freyn

foliosus Somm. and Lev.
transcaucasicus [lyin
szovitsii Fisch. and Mey.

. ijinii Mulk.
. sevanensis Mulk.
. karabachensis Mulk.

Section 4. Pleiacme Bornm.

tournefortii Ldb.

Section 5. Echinops

. Sphaerocephalus L.

erevanensis Mulk..-
dagestanicus I\jin

. viridifolius Ijin

galaticus Freyn
colchicus Sosn.
ossicus C. Koch

. latifolius Tausch

. dissectus Kitag.

. tricholepis Schrenk
. abstersibilis Tin

Section 6. Terma Endl.

. exaltatus Schrad.

Section 7. Chamaechinops Bge.

integrifolius Kar. and Kir.
humilis MB.

25
26
Df
28

30
32
32
35
34
34
35
36
36
37
38
38

39

40
41
41
42
43
43
44
44
45
46
46

47

48
50

Vil

15110.

15120.

54. E. tschimganicus B. Fedtsch.
55. E. saissanicus (Keller) Bobr.
Section 8. Nanechinops Bge.

56. E. gmelini Turcz.
57. E. nanus Bge.
58. E. knorringianus Ijin

Genus 1568. Acantholepis Less.

1. A. orientalis Less.

Tribe 12. Cynareae Less.

Subtribe 1. Carlininae O. Hoffm.

Genus 1569. Cousiniopsis Nevski

1. C. atractyloides (Winkl.) Nevski
Genus 1570. Xeranthemum L.

Section 1. Xeranthemum

. X. annuum L.
. X. squarrosum Boiss.
. X. inapertum (L.) Mill.

hwWN —

Section 2. Xeroloma (Cass.) DC.

5. X. cylindraceum Sm.

Genus 1571. Siebera J. Gray
1. S. pungens (Lam.) DC.

Genus 1572. Chardinia Desf.

1. C. orientalis (L.) Ktze.
2. C. macrocarpa C. Koch

Genus 1573. Amphoricarpus Vis.

Section. 1. Chodatella Alb.
1. A. elegans Alb.

. X. longipapposum Fisch. and Mey.

XV

50
51

a2
52
53

54

58

62
63
64
65

66
68

70
71

75

Vill

Xvi

15130.

—

N

ae iS
© Oey CHe@me@

ee a
Sw mw ww

Genus 1574. Carlina L.

vulgaris L.

. biebersteinii Bernh. ex Hornem.
lanata L.

. onopordifolia Bess. ex. Szafer, Kulcz.,
Pawl.

. acaulis L.

. cirsioides Klok.

Genus 1575. Atractylodes DC.
. A. ovata (Thnb.) DC.

Subtribe 2. Carduinae O. Hoffm.
Genus 1576. Arctium L.

Section 1. Eglandulosa Aréne

. platylepis (Boiss. and Bal.) Sosn.
. lappa L.

. leiospermum Suz. and C. Serg.

. nemorosum Le}.

. minus Bernh.

Section 2. Glandulosa Aréne

. A, palladinii (Marc.) Grossh.
. A, tomentosum Mill.
. A. radula Juz. and C. Serg.

Genus 1577. Anura (Juz.) Tschern.
. A. palidivirens (Kult.) Tschern.

Genus 1578. Cousinia Cass.

Section 1. Badghysia Tschern.
. C. badghysi Kult.

Section 2. Leucocaulon Tschern.

. C. turcomanica Wink.
. C. linczewskii Juz.

M9
82
83

84
86
87

89

95
97
98
99
100

103
104
106

107

144

145
145

15140.

15150.

15160.

29:

30.

ah
32.
33.

Section 3. Leptocephalae Tschern.

OVID

QO OA OV Oo OVO TEVIGioKic

DON DAO} (9) 4) 1

. leptocephala Fisch. and Mey.
. albiflora (Bornm. and Sint.) Bornm.
. hypopolia Bornm. and Sint.

mucida Kult.

. stahliana Bornm. and Gauba
. oreoxerophila Kult.
. chaetocephala Kult.

Section 4. Leiocaules Bge.

. astracanica (Spreng.) Tamamsch.

affinis Schrenk

. talassica (Kult.) Juz.

pseudoaffinis Kult.
hamadae Juz.
xanthiocephala Tschern.

. crytadena Juz.
. antonowii Wink}.
. glandulosa Kult.

Section 5. Chrysoptera Tschern.

dissecta Kar. and Kir.

. perovskiensis (Bornm.) Juz.
. sogdiana Bornm.

dolichoclada Juz.
alata Schrenk
oxiana Tschern.

. psammophila Kult.
. Schisoptera Juz.

turkmenorum Bornm.

Section 6. Olgaeanthe Tschern.

. olgae Rgl. and Schm.

Section 7. Chrysantha Tschern.

. pygmaea Winkl.

Section 8. Actinia Bge.

albertoregelia Winkl.
bobrovii Juz.
oopoda Juz.

XVil

146
147
148
150
150
150
151

152
153
154
155
155
156
157
158
158

160
161
161
162
163
163
164
165
165

166

167

168
169
170

XVil1

1570. 34.
a5:
36.

Be
38.
32

40.

41.

42.
43.
15180. 44.

45.

46.

47.
48.
49.
50.

Sik.
52:

33°
15190. 54.

C. gontscharowii Juz.
C. stricta Tschern.
C. sclerophylla Juz.
Section 9. Pseudactinia Tschern.

C. oreodoxa Bornm. and Sint.
C. apiculata Tschern.
C. glochidiata Kult.
Section 10. Hoplophylla Tschern.

C. raddeana Winkl.

Section 11. Nanarctium Juz.

C. arctioides Schrenk

Section 12. Pseudarctium Juz.

. umbrosa Bge.
. tomentella Winkl.
. pseudarctium Bornm.

DQ) O) ©)

Section 13. Lappaceae Bge.
C. lappacea Schrenk

Section 14. Ctenarctium Juz.

C. anomala Franch.

Section 15. Molles Bge.
C. pseudomollis Wink.
C. leptacantha (Bornm.) Juz.
C. fascicularis Juz.
C. mollis Schrenk
Section 16. Sciadocousinia Tschern.
C. eryngioides Boiss.
C. margiana Juz.

Section 17. Cousinia

Subsection 1. Irano-armeniacae Takht.

C. chlorocephala C.A.M.
C. daralaghezica Takht.

170
Mii
173

174
174
175

176
177
178
179
179
180

182

183
184
184
185

186
186

188
189

15200.

15210.

55
56

63.
64.
65.
66.
67.
68.
69.
70.
TK.
13
73.
74.
WSs
76.
Wa:
78.
79:
80.
81.
82.
83.
84.
85.
86.
87.
88.
89.
90.
Oe
92.

nie.
: G
Bi).
58.
So:
60.
61.
62.

OLOMO@V eM e)

Subsection 2. Irano-turkestanicae Takht.

. orientalis (Adams) C. Koch
. microcephala C.A.M.
. dolicholepis Schrenk
. vicaria Kult.

POLSON ONe TS OMOTeTe Te Te TOMeTeowoiortorte! Clie fee) elelie) erie!

armena Takht.
fedorovii Takht.

. brachyptera DC.

. araxena Takht.

. erivanensis Bornm.
. Ujinii Takht.

. lomakinii Winkl.

. hohenackeri Fisch. and Mey.

angreni Juz.
scabrida Juz.
minkwitziae Bornm.
tenuisecta Juz.
syrdariensis Kult.

turkestanica (Rgl.) Juz.

juzepczukii Tschern.
umbilicata Juz.
pungens Juz.
scleracantha Kult.
polycephala Rupr.
finitima Juz.

. Sporadocephala Juz.

spiridonovii Juz.
transoxana Tschern.
ambigens Juz.
resinosa Juz.

kazachorum Juz. ex Tschern.

horridula Juz.
maracandica Juz.
platystegia Tschern.
dichromata Kult.
egens Juz.

. congesta Bge.
. agelocephala Tschern.
. androssovii Juz.

X1X

189
190
122
192
193
194
194
195

196
196
197
198
199
199
200
202
202
203
203
204
205
205
206
207
207
208
208
209
210
212
212
213
214
214
215
216
216
27,

XI

XX

15230.

15240.

15250.

15260.

QO)

yO) (A) ©

OOO Or) ©)

OY CVOr GO @Ve) OV) ©

Section 18. Abolinia Tschern.

C. abolinii Kult.
C.
C. dolichophylla Kult.

egregia Juz.

Section 19. Eriocusinia Tschern.

. pannosa Winkl.

franchetii Winkl.

. pannosiformis Tschern.

semidecurrens Winkl.
fedtschkenkoana Bornm.

. korshinskyi Wink\.
. pterolepida Kult.
. macilenta Wink.

Section 20. Homalochaete Juz.

leptacma Tschern.

| litvinovii Kult.

amoena Winkl.

. campyloraphis Tschern.

proxima Juz.
subcandicans Tschern.
candicans Juz.
podophylla Tschern.
leptocladoides Tschern.
gnezdilloi Tschern.

. leptoclada Kult.
. hoplophylla Tschern.

corymbosa Winkl.

. coerulea Kult.

Section. 21. Lopholepis Tschern.

. rigida Kult.
. mindshelkensis B. Fedtsch.
. gomolitzkii Juz.

Section. 22. Pectinatae Winkl.

. albertii Rgl. and Schm.

. pentacantha Rgl. and Schm.
. horrescens Juz.

. pentacanthoides Suz.

218
218
219

220
221
222
222
224
225
225
226

227
227
228
228
229
230
230
232
232
233
234
234
235
235

236
237
238

239
239
240
241

XII

15270.

15280.

125.

126.
127:
128.
129.
130.
13k:
132.
133.

134.

135)

136.
1a.
138.
139.
140.
141.
142.

143.
144.
145.
146.
147.

148.

149.

Section 25. Serratulopsis Tschern.

C.

Section 26. Amberbopsis Tschern.

Cc:

SOUS) Oy or or

SESE OOS oeose)

SSNS) @) Loic 10}

Section 23. Oligantha Juz.

. triflora Schrenk

Section 24. Chrysis Juz.

aurea Winkl.

medians Juz.

refracta Juz.
schmalhausenii Winkl.
karatavica Rgl. and Schm.

. haesitabunda Juz.
. Chlorantha Kult.

korolkovii Rgl. and Schm.

vavilovii Kult.

grandifolia Kult.

Section 27. Stenoloma Juz.

. komarowii (O. Ktze.) Winkl.
. stenophylla Kult.
. leucantha Bornm. and Sint.

grisea Kult.
alaica Juz.
fallax Winkl.
baranovii Juz.

Section 28. Kuekenthalia Juz.

kuekenthalii Bornm.
neglecta Juz.
dissectifolia Kult.

. decurrentifolia Juz.
. ulotoma Bornm.

Section 29. Lasiandrae Bge.

. lasiosiphon Juz.

Section 30. Cynaroideae Bge.

. onopordioides Ldb.

XXi

242

244
245
245
246
246
247
247
248

249

249

250
251
252
252
254
255
255

256
256
257
257,
258

259

260

XXII

15290.

15300.

15310.

150.
tsuile
152°
L353:
154.
55:

156.
157.

158.

159.
160.
161.
162.
163.
164.
165.
166.
167.
168.
169.
170.

7s

GP,

173.
174.
175.

C. cynaroides (MB.) C.A.M.
C. macrocephala C.A.M.

C. lyrata Bge.

C. purpurea C.A.M.

C. freynii Bornm. and Sint.
C. beckeri Trautv.

Section 31. Kopetdagia Tschern.

C. smirnowii Trautv.
C. botschantzevii Juz.

Section 32. Helianthae Bge.
C. spryginii Kult.

Section 33. Dichotomae Bge.

. dichotoma Bge.

. erectispina Tschern.
polytimetica Tschern.
. patentispina Tschern.
sylvicola Bge.
bungeana Rgl. and Schm.
pusilla Winkl.
orthacanha Tschern.
tedshenica Tschern.
murgabica Tschern.

. angusticeps Juz.

. eugenii Kult.

VCO) OO Vo

Section. 34. Xiphiolepides Bge.

C. hystrix C.A.M.

Section. 35. Pugioniferae Bge.

C. macroptera C.A.M.

Section 36. Microcarpae Bge.

Subsection 1. Coronophora Juz.

C. radians Bge.
C. coronata Franch.
C. pulchra Winkl.

262
263
263
264
265
265

266
267

268

269
269
270
270
22
272
DIS
2H
274
274
275
275

276

20%

278
279
280

XII

15320.

15330.

15340.

15350.

176.
Li 7.
178.

179.
180.
181.
182.
183.
184.
185.
186.
187.
188.
189.
190.
191.
192.
193%
194.
195:
196.
Ws
198.
199.
200.
201.
202.
203.
204.
205.
206.
207.
208.
209.
210.
2M.
212.
213.
214.
PANS),
216.

See)

SUP SUG @ Ole) OLe Toren erere ever ey ere) ley erevelorerero oleic joie

. alpestris Bornm.
. mulgediifolia Bornm.
. leptocampyla Bornm.

Subsection 2. Carduncellus Juz.

. alpina Bge.
. carduncelloides Rgl. and Schm.

outichaschensis Franch.
stellaris Bormmm.
buphtalmoides Rgl.
hilariae Kult.
knorringiae Bornm.
grigoriewii Juz.

rava Winkl.

glabriseta Kult.

triceps Kult.

. fetissowii Winkl.

schischkinii Juz.

speciosa Winkl.

batalinii Winkl.

calva Juz.

oxytoma Rech. f.

ovcezinnikovii Tschern.
peduncularis Juz.

gulczensis Kult.
schtschurowskiana Rgl. and Schm.

pseudodshisakensis Tschern. and Vved. ...

dshisakensis Kult.
princeps Franch.

. laniceps Juz.
. tianschanica Kult.

mogoltavica Tschern. and Vved.
butkovii Tschern. and Vved.
praestans Tschern. and Vved.

. strobilocephala Tschern. and Vved.

rosea Kult.

. rhodantha Kult.
. rotundifolia Winkl.

C. ferruginea Kult.

c
C.
C.
C.

dimoana Kult.
darwasica Winkl.
integrifolia Franch.
divaricata Winkl.

XX1il

282 ©
282
283

284
285
285
287
287
288
289
290
290
292
292
293
294
294
295
295
296
296
297
22355)
298
299
300
300
302
303
303
304
305
305
306
307
307
308
309
309.
310
310

XIV

XX1V

15360.

15370.

15380.

217. C. centauroides Fisch. and Mey.

Subsection 3. Kokuria Juz.

218. C. severtzovii Rgl.

219. C. leiocephala (Rgl.) Juz.

220. C. pulchella

221. C. pauciramosa Kult.

222. C. regelii Winkl.

223. C. arachnoidea Fisch. and Mey.
224. C. microcarpa Boiss.

225. C. ninae Juz.

226. C. platylepis Schrenk

227. C. ortholepis Juz.

Subsection 4. Subappendiculatae Tschern.

228. C. ferganensis Bornm.
229. C. subappendiculata Kult.
230. C. simulatrix Wink.

231. C. wedenskyi Tschern.

Section 37. Microcousinia Tschern.

232. C. minuta Boiss.

Section 38. Jurineopsis (Juz.) Tschern.

233. C. krauseana Rgl. and Schm.
234. C. submutica Franch.

235. C. tamarae Juz.

236. C. omphalodes Tschern.
237. C. dubia M. Pop.

238. C. kokanica Rgl. and Schm.
239. C. hastifolia Winkl.

240. C. transiliensis Juz.

241. C. waldheimiana Bormnm.
242. C. margaritae Kult.

243. C. jassyensis Winkl.

Section 39. Tenellae Bge.
244. C. tenella Fisch. and Mey.

Section 40. Glaphyrocephalae Tschern.
245. C. glaphyrocephala Suz.

S12

313
314
314
StS
316
316
lig
318
318
S18)

320
320
322
323

324

324
325
326
326
327
328
328
329
330
330
332

333

334

XVIi

XVII

15390.

15400.

15410.

Section 41. Alpinae Bge.

246. C. magnifica Juz.

247. C. lanata Winkl.

248. C. pseudolanata M. Pop.
249. C. adenophora Juz.

250. C. splendida Wink.
251. C. xanthina Bornm.
252. C. trachyphylla Juz.
253. C. chrysantha Kult.
254. C. stephanophora Wink.
255. C. semilacera Juz.

256. C. allolepis Tschern. and Vved.
257. C. newesskiana Winkl.
258. C. trichophora Kult.
259. C. schugnanica Juz.
260. C. sororia Juz.

261. C. abbreviata Tschern.
262. C. ramulosa Rech. f.
263. C. acrodroma Tschern.
264. C. rubiginosa Kult.

265. C. multiloba DC.

266. C. eriotricha Juz.

267. C. laetevirens Winkl.
268. C. verticillaris Bge.
269. C. bonvalotii Franch.
270. C. ceratophora Kult.
271. C. serawschanica Winkl.
272. C. caespitosa Wink.

Doubtful Species
. bucharica Winkl.

VOTO

. xiphiolepis Boiss.

Genus 1579. Lipskyella Juz.
1. L. annua (Winkl.) Juz.

Genus 1580. Hypacanthium Juz.
1. H. echinopifolium (Bornm.) Juz.

. trachylepis var. integrifolia O. Ktze.

XXV

334
335
336
337
337
338
339
340
340
342
343
344
345
346
346
347
348
348
349
350
352
352
352
354
355
355
356

357
35)/
35)

358

360

XVI

XXV1

15420.

15430.

Genus 1581. Schmalhausenia Winkl.
1. S. nidulans (Rgl.) Petrak

Genus 1582. Saussurea DC.

Subgenus 1. Stephanodontos Lipsch.

1. S. carduicephala (Ijin) Ijin

Subgenus 2. Amphilaena (Stschegl.) Lipsch.

2. S. involucrata Kar. and Kir.

aS

8.
2)
10.
jul
12
13.

Subgenus 3. Frolovia (DC.) Lipsch.

3. S. frolovii Ldb.
4. S. asbukinii lin
5. S. sulcata Vin

Subgenus 4. Eriocoryne Wall. ex DC.

oy
ney

ANnNNHHHYN

glacialis Herder

gnaphalodes (Royle) Sch. Bip. ex Klatt...

Subgenus 5. Saussurea

Section 1. Pycnocephala Lipsch.

. baicalensis (Adams) Robinson
. dorohostaiskii Palibin
. krylovii Schischk. and Serg.

schanginiana (Wydl.) Fisch. ex. Herd.

. leucophylla Schrenk
. sordida Kar. and Kir.

Section 2. Laguranthera (C.A.M.) Lipsch.

14.
15.
16.
17.
18.
19.
20.
21.
22%

M~HAANHANANANRNHN

. rigida Ldb.
. ninae Ijin
. coronata Schrenk

blanda Schrenk

. mikeschinii Tin

. ispajensis Ijin

. Salicifolia (L.) DC.
. lomatolepis Lipsch.
. cana Ldb.

361

382

383

384
387
388

389
390

393
394
396
By /
398
400

403
404
406
407
408
409
410
412
413

15440.

15450.

15460.

23%
24.
25:
26.
Pale
28.
29.
30.
31.
32.
33.

34.

S.

ANNNNHANNNHNKNANNANNNNNNNHAY

ANNNNHRHANHUNNHN

. Sumneviczii Serg.
. serratuloides Turcz.
. caespitans Tin

canescens Winkl.

vvedenskyi Lipsch.
salemanii Wink}.

elata Ldb.

. elegans Ldb.

. larionovii Wink.

. alberti Rgl. and Winkl.
. pulviniformis Wink.

Section 3. Benedicta DC.

Subsection 1. Odontolepis Kitam.

odontolepis Sch. Bip.

Subsection 2. Hololepis Lipsch.

. jadrinzevii Kry]l.
. tomentosa Kom.
. porcellanea Lipsch.

Kitamura Miyabe and Tatewaki
sajanensis Gudoschnikov

. sovietica Kom.

Squarrosa Tutcz.

pseudosquarrosa M. Pop. and Lipsch.

controversa DC.

. discolor (Willd.) DC.

hypargyrea Lipsch. and Vved.

. ajanensis (Rgl.) Lipsch.

lenensis M. Pop. ex Lipsch.
S. uralensis Lipsch.
splendida Kom.

elongata DC.

. recurvata (Maxim.) Lipsch.

ussuriensis Maxim.
maximoviczii Herd.
nupuripoiensis Miyabe and Miyake

. kurilensis Tatewaki

. riederi Herd.

. acuminata Turcz.

. sachalinensis Fr. Schm.

XXVIl

414
416
417
418
420
421
422
423
425
426
427

428

429
430
431
432
433
434
436
437
438
439
440
441
441
442
443
444
445
446
448
450
452
453
454
455

XXVill

15470.

15480.

15490.

15500.

15510.

oS)
96.
ile

98
99

ANANNDNANHANNUNANRHUHUANDUNAHHRHRUAHHHAUHHRHRHDUHHANHAUNY

. amurensis Turcz.

. duiensis Fr. Schm.

. Shiretokoensis Sugaw.
. dubia Freyn

umbrosa Kom. —
grandifolia Maxim.
subtriangulata Kom.
sinuata Kom.

. triangulata Trautv. and Mey.

manshurica Kom.
parviflora (Poir.) DC.
porcii Degan

. neoserrata Nakai

fauriei Franch.

. latifolia Ldb.

foliosa Ldb.

. pricei Simps.

congesta Turcz.

. poljakovii Glehn
. tilesii Ldb.

soczavae Lipsch.
pseudo-tilessii Lipsch.
oxyodonta Hult.

. alpina (L.) DC.

sukaczevii Lipsch.
esthonica Baer

. Stubendorfii Herd.

ambigua Kryl.
saxosa Lipsch.
angustifolia (Willd.) DC.

. viscida Hult.
. pseudoangustifolia Lispch.
. kuschakeviczii Winkl.

ovata Benth.

. salsa (Pall.) Spreng.

. paradoxa Lipsch.

. daurica Adams

. turgaiensis B. Fedtsch.
. kaschgarica Rupr.

. S. faminziana Krassn.

. S. karaartscha Saposchn.

456
458
459
460
462
464
465
468
469
471
472
474
475
476
478
479
480
482
483
484
486
487
488
489
492
493
495
496
497
498
500
501
502
503
504
506
507
508
509
510
511

XVII 15520.

15530

XX1X

Section 4. Jurineiformes (Lipsch.) Lipsch.
100. S. chondrilloides Wink. ee 512

Section 5. Jacea Lipsch.

101. S. masarica Lipsky PB eS
102. S. tadshikorum Ijin Soe 514

Section 6. Depressae Clarke

103. S. nuda Ldb. ih 515
104. S. tschuktschorum Lipsch. ° Me 518
105. S. caprifolia Ijin At 519

Subgenus 6. Theodorea (Cass.) Lipsch.

106. S. amara (L.) DC. ae 520
107. S. pulchella Fisch. se 522
108. S. neopulchella Lipsch. ae 525
109. S. alata DC. oa 526
110. S. crepidifolia Turcz. Ae Sd)
111. S. runcinata DC. ae 528
112. S. laciniata Ldb. ae 529
113. S. ceterahifolia Lipsch. ne 532
114. S. robusta Ldb. acs: 555
115. S. prostrata Wink. ne 534
Genus 1583. Polytaxis Bge.
1. P. lehmannii Bge. a 536
2. P. winkleri \\jin ue ay,
Genus 1584. Jurinea Cass.
Section. 1. Cyanoides (Korsh.) Ijin
1. J. cyanoides (L.) Rchn. ie 546
2. J. pseudocyanoides Klok. aes 547
3. J. schischkiniana \\jin Ht 548
4. J. xerophytica Ijin ies 549
5. J. eversmannii Bge. ae 549
6. J. charcoviensis Klok. a 550
7. J. granitica Klok. Ai 552
8. J. salicifolia Gruner ee 352
9. J. albicaulis Bge. ie 554

XIX

XXX

15540.

15550}

15560.

15570.

32)
32a.

Section 3. Stenocephalae Benth. and Hook.

See
34.
3.
36.

SANNA NNN

SNNNNNNN

SSN NNNNNNNNSS

J:

vf
Ji
J.

gorodkovii Ijin ©

creticola Ijin

laxa Fisch.

amplexicaulis (S.G. Gmel.) Bobr.

. altaica {lin

transuralensis jin

. ciscaucasica (Sosn.) [jin

kazachstanica Ijin

. pineticola Ijin

paczoskiana I\jin

. tenuiloba Bge.
. tanaitica Klok.
. thyrsiflora Klok.

Section 2. Bellae Iljin

pumila Alb.
woronowii Ilin
bellidioides Boiss.
iljinii Grossh.
venusta Ijin
sosnovskyi Grossh.
annae Sosn.
levieri Alb.

. coronopifolia Somm. and Lev.
. cartaliniana Boiss.

filicifolia Boiss.

multiflora (L.) B. Fedtsch.
stoechadifolia (M.B.) DC.
pulchella (Fisch. and Mey.) DC.
kapelkinii O. Fedtsch.

Section 4. Integrae [lyin

. abolinii Thin

. Schachimardanica {in
. Stenophylla Mjin

. caespitans lyin

eduardi-regelii \\jin
capusii Franch.

. tortisquamea Ijin
. androssovii Ijin

554
555
556
558
559
559
561
562
563
564
565
566
567

569
570
570
571
302
574
574
575)
576
a7
Sa

379
580
581
582

584
585
586
587
587
588
590
590

15580.

15590.

15600.

45.
46.
47.
48.

49.
50.
51.
52.
D3:

54.
a5:

56.

She
58.
Soh

60.
61.
62.
63.
64.
65.
66.
67.
68.
be:
70.
wl.
M2.

macranthodia Ijin
kokanica Vin
rhizomatoides I\jin
trifurcata [lyin

SNNN

Section 5. Suffrutices I]jin

. suffruticosa Rgl.

. multiceps Min

. mugodsharica {jin
. sintenisii Bornm.

. orientalis Ijin

SNNNN

Section 6. Floccosae (Sosn.) Ijin

J. armeniaca Sosn.
J. elegans (Stev.) DC.

Section 7. Penduliflorae Iljin

Subsection 1. Robustae [lyin

J. robusta Schrenk

Subsection 2. Pappilligerae Ijin

J. komarovii jin
J. kamelinii Win
J. elegantissima I\jin

Subsection 3. Persistentes Iljin

. trautvetteriana Rgl. and Schm.
. gracilis Ijin

. popovii Tin
cephalopoda I\jin

. bucharica Wink\l.

. psammophila Ijin

. lasiopoda Trautv.

. karatavica {hin

. botschantzevii Wyin
. winkleri Win

. tadshikistanica in
. impressinervis lin
. apoda Ijin

i

XXXi

591
592
593
593

>) /5)
596
596
DoT
598

600
601

605

606
607
608

608
609
610
610
611
612
614
615
615
616
617
618
619

| XXXil

XX

15610.

15620.

15630.

73.
74,
75.
76.

77.
78.
79.
80.
81.
82.
83.
84.
85.
86.
87.
88.
89.
90.
91.

Section 10. Derderia (Jaub. and Spach) Boiss.

92°
O85:
94.
95.
96.

Oe
98.
ve).
100.

101.
102.
103.
104.

SANS

WANA NS

SANS

NWN SN

SN NNN NNN NNN NNN SN

Section 8. Olgae Iljin
olgae Rgl. and Schm.

ferganica (Ijin) Ijin

kuraminensis I\jin
xeranthemoides I\jin

Section 9. Insculptae Iljin

maxima Winkl.

. bipinnatifida Winkl.
. pjataieviae lljin

. serpenticaulis I\jin
. asperifolia Ijin

propinqua {hin

. helichrysifolia M. Pop.
. zakirovii Ijin

semenovii Winkl.
hamulosa Rubtz.
lithophila Rubtz.

. serratuloides Iljin

. atripurpurea Winkl. ex Iljin
. sangardensis I\jin

. baissunensis Iljin

. auscheriana DC.

. derderioides Winkl.
. transhyrcanica \\jin
. eximia Tek.

. karabugasica I\jin

Section 11. Stechmannia Boiss.

. antonovii Winkl.
. kultiassovii Win
. lydiae Ijin

. ludmilae Ijin

Section 12. Nanae Iljin

. abramovii Rgl. and Herd.
. tapetodes Ijin

. bobrovii Ijin

. antoninae Ijin

620
621
622
623

627
628
629
630
631.
632
633
634
635
636

O37:

638
640
641
642

644
645
645
646
647

648
649
650
651

652
654
655
656

XXI

15640.

15650.

15660.

105.

106.
107.
108.
109.
110.
11d:
112:
113.
114.
115.
116.
lees

118.
Wo),
120.
121.

122.
123.
124.
125:
126.
127
128.
Wh
13.0:
Tene
132.
133.
134.
135:
136.
137,
138.
139:
140.

SAG ae Re A Bg Spee mee

J:
J.
J.
Jb

WANNA NNN NNN

knorringiana I\jin

Section 13 Molles Korsch.

. arachnoidea Bge.
. talijevii Klok.
. cretacea Bge.

kasakorum (jin
ledebourii Bge.

. mollissima Klok.
. calcarea Klok.

michelsonii jin

. sordida Stev.
. blanda (M.B.) C.A.M.
. exuberans (Trautv.) Sosn.

demetrii lyin

Section 14. Jurinea

alata (Desf.) Cass.
spectabilis Fisch. and Mey.
grossheimii Sosn.
ruprechtii Boiss.

Section 15. Chaetocarpae (Korsh.) Iljin

chaetocarpa Ldb.
persimilis jin

suidunensis (Winkl.) Korsh.
margalensis Ijin

. multiloba yin

dshungarica (Rubtz.) [jin

. longicorollaris (Ijin) Tin

krascheninnikovii Ijin
lipskyi \\jin
adenocarpa Schrenk

. tianschanica Rgl. and Schm.
. foliosa (Tin) Ihjin

almaatensis lin

. lanipes Rupr.

algida lin
darvasica \\jin
densisquamea l\jin
nivea Wink.

. grumosa I\jin

XXXil1

656

659
660
661
662
663
664
665
666
667
668
670
671

673
674
675
676

680
680
681
682
682
683
684
686
687
687
688
689
690
690
691
692
693
694
694

XXII

XXX1V

15670.

15680.

15687.

—

1.

eee

. poacea Ijin
baldshuanica Winkl.
. kirghisorum Janisch.
. Spissa I\jin

. czilikinoana I\jin

. fedtschenkoana Ijin

Species of Unknown Position

. pteroclada I\jin

. bracteata Rgl. and Schm.
. spiridonovii Tin

. mariae Pavl.

. monticola [lyin

SNA NN

Genus 1585. Jurinella Jaub. and Spach

. J. moschus (Habl.) Bobr.
. J. subacaulis (Fisch. and Mey.) Iljin
. J. squarrosa (Fisch. and Mey.) Iljin

Genus 1586. Perplexia Iljin

. P. microcephala (Boiss.) Ijin

Genus 1587. Pilostemon Ijin

. P. karateginii (Lipsky) Tin
. P. filifolia (Winkl.) Tin

Genus 1588. Nikitinia lyin
N. leptoclada (Bornm. and Sint.) [jin

695
696
698
699
700
700

701
701
702
702
703

705
706
707

709

710
711

9/1193

1

Tribe 11. ECHINOPSIDEAE Cass. in Bull. Soc. Philom. (1815)
173.—Trib. Echinopseae Cass. in Dict. sc. nat. X (1818) 156; XIV (1819)
200; XX (1821) 362; Opusc. phyt. I (1826) 299; II (1826) 203.—Tnib.
Echinopodeae Cass. in Dict. sc. nat. XLI (1826) 314; LX (1830) 572.—
Trib. Cynareae subtrib. Echinopsideae Less. in Linnaea, VI (1831) 88
and Synops. Comp. (1832) 13; DC. Prodr. VI (1837) 522; Boiss. FI. or.
Ill (1875) 423.—Trib. Cynaroideae subtrib. Echinopsideae Benth. in
Benth. and Hook. f. Gen. Pl. II (1873) 212.—Trib. Carduees Baill. Hist.
des Plantes, VIII (1882) 69 p. p—Trb. Cynareae subtrib. Echinopsidinae
O. Hoffm. in Pflanzenfam. IV, 5 (1894) 313.—Capitula one-flowered,
aggregated in globose or ovoid heads, with one- or two-rowed involucre.
General involucral bracts very small, usually setaceous, recurved and
hidden under head, sometimes almost undeveloped, or large, leafy, erect,
exceeding and surrounding head. Each capitulum with its own involucre,
consisting of outer row of numerous bristles and many-rowed involucral
bracts, of which outer bracts shorter than inner. All flowers [florets]
tubular, bisexual and fertile, white to blue; their limb pentapartite.
Anthers sagittate at base, with ciliate appendages. Style with short and
thick branches having crown of pollen-collecting hairs below point of
branching. Achenes with appressed hairs. Pappus of short, free or basally
fused bristles, forming crown. Phyllotaxy alternate.

Herbs, distributed in southern Europe, Mediterranean, in the steppes
and semi-deserts of Asia, including Japan, and North Africa.

KEY TO GENERA OF TRIBE ECHINOPSIDEAE

1. Heads consisting of densely aggregated, one-flowered capitula,
globose or ovoid. General involucral bracts very small, usually
setaceous, recurved, or hidden under head, sometimes almost
TAT ENTS! LOY BY= Ts Re ee ee ae ne er 1567. Echinops L.

+ Heads hemispherical, or later becoming almost globose. General
involucral bracts large, outer bracts leafy, exceeding and surrounding
CANON EL Ht Ne ener a i eine 1568. Acantholepis Less.

GENUS 1567. Echinops L.':?

Echinops L. Sp. pl. (1753) 814; Trautv. Diss. Echin. (1833) 16;
Bge. in Bull. Acad. Sc. Pétersb. VI (1863) 390-412; Hoffm. in
Pflanzenfam. IV, 5 (1891) 312; Mulk. in Tr. Bot. Inst. Akad. Nauk
ArmSSR, VIII (1950) 5—92.—Echinopus Adans. Fam. II (1790) 114.—
Echinanthus Neck. Elem. 1 (1790) 91.—Echinopsus St.-Lag. in Ann.
Soc. Linn. Lyon, VII (1880) 109.—Sphaerocephalus O. Ktze. Rev. gen.
I (1891) 366.

Capitula one-flowered, surrounded by numerous bristles; involucral
bracts in three to five rows, membranous or coriaceous; outer bracts
shorter, linear, broadened in upper part; middle bracts lanceolate, carinate,
attenuate into subulate cusp; inner bracts sometimes not longer than
middle ones, free, partly fused in lower part, or fused into tube, and
free above, pointed or fimbriate-ciliate. Corolla regular, white or pale
blue to deep blue, with narrow deeply incised tube protruding from
capitulum; stamens five; anthers fused into tube with tuft of hairs below;
stigma bifurcate, with ring of hairs in upper part of style. Achene
fusiform, slightly ridged, densely covered with long appressed hairs,
often covering pappus; pappus of serrate bristles, more or less fused in
lower part. Numerous capitula terminal on stem and its branches,
aggregated into globose heads with globose or obovoid, glabrous, pitted
receptacle; head surrounded by recurved and compressed common
involucre below receptacle, consisting of scaly and apically broadened
flat bristles like those surrounding individual capitula and forming so-
called brushes. Stems erect, usually sparsely branched in upper part,
sulcate, arachnoid-hairy tomentose, sometimes glandular, very less often
glabrous. Leaves alternate, pinnatifid to bipinnatisect, usually with prickly
toothed lobes, reduced from stem base upward; lower leaves petiolate;
upper sessile; leaves usually pubescent above, sometimes also glandular,
less often almost glabrous; usually almost tomentose beneath. Perennials,
less often surviving only for few years and occasionally annual herbs,
distributed in steppe and semidesert zones, especially in the zone of
mountainous semideserts and steppes.

The genus includes 125-130 species. The distribution area of the
genus is in the eastern hemisphere and extends from the Iberian Peninsula
to Southern Europe and the steppes and semideserts of Asia as far as
Japan; in the south, it occupies the Mediterranean Region and the
northeastern part of Africa.

'Treatment by E.G. Bobrov.
*From the Greek words echinos—hedgehog, and ops(is)—appearance; named on the
basis of spherical and often prickly heads.

Type species: E. sphaerocephalus L.

A.A. Bunge published the most complete review of the genus.
However, published 100 years ago, it, naturally, is antiquated. Out of
the 11 sections included in this review under Echinops, 8 were established
by Bunge. Subsequent authors, Boissier (Flora orientalis) and Hoffmann
(Pflanzenfamilien), slightly enlarged the sections of the genus, a course
that in my opinion is without sufficient basis. Although the number of
Echinops species has almost doubled since Bunge’s publication, we prefer
the classification of the genus based on the subdivisions and the system
proposed by Bunge, even though it is imperfect. Only a new overall
revision of the genus would provide the basis for a total re-examination
of the classification of Bunge and the development of a new
classification. However, such a work is difficult at present because the
collections still have very few plants from Northwest Asia, Iran,
Afghanistan, and Northeastern Africa.

It should also be added that the sections of Echinops are not very
clearly demarcated, which, incidentally, is typical of many genera of
Compositae.

As regards the Echinops species in the Soviet flora, the present
review most probably does not cover them completely. This is because
there are few Echinops specimens in herbaria, because botanists working
in the field often avoid them because of their prickly nature and because
their inflorescences crumble on drying.

Further study undoubtedly will increase somewhat the number of
Soviet Central Asian species.

Economic importance. The globe-thistle species do not have any
significant economic value, but they also are not always referred to
entirely in negative terms, as the troublesome, weedy, ruderal plants.
They do not pose a big threat as weeds, as they are easily destroyed
with the usual control measures, post-harvest soil plowing and inter-row
tillage. The spreading of globe-thistles is greatly facilitated by the limited
grazing of them because of the prickliness.

Some species of globe-thistles have recently attracted attention as
ornamental plants and are fairly widely cultivated for this purpose in
western Europe. The list of ornamental globe-thistles may be significantly
enlarged because of a few attractive Caucasian species, recommended
by Ya.I. Mulkijanyan (Bull. Bot. Sada Akad. Nauk ArmSSR, No. 12,
1951). No less interesting in this respect are a few Soviet Central Asian
species. The ornamental value of many globe-thistle species is somewhat
reduced because they shed the lower leaves early, which is a typical
feature of steppe and semidesert plants.

Some species are valuable as melliferous plants and cultivated for
honey production. Very interesting in this respect are many native

4

species, which deserve to be planted more in open areas, gravelly slopes,
along gorges, etc. Some species contain alkaloids.

Mulkijanyan provided brief information on the biology of the globe-

thistles (Bull. Bot. Sada Akad. Nauk ArmSSR, No. 10, 1951).

+N+ 4

. Plants perennial or surviving few yeafrs ..............:cceesseeeeeeeneeeeeeees 2.
Plants annual (section Nanechinops) ..........cccccccssscceeessceeeeeneeeeees 56.
. Inner involucral bracts connate for almost half their length...... 3.

Inner involucral bracts free, sometimes searcely connate only at

. Involucral bracts 12-15, in three rows; inner involucral bracts

connate, forming pentagonal coriaceous tube (section Oligolepis)

Involucral bracts 16—25, in four or five rows; inner involucral bracts
connate in membranous cylindrical tubes (section Ritrodes) ...

. Outer and middle involucral bracts with straight, usually nee

hairs yalong imargintet.2ks. hago eee BORO ee, Wem 5.
Outer and middle involucral bracts with recurved acicular spinules
in upper part along margin, appearing almost like branched hom
(section: (Pleiaeme) eo uaeRt Sena ee 39. E. tournefortii Ldb.

. Pappus cup-shaped, cleft in upper part (section Therma)..............

AHS 8 SE: 8 AT Ol OED 8 UE. hy 51. E. exaltatus Schrad.
Pappus bristles free almost entirely or over large part............... 6.

. Pappus bristles connate, linear, obtuse, and densely barbulate ... 7.

Pappus bristles subulate, sparsely toothed in upper part, not
barbulate; plants of northeastern Soviet Central Asia and Siberian
mountains from Tien Shan to Sayans (section Chamaechinops) ..
1h cheer) Mika oe pala dedi Gude) ole ipenvesienll ait earel nen Niels atin 53.

. Involucral bracts 16-18; outer bracts usually glandular; leaves

thinner; plants. of forest edges and shrubby thickets (section
EChIinODS)>ehhrech lle RS. dR eal e ae ene ANY 43.
Involucral bracts 18-25; outer bracts without glandular hairs; leaves
coriaceous; plants of plains, mountain steppes, and semideserts
(SectiontRitzo) hanna De. wlel..aue.. te. ce eee 32.

. Outer involucral bracts tomentose .....1. E. transcaspicus Bornm.

All involucral bracts glabrous, smooth, or rough, but not pube-
SCente ZAHER. Ae. SR 2 ee eS ie i AS aT 9.

. Stem entirely or at least in upper half white-tomentose; leaves

oblong-lanceolate; inner-involucral bracts connate in thin, coriaceous
light: browhetubérss..oseea)..reyenl..oet. bade. oes. ceed Bese 10.
Stem glandular, not white-tomentose, sometimes white arachnoid-
hairy above; leaves broadly lanceolate; inner involucral bracts
connate in coriaceous dark brown tube ..................ccccceceeeeeeeeeee LZ.

+

We

5

Stem white-tomentose only in upper half or one-third; upper surface
of leaves green, glandular, glands eventually disappearing but not
-arachnoid-hairy or sparsely white-tomentose; bristles surrounding
mature capitula not shorter but longer than capitula at fruiting...
PN OES AR SE AL Oe Ee 2. E. maracandicus Bge.
Stem entirely white-tomentose and sulcate; leaves with glandular
hairs on upper surface and arachnoid-hairy or white-tomentose
along veins; bristles surrounding capitula not more than half as
AGH EMASM CA HCL aT os FEE TRE EY HED SORE Aer 11

. Leaves grayish-green above and white-tomentose along veins; stem

SEALE SUC ALC sresectee eevee teed eet Renee oe 3. E. polygraphus Tschern.
Leaves green above, arachnoid-hairy (along veins), with scattered
glandular hairs; Stems Slimtthy Sulcate ..0.2:......cctteeceetacesccetseec-ssssceeee
EES mee pe sass 4. E. brevipenicillatus Tschern.

. Stem white arachnoid-hairy above; leaves on upper surface with

numerous, stalked, glandular hairs, somewhat arachnoid-hairy or
white arachnoid-hairy only along veins ................:ccccceesteeeteeees 13:
Stem with short or long-stalked glandular hairs, but not arachnoid-
hairy; leaves rough with numerous short-stalked or sessile glandular
hairs, but without arachnoid-hairy pubescence ..................::00 14.

. Stem deeply sulcate, appearing longitudinally striate; upper surface

of leaves with white arachnoid hairy pubescence along veins; outer
involucral bracts 12-13 mm long.......... 5. E. leucographus Bge.
Stem shallow-sulcate; leaves somewhat white arachnoid-hairy on
upper surface, without distinctive pubescence along veins; outer
involucral bracts about 8 mm long...... 6. E. leiopolyceras Bornm.

. Lower cauline leaves up to 25 cm long and 8 cm wide, crenate-

lobate; bristles surrounding capitula up to 10 cm long.................
Keres. nee 2 a ane caution act crt cum. wouldnt asboor ae 6. E. lipskyi [jin
Lower cauline leaves 10-13 cm long and 4-5 cm wide, widely
shallow crenate-toothed; bristles surrounding capitula 12-14 mm
LONG ceustivca lamisil n jocte: cebiaeren cee 7. E. praetermissus Nevski

. Involucral bracts arachnoid-hairy pubescent in exposed part........

Nit. 2 ee roe eid Amy nto! Me ener sctll a. 15. E. pubisquameus [ljin
Involucral bracts glabrous, smooth, or scabrouS...............cc0008 I.

. Stem 1—1.5—2 m high; lower cauline leaves 25—40 cm long; capitula

8-10 cm wide.............. 26. E. macrophyllus Boiss. and Hausskn.
Stem not more than | m high; capitula 2.5-4 or 5-6 cm wide;
basal and lower cauline leaves up to 25 cm (occasionally 30 cm)
NTN eras oc ae oe RN, ete LO: 82. FEDER BA A eh D7.
Stem white-tomentose or sparsely arachnoid-hairy pubescent without
STU G (UE ET RS anaes Aaa has a cen etn dias ate raal imei sald elbeaeee 18.

20.

Dil

WI

8)

24.

Stem glandular, besides white-arachnoid or sparsely white-tomentose
IN ALOWEL, Dah jOL AONE MAGES. tie. au. te cbmc nto: <cpbaceer eee eee 24.

. Receptacular bristles half or almost half as long as capitulum ....

Receptacular bristles one-third—one-fourth—one-fifth as long as
capitulum .......:... PN: Sah cate Sen memati er RLF 2M
Leaves same colored on both surfaces, gray and finely white-
LOMIENTOSE) ea tecese sci Mi nes crue ha ceanabactee tise, 13. E. dubjanskyi [lyin
Leaves greenish-gray above from sparse arachnoid-hairy pubescence,
almost WIte-FOMERTOSE OCMC ANN atic eae ate ier aae eae ee 20

Lamina pinnatifid in upper part, pinnatisect in lower; capitula 12—
14 mm long, receptacular bristles about 5 mm long, 1.e., less than
half as long as capitulum; corolla tube below throat sparsely.
covered with short bristles (on sands of southern Kazakhstan)...

aR aS RI 7 NAR ae aa 11. E. albicaulis Kar. and Kir.
Lamina pinnatisect, with 34 paired and one terminal lobe; capitula
14-15 mm long, receptacular bristles 8-9 mm long, i.e. exceeding
half their length; corolla tube below throat with few glandular
hairs. (Semidesert, ‘southern: kazakhstan): .A4. 484 oe ae

NN SR Neh TRS IR Ete ona 12. E. kafirniganus Bobr.
Leaves grayish-green above, sparsely arachnoid-hairy; lower cauline
leaves up fo 20°25) Cin TOMS, va... sssctececseuece ete Mee yee ee DD
Leaves green above, rough, without arachnoid-hairy pubescence;
lower Cauline: leaves, 30-40) cm ONS css cence esteeenas tee ee ee

Menace te eda NVA ne ela nett 18. E. chantavicus Trautv.
Basal leaves pinnately divided with prickly, crenate, narrow (0.5
cm) rachis and narrow, deltoid, spinescent lobes; corolla tube
slandular and. Spreatlime. Watiy aDOVE c2r.c.4. cents: cc-eeces coe aeces eens 23%
Basal leaves pinnately divided into narrow-lanceolate lobes, densely
prickly with fine and short spinules along margin; corolla
tube Eucent on outer side of throat (Mountainous Turkmenia)

A coeibe RSA anA toy RRL RONaL eas an UNOS AEE 9 LA LRCIEN wads 9. E. ritrodes Bge.

Leaves sparsely arachnoid-hairy above, without glandular hairs;
corolla tube densely covered with long and spreading hairs in upper
part (northern ranges of Pamiro-Alai, Shakhimardan) ...................

snshwtbsncieensieaiaue MRR: Bes ee 16. E. dasyanthus Rgl. and Schm.
Leaves sparsely arachnoid-hairy above, together with fine, sessile,
glandular hairs, short-stalked on petioles; corolla tube very sparseiy
pubescent, (southernyranges off fiem, Sham) ie. cseecnae-teuteoeeee ode

Pe SR a eae tee RRA NE 17. E. tjanschanicus Bobr.
Leaves green on both sides, rough above, densely glandular beneath,
along petiole and veins (southern Balkhash region, southeastern
Atalo-Caspiane TePiOM)s .4..6.65.) ee ee 14. E. subglaber Schrenk

27

28.

29;

30.

ons

Sp!

Ba:

7

Leaves grayish-green above, usually with arachnoid-hairy pubescence
or glandular, white- or grayish-tomentose beneath.................... 25.

. Leaves grayish-green above, with arachnoid-hairy pubescence but

not glandular, grayish-tomentose beneath, without glandular hairs;
stem in lower part white arachnoid-hairy along ridges, yellowish-
brown along grooves, with stalked glandular hairs.............0....00..
ee Oe a oh ae eeerten Penner et cee 22. E. fedtschenkoi [jin
Leaves always with stalked or sessile glandular hairs above,

‘sometimes scattered; lower surface of leaves usually without

glandular hairs or glands present only along veins .................. 26.
. Stem 40-80 cm high; capitula 3.5-6 cm wide......... eee Zi
Stem 20-35 cm high; capitula 3-4 cm wide............ eee Sule
ants solwsoviet ential A Stat. ..2 cen ckecocsnncenetauerenae tae ees 28.
Platts, of, sliransCaucaSlanscced otcboy. - i g-surged <i} oaoauphenied onal: oO:

Stem with arachnoid-hairy pubescence and glandular in lower part;
leaves with scattered stalked glandular hairs above; capitula 2—2.5
COON Gia cris eri Mia dees Bscaod 19. E. karatavicus Rgl. and Schm.
Stem floccose and loosely tomentose, with short-stalked glandular
hairs; leaves with scattered sessile glandular hairs above; capitula

IVa Reva 100 C0) one, aan ee 20. E. obliquilobus [jin
Stem 2—5 mm thick, short-glandular in lower part; leaves finely
and very loosely tomentose above............. 24. E. opacifolius [ljin

Stem more than 6 mm thick, entirely glandular; leaves without
pubescence above, but glandular or somewhat arachnoid-hairy and
rove) due, tomstalked elandulanthaitsj2's..aec beth. Bee eee.. 30.
Leaves hard and rough above due to dense cover of stalked
glandular hairs; glandular hairs on stem numerous, surrounding
flat and curly brownish bristles .............. 23. E. orientalis Trautv.
Leaves not pubescent above, but with thick-stalked glandular hairs;
stem with numerous similar glandular hairs................:ccsccceeeeeeeee
2S eepemenei site tye. IU, LONE, PER 25. E. connatus C. Koch
Stem with finely tomentose pubescence and glandular hairs; leaves
with sessile glandular hairs above; capitula 18—20 mm long.......
ws LENE. MOEA. Bakes 8 SONS. SO, 21. E. babatagensis Tschern.
Stem glandular and white arachnoid-hairy only above; leaves with
short-stalked glandular hairs above; capitula 13-15 mm long......
Pea hry, PRU YER ret kacidl heehee yn 10. E. multicaulis Nevski
Stem with white arachnoid-hairy or white-tomentose pubescence,
watitout, obardular hairs. fects cee seine eae an ea ee 34.

‘Stem glandular entirely or only in lower and middle parts, usually

Witte tOMEMLOSS -AD OVE) tu. au.---easee-sceeberbere ataastasals SHR eee Sa 33).
Stem glandular hairy only in lower or lower and middle parts,
Wihite-tomentase, Only MN WIPPer PAalts:0...cc..c...c-ccaceertensapement®-. Be -- BO)

34.

35

36.

oy

38.

39%

40.

41.

Stem entirely glandular hairy and without tomentose pubescence
STLIET ots A nener: emer eraeeen eee ante Ne ee ee 28. E. meyeri (DC.) Iljin
Lobes of leaves lanceolate, more or less flat, in same plane as

E091 50 lea MnO M eR eo batt bbe ol hdd iet Mei ac Me fanaa ite core meeD 3)3)5
Lobes of leaves spinulate, divergent in different planes. Capitula
3-3.5 cm wide (Greater Balkhans)................ 29. E. spiniger Iljin

Leaves green arachnoid-hairy above, grayish-tomentose beneath,
with prominent veins; both upper and lower surfaces eglandular

Leaves grayish-green, weakly arachnoid-hairy above, with few
short-stalked glandular hairs; grayish-tomentose beneath, with
prominent veins and scattered glandular hairs ...................cccceee
RE Den Ri cone treet priory tener en pea Ansar 31. E. polyacanthus Iljin
Outer involucral bracts light brown in broader part scarcely
brownish, less than half as long as involucre ...............cc:ccceee Sule
Outer involucral bracts dark brown in broader part, more than half
asslongaas TMVOlUcTe me rete cece 34. E. transcaucasicus [jin
Stem up to 60 cm high; lower leaves up to 45 cm long, midrib of
lower leaves up to 5 mm wide beneath, yellow (Karabakh) ........
mites acres learn rrer meter cee ene snercrton aan 38. E. karabachensis Mulk.
Stem 1 m or more high; lower leaves 50-75 cm long and 25-30
cm wide (Lake Sevan basin, Vedinskii Region of Armenia) .. 38.
Capitula narrow at base, somewhat conical; bristles surrounding
capitula as long as outer involucral bracts; basal leaves up to 75
CMolONnGecrcsassenesrsaana ee 36. E. iljinii Mulk.
Capitula broad conical; bristles surrounding capitula shorter than
outer involucral bracts; basal leaves up to 50 cm long ................
BL RAE IO Se RO eae eh Soe OE 37. E. sevanensis Mulk.
Leaves dark green above, glabrous or weakly arachnoid-hairy
pubescent; white-tomentose beneath; both surfaces without
Slanidilar ais?) 4k: a a ee 27. E. ritro L.
Leaves arachnoid-hairy pubescent and always glandular above;
white-tomentose beneath, occasionally weakly pubescent and
without *elandular “hairseie8 Bn OR OE ee 40.
Leaves arachnoid-hairy and glandular above, slightly incurved;
white-tomentose beneath, without glandular hairs .................06:
= RS SEP RORE A STE ea oe Ee See AH 30. E. pungens Trautv.
Leaves with stalked glandular hairs beneath all over lamina or
only “along--veinse: fy. Bi nk Se EE A Heine toe 41.
Leaves arachnoid-hairy above, rugose and scabrous due to numerous
short-stalked glandular hairs; grayish-tomentose beneath, with
numerous light brown, stalked-glandular hairs along veins. Capitula
57 emi wide: sheet e Va 35. E. szovitsii Fisch. and Mey.

42.

43.

44.

45.

46.

47.

48.

49.

5)

Leaves arachnoid-hairy and glandular above but not scabrous;
white-tomentose beneath, with numerous long-stalked, brown,
glandular hairs along veins, or weakly glandular-hairy to white-
tomentose, with few stalked glandular hairs ........00. eee 42.
Leaves arachnoid-hairy above, with scattered short-stalked glandular
hairs; white-tomentose beneath, with few long-stalked glandular
hairs along veins. Capitula 4-5 cm wide, on lateral branches 3—
SPOVCMMS «. nteence sate sce aetna. -eeteasens- 33. E. foliosus Somm. and Lev.
Leaves arachnoid-hairy and glandular above; weakly glandular-
pubescent to gray-tomentose beneath with thick glabrous veins,
bearing, few stalked-glandular hairs. Capitula 6—6.5 cm wide.....
Auriinn CMe ASE PS CAENG 2hr 8 setae REET SERA ee ea 32. E. conrathi Freyn
Stem arachnoid-hairy or loosely tomentose, always with stalked-
glandular hairs over entire length, in middle, or only in lower part
PRR RRA N A REREEE AAP: MENS cust chehanaaneretiabeuetet vsthasuscinban Wan cakinmchee tht piias 44.
Stem floccose-white tomentose, glabrescent in parts, without glandular
hairs; leaves without glandular hairs above as well as beneath

IGM OM VSTIIS J. c0F sever cnet a omeneniarnmitenr ete’ 50. E. abstersibilis [ljin
Leaves very sparsely arachnoid-hairy beneath, slightly lighter in
color than grayish-green upper surface ............c:ceeeeeeseteeereeeees 45.
Leaves grayish arachnoid-hairy or white-tomentose beneath,
differenteinicolorntiromupper Surfaces: 2 ae: 1. 46.
Involucral bracts 15—18; outer bracts glandular dorsally (Yerevan
Resion)iunadanm..2nsd xeloboele, bodks 41. E. erevanensis Mulk.

Involucral bracts 20—23; outer bracts without glandular hairs
dorsally and only very short hairy, other bracts scabrulous. ........
op de Cleat, shite. itucl | io: Maren hie, 43. E. viridifolius []jin
Leaves almost glabrous or scarcely arachnoid-hairy above, without
glandular hairs, sometimes scabrous due to numerous bristles.....

ERIE ss el Reh. eegila PR. delay. Hct A ee. 47.
Leaves scabrous above from stalked glandular hairs or bristles
sumounded pby,clandularghairs .)8i)..f4A shoes en Le Sill.

Leaves scarcely arachnoid-hairy and scabrous above from numerous
bristles; capitula 15—18 mm long, receptacular bristles little less
than half as long as involucre .................. 46. E. ossicus C. Koch
Leaves almost glabrous above or scarcely arachnoid-hairy, not
Seabroustudank “onecmodite.... setagath fds elisen se. tmleme . enkanel, ee, 48.
Plants of eastern Kazakhstan or western Transcaucasia ........... 49.
Plants oipEastermenSiberiagandibar J2aSter.. c.aseecte.c noes, ease -cs 50.
Stem 1—1.5 m high; heads 4-6 cm wide; capitula 20-23 mm long;
leaves lanceolate, finely tomentose and glandular beneath (eastern
Kazakhtain) or a. sessed depeteawrenee. Sete cee 49. E. tricholepis Schrenk
Stem 0.6—1 m high; heads 3—4.5 cm wide; capitula 15-20 mm
long; leaves almost lyrate, loosely white-tomentose and without

10

50.

Sali:

§2:

56.

aye

glandular hairs beneath (western Transcaucasia).............::000ce
Ee Ee Pe cena ROY Nie MENSee seeeeeee 45. E. colchicus Sosn.
Stem weakly arachnoid-hairy below, upper surface of leaves weakly
arachnoid-hairy (Eastern Siberia)............ 47. E. latifolius Tausch.
Stem almost glabrous below, with isolated brown hairs; leaves
glabrous above, more deeply incised (Far East) .......... cc ceeeee eee
Piecing ai at te tiara cece he erm tern 48. E. dissectus Kitag.
Lower leaves lyrate, with small, ovate, remote lower lobes and
large elliptical terminal lobe, short prickly-toothed along margin
o deaponsanyadDaeT das QM i ee ri Nai a sara Behe 44. E. galaticus Freyn.
Lower leaves lanceolate, with ovate to deltoid lobes, with short
acute teeth on margin and terminating in short spinules ......... Sw.
Stems ribbed, glandular along sides and ribs, appressed-hairy above,
appearing white arachnoid-hairy; leaves few, distant ....................
» Riwsictiawude le lisaat eae Be Ok Aaa unig 40. E. sphaerocephalus L.
Stem sulcate, with numerous brownish-glandular hairs along sides
white arachnoid-hairy along ridges, leafy up to heads (eastern
CaUcasusy rc. RAs deed hess Serva aaa: 42. E. dagestanicus I]jin

. Leaves undivided, with one or several teeth only at base............

Ber cence ost Mer. 52. E. integrifolius Kar. and Kir.
Leaves ipinnatilobate ,orapinnatisectecds 20s.. Abeer one) 54.

. Stem white-tomentose, without glandular hairs; leaves pinnatilobate;

lower leaves with obtuse lobes ................0006 53. E. humilis MB.
Stem covered with stalked glandular hairs, arachnoid-hairy only in
uppemipant..waten etree, ele tS Ol ato e mie ieay S)5)5

. Stem 10-30 cm high; lobes of leaves and leaf axis very narrow or

linear; corolla glandular-hairy on outer side below throat; plants
of they Tien! Shan aa. “90 54. E. tschimganicus B. Fedtsch.
Stem 25-45 cm high; lobes of leaves and leaf axis wider; corolla
glabrous below throat; plants of southern Altai and Zaisan Basin
Oo TUR SEO DOANE. ft 55. E. saissanicus (Keller) Bobr.
All leaves arachnoid-hairy-woolly or tomentose on both sides; stem
White-tomeritose Oss ok INS Aer Oe hw 57. E. nanus Bge.
All leaves, or at least upper, green above due to arachnoid-hairy
pubescence or absence of tomentum, with stalked glandular hairs;
stent” elandullat...5 earee es eet, SA ee DRE ee ee ie
Lower leaves scarcely tomentose due to long entangled hairs,
undivided or more or less lobed (plants of Central Asia; in Soviet
Union found only in southern Altat)........... 56. E. gmelini Turcz.
All leaves without tomentum or arachnoid-hairy pubescence above,
green or yellowish-green, with numerous stalked glandular hairs,
undivided or crenate-spiny toothed (Fergana Valley) .................---
VAR a sts. RAE Nada ded gels aicvias {anGacae coat teem ee 58. E. knorringianus [lyin

11

Section 1. Oligolepis Bge. in Bull. Acad. Sc. Pétersb. VI (1863)
400, 403; Boiss. FI. or. III, 424.—Involucral bracts 12—15, in three rows;
inner bracts connate, forming pentagonal coriaceous tube.

Lectotype of section: E. leucographus Bge.

1. E. transcaspicus Bornm. in Beih. bot. Centralbl. XXXVI, 2 (1918)
226.

Perennial. Stem glabrous below and in middle, glandular above.
Leaves with sessile glandular hairs above. Outer involucral bracts rather
hard, tomentose; inner bracts sometimes attenuate into long spinule in
sterile capitula.

Soviet Central Asia—mountainous Turkmenia (Middle Kopetdag
Sulyuklyu-Saratovka near Iran border). Described from indicated place.
Type in Weimar.

Note. An enigmatic plant, incompletely described on the basis of
immature specimens collected by T. Sintenis (No. 688, 10.VII.1902) in
Saratovka. It is necessary to search in this region for a globe-thistle
with leaves glandular above and tomentose involucral bracts.

The original diagnosis is insufficient to confirm the placement of
this species in the section Oligolepis.

Series 1. Maracandici Bobr.—Stem white-tomentose entirely or at
least in upper half.

2. E. maracandicus Bge. Beitr. z. Kenntn. d. Fl. Russl. u. Stepp.
Centr. As. (1851) 175; Mém. Sav. Etr. VII (1854) 351; Bge. in Bull.
Acad. Sc. Pétersb. 6, 403; O. and B. Fedtsch. Perech. Rast. Turk. IV,
222; Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada, III, 99 —E. xanthocanthus
Rgl. and Schm. in Izv. Obshch. Lyubit. Estestv. Antrop. Entnogr. 34, 2
(1882) 46.—O. and B. Fedtsch. op. cit—E. argyrocomus Bornm. in
Beih. bot. Centralbl. XXXVI, 2 (1918) 222.

Perennial. Root many-headed. Stem 40-100 cm high, branched
above; glabrous in lower half, smooth and yellowish, sometimes with
fewer glandular hairs; densely white-tomentose in upper half or one-
third, leafy throughout. Leaves green above, with sessile or almost sessile
glandular hairs, glabrescent, densely white-tomentose'and eglandular
beneath; all leaves lanceolate, pinnatifid; lower leaves very short-
petiolate; upper leaves pinnatilobate with remote deltoid or ovate toothed
lobes terminating in long, yellow, bare spines; sinuses wide, roundish,
usually without spines and entire. Heads 5-6 cm wide. Capitula about 3
cm long; bristles surrounding capitulum numerous, silvery, longer than
capitulum at fruiting, initially shorter. Involucral bracts 15-17, glabrous;
inner bracts connate in larger part into membranous cylindrical tube,
brownish in fused part, greenish in free part, broken-toothed along

12

margin; outer bracts shorter than bristles, rhomboid above, ciliate; middle
bracts cuneate, broadened above and terminating into spine; corolla pale
yellow; apical appendages of anthers longer and compressed. Achenes
with numerous yellowish bristles; pappus of unequal, yellowish bristles,
connate up to half length. Flowering VIJ-—VIII.

Shallow and gravelly slopes of middle and lower mountain zone.—
Soviet Central Asia: Tien Shan, Syr-Darya (Fergana), Pamiro-Alai (from
Zeravshan to western Pamir). Endemic. Described from upper Zeravshan.
Type in Paris.

3. E. polygraphus Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVI (1961) 52.

Perennial. Stem 40-80 cm high, striate-sulcate, white-tomentose,
with few short branches above, leafy. Leaves coriaceous, grayish green
and loosely white-tomentose along veins on upper surface, with very
prominent veins and grayish-tomentose beneath, lanceolate; basal leaves
petiolate; cauline leaves short-petiolate, pinnatipartite, with narrow and
entire axial part and narrow-deltoid entire lobes, terminating in spine,
with two short basal spinules; upper leaves narrow-lanceolate, large spiny-
toothed. Heads 44.5 cm wide. Capitula 28-30 mm long, surrounded by
numerous, somewhat flat, dirty-white, 15—16 mm long bristles. Involucral
bracts 13—14, glabrous, thin; outer bracts linear, 11-12 mm long, deltoidly
broadened above, short-toothed, acuminate, scarious; middle bracts
lanceolate, 17-20 mm long, carinate, with wide membranous margin,
acute; inner bracts connate over three-fourths in membranous tube, free
above, creamish, short-ciliate, and pointed. Corolla white, deeply parted,
with glandular tube. Achenes densely covered with yellowish hairs;
pappus of scabrous bristles fused at base. Flowering VI—VII; fruiting
VII-VUI.

Red sandstone foothills ——Soviet Central Asia: Pamiro-Alai (extreme
west, Yakkabag Region). Described from the vicinity of Kipchak winter
camp. Endemic. Type in Tashkent.

4. E. brevipenicillatus Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVI (1961) 51.

Perennial. Stem erect, 30-60 cm high, weakly sulcate, white-
tomentose, leafy, branched above. Leaves coriaceous, green above with
scattered glandular hairs and arachnoid-hairy, white-tomentose beneath
with prominent veins; basal leaves usually short-petiolate; cauline leaves
sessile, auriculate, and spinulate at base, lanceolate, acuminate, pinnatifid
or partite, with deltoid entire lobes, terminating into spinule; upper leaves
lanceolate, auriculate, crenate-toothed, with prickly pointed teeth. Heads
about 5 cm wide; capitula 2.5 cm long, bristles surrounding capitula

14

half as long as capitulum at end of flowering, somewhat flat and white.
Involucral bracts 16—19, glabrous; outer bracts free, inner bracts connate
up to half their length in membranous tube; outer bracts about 8 mm
long, linear, broadened above and short-acuminate, narrow-membranous;
middle bracts lanceolate, carinate, broadened above and attenuate in
cusp; inner bracts lanceolate in upper free part, acute, variable in length,
greenish. Corolla whitish, deeply parted with glandular tube. Achenes
densely covered with yellowish hairs; pappus of scabrous yellowish
bristles, fused at base. Flowering VI; fruiting VII.

Red sandstone foothills.—Soviet Central Asia: Pamiro-Alai
(southwest Babatag). Endemic. Described from vicinity of Dzhida-Bulak
in Babatag. Type in Tashkent.

Series 2. Leucographi Bobr.—Stem glandular, not white-tomentose,
sometimes white arachnoid-hairy above.

5. E. leucographus Bge. in Bull. Acad. Sc. Pétersb. VI (1863) 406;
O. and B. Fedtsch. Perech. Rast. Turkm. IV, 223; Fl. Turkm. VU, 145.—
E. grossheimii \\jin Fl. Kavk. IV (1934) 160; Mulk. in Tr. Bot. Inst.
Akad. Nauk ArmSSR, VIII, 79.

Perennial. Stem 40-60 cm high, branched above, deeply sulcate
and hence appearing striated, increasingly densely white arachnoid-hairy
above, with scattered glandular hairs, encircling curly brown hairs, weakly
arachnoid-hairy below, with numerous curly, stalked-glandular hairs.
Leaves coriaceous, green above with numerous stalked-glandular hairs
and white arachnoid-hairy veins, lanceolate; basal leaves 20-30 cm long,
oblong, narrowed toward base; cauline leaves wider, with slightly
decurrent base; all leaves boldly sinuate-toothed, with broad triangular
teeth, prickly-toothed along margin and terminating in spinule, sinuses
with shorter spinules, teeth reduced toward base of leaves, leaves thus
narrowed appearing petiolate; upper leaves small, with broad base, prickly
toothed. Heads 3—4.5 cm wide; capitula 2—2.5 cm long, bristles
surrounding them about 15 mm long. Involucral bracts 15-16; outer
bracts 12-13 mm long, broadened above and terminating in cusp,
carinate, scarious along margin, connate at base; middle bracts 18—20
mm long, like outer, but with longer cusp, also connate at base; inner
bracts about 20 mm long, fused over two-thirds length into brown
pentagonal tube, free above, cristate and cuspidate. Corolla pale blue,
deeply parted, with a glandular tube. Achene densely covered with
yellowish hairs; pappus of scabrous bristles fused at base. Flowering
VI-VII; fruiting VII.

Lower mountain zone.—Caucasus: Southern Transcaucasia (Megry),
eastern Transcaucasia (Ordubad); Soviet Central Asia: mountainous

15

Turkmenia, Pamiro-Alai. General distribution: Iran. Described from
northern Iran. Type in Leningrad.

6. E. leiopolyceras Bornm. in Beih. bot. Centralbl. XXXVI, 2 (1918)
208; Fl. Turkm. VII, 146.—E. griffithianus auct. Fl. As. Med. non
Boiss.—E. armenus Grossh. in Tr. Tifl. Bot. Sada, II, 1 (1920) 36; Iljin
in Grossh. Fl. Kavk. IV, 160; Mulk. in Tr. Bot. Inst. Akad. Nauk
ArmSSR, VIII, 70.—E. araxinus Mulk. op..cit. (1950) 78; Dokl. Akad.
Nauk ArmSSR, XII, 1 (1950) 29.

Perennial or biennial. Stem 40—70 cm high, one or two-head, sulcate,
branched and more densely arachnoid-hairy above, densely glandular
along entire length, glandular hairs on curly, divergent, brownish bristles.
Leaves coriaceous, grayish-green, very weakly arachnoid-hairy, and
densely glandular above, white-tomentose beneath with prominent veins,
covered with stalk glandular hairs, lanceolate; basal leaves 20-25 cm
long; cauline leaves broadly amplexicaul; all leaves crenate-toothed, teeth
long-spinose along margin and leaves terminating in very long spines;
upper leaves small, sessile, auriculate, with spines longer than lamina.
Heads 5—6 cm wide, greenish. Mature capitula 2.5-3.5 cm long,
surrounded by bristles, up to 16-18 mm long’ at maturity. Involucral
bracts 15, finely scabrous; outer bracts about 8 mm long, carinate,
membranous along margin, broadened above and more prickly acuminate;
middle bracts narrowed above into long, stiff, grooved, 1.5-3 cm long
cusp; five inner bracts connate over two-thirds their length into dark
pentangular tube, free part attenuate into grooved acute tip, 3-4 cm
long, corniculate in sterile capitula. Corolla whitish-blue, deeply parted,
with glandular tube. Achenes yellow-hairy; pappus of sulcate bristles
fused at base. Flowering VI—VII; fruiting VII.

Gravelly slopes of lower hills, red sandstones—Caucasus: Southern
and eastern Transcaucasia; Soviet Central Asia: mountainous Turkmenia,
Pamiro-Alai (in the west—Surkhan Darya and Kashka Darya provinces).
General distribution: \ran. Described from northern Iran. Type in Weimar.

Note. The formal difference between this species and E. griffithianus
Boiss., described from the Kabul region, is that the leaves are glandular
above, whereas the leaves of the latter species, according to Bornmiiller,
are not glandular above. If this were to prove wrong and in authentic
specimens of E. griffithianus Boiss. the leaves were found to be glandular
above, then it would follow that the name proposed by Boissier should
be considered to have priority

7. E. praetermissus Nevski in Tr. Bot. Inst. Akad. Nauk SSSR,
Ser. 1, 4 (1937) 206, Fig. 9; Fl. Turkm. VII, 145.

16

Perennial. Stem 50-70 cm high, simple, covered almost entirely
and especially densely in upper part with rusty long-stalked glandular
hairs. Leaves coriaceous, green above, densely glandular and scabrous
due to numerous short-stalked glandular hairs; white-tomentose beneath
with rather prominent midrib; lower cauline leaves oblong, 10-13 cm
long, 4-5 cm wide, terminating in 1—1.5 cm long spine, broadly and
shallow crenate-toothed, prickly along teeth, with two spinules near base.
Leaves reduced toward upper part, ovate, prickly-toothed, slightly curved
in lower part. Heads about 4.5 cm wide. Capitula 2—5 cm long; bristles
surrounding capitula 12-14 mm long, white, rusty below, half as long
as capitulum. Involucral bracts 14-16; outer bracts 10 mm long,.
lanceolate, spa‘ulate in upper part and abruptly broad truncate with short
acute cusp; middle bracts 11-17 mm long, cuneate, abruptly pointed
and prickly in upper part; inner bracts 20 mm long, .connate over two-
thirds length in brown coriaceous tube, free above, toothed and attenuate
into subulate cusp, unequal. Corolla light bluish, with glandular tube;
pappus of bristles fused at base. Flowering VI; fruiting VII.

Red sandstone lower hills.—Soviet Central Asia: Pamiro-Alai
(Kugitang Range near the village of the same name). Endemic. Described
from the place indicated. Type in Leningrad.

8. E. lipskyi Iljin in Bot. Mat. Gerb. Bot. Sada, IV (1923) 97.

Perennial. Stem 40-70 cm high, with remains of leaves at base,
branched, densely glandular with short-stalked hairs. Leaves coriaceous,
green on upper surface, scabrous due to short-stalked or almost sessile
glandular hairs; white-tomentose with prominent glandular hairs beneath;
lower cauline leaves up to 25 cm long and 8 cm wide, narrowed toward
base, with obtuse apex, attenuate into stiff spine, crenate-lobed, with
broad deltoid entire lobes, with one terminal and two basal spinules;
leaves reduced toward stem apex, ovate, prickly-toothed along margin.
Heads solitary; fully developed capitula about 2.5 cm long; terminal on
branches; bristles surrounding capitula unequal, up to 10 mm long, rusty-
white, slightly flat. Involucral bracts 10—20; outer bracts lanceolate,
membranous along margin, broadened above and short-acuminate at tip;
middle bracts 15-17 mm long, carinate, spatulate above and attenuate
into long spine, connate at base; inner bracts up to 25 mm long, connate
over larger part in coriaceous, pentagonal, dark tube, terminating in
spine. Corolla light bluish, with glandular deeply parted tube. Achene
densely covered with short, yellowish hairs; pappus of bristles fused at
base, free above. Flowering VI-—VII; fruiting VII. (Plate I, Fig. 3).

Gravelly slopes of lower mountain zone.—Soviet Central Asia:
Pamiro-Alai (Gissar Range, Denau, Dagana, Boisun, Derbent). Endemic.
Described from Dagana. Type in Leningrad.

)

Section 2. Ritrodes Bge. in Bull. Acad. Sc. Pétersb. VI (1863) 401,
406; Boiss. Fl. or. III, 424; Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR,
VIII, 169.—Involucral bracts 16—25, in four or five rows; inner involucral
bracts connate in cylindrical membranous tube.

Type of section: E. ritrodes Bge.

Series 1. Ritrodes.—Basal leaves pinnatifid into narrow-lanceolate
lobes, densely covered with thin and short spines along margin.

9. E. ritrodes Bge. in Bull. Acad. Sc. Pétersb. VI (1863) 407; Boiss.
Fl. or. III, 428.

Perennial. Root two or three-headed, forming caudex with floccose-
tomentose stem bases surrounded by remains of old leaves. Stem 30—60
cm high, branched above; branches in groups of two or three, entirely
white tomentose-arachnoid-hairy, leafy in basal and lower parts. Leaves
lanceolate; basal leaves 20-25 cm long, 4-5 cm wide below midpoint
in widest part, pinnatifid into lanceolate lobes, with thin: and short spines,
increasingly reduced upward along stem; upper leaves 1-3 cm long,
narrow-lanceolate, prickly-toothed; upper surface of leaves grayish-green,
finely sparsely arachnoid, glabrescent; lower surface grayish, densely
gray-arachnoid-hairy, like petioles and prominent veins. Heads solitary
terminal on branches, one to three on single stem, 3—3.5 cm wide.
Capitula 13-15 mm long; bristles surrounding capitula unequal, somewhat
flat, yellowish-white, up to 4 mm long. Involucral bracts 20-23; outer
bracts rhomboidal at apex, acuminate, short-ciliate; outermost bracts
scarcely longer than bristles; middle bracts up to 10 mm long, smooth
or very finely scabrous, carinate, acuminate in spine, ciliate along margin;
inner bracts even more attenuate, up to 12 mm long, connate at base.
Corolla bluish (or pinkish?) deeply parted; tube pubescent on outer side
near throat. Pappus of scabrous bristles fused at base. Flowering VII;
fruiting VIII.

Gravelly slopes of lower mountain zone in deserts.—Soviet Central
Asia: mountainous Turkmenia (Firyuza, Desht, Aidere). General
distribution: Iran. Described from the vicinity of Mashhad. Type in
Leningrad.

10. E. multicaulis Nevski in Tr. Bot. Inst. Akad. Nauk SSSR, Ser.
1, 4 (1937) 286; Fl. Turkm. VII, 144.

Perennial. Plants with three or four-headed root, producing three or
four stems, surrounded by numerous remains of old leaves. Stems 20—
35 cm high, densely covered with short-stalked glandular hairs, moreover
white arachnoid-hairy in upper part, with few branches above. Leaves

18

coriaceous, oblong-lanceolate, grayish-green above, with short-stalked
glandular hairs, scabrous; leaves white-woolly beneath, glandular and
greenish along prominent veins; leaves pinnatifid, with linear-lanceolate
lobes; basal leaves 15—25 cm long and 3—4 cm wide with lobes; cauline
leaves shorter and narrower, leaf axis between lobes 0.5—0.8 cm wide,
veins terminating in spinules, hence even linear lobes appear spiny;
upper cauline leaves narrow-lanceolate, simple prickly-toothed and not
pinnatipartite. Heads 3—4 cm wide. Capitula 1.3—1.5 cm long, surrounded
by numerous somewhat flat and white 3-4 mm long bristles, one-third
to one-fourth as long as capitulum. Involucral bracts glabrous, scabrous,
bluish; outer bracts 3-5 mm long, lanceolate, broadened above, long-
ciliate; middle bracts carinate, grooved on inner side, long-ciliate and
terminating into spine, slightly recurved above; outer and middle bracts
free at base; inner bracts connate at base, brownish below on inner side,
bluish above on outer side, long-ciliate and attenuate into cusp, longest
reaching 15 mm in length. Corolla blue(?), glandular on outer side.
Achenes densely hairy; pappus of bristles, light colored. Flowering VII;
fruiting VIII.

Middle mountain zone.—Soviet Central Asia: Pamiro-Alai (Kugitang
Range). Described from the rise to Kugitang along the western slope, in
the middle of the range. Type in Leningrad.

Series 2. Albicaules Bobr.—Stem white-tomentose or more sparsely
arachnoid-hairy, without glandular hairs. Plants of foothill semideserts
of northern Tien Shan, southern Pamiro-Alai, and Ust-Urt. E. albicaulis
Kar. and Kir., E. dubjanskyi I\jin and E. kafirniganus Bobr. belong to
this series.

I tentatively include here also the morphologically distinct £.
subglaber Schrenk and E. pubisquameus I\jin, which differ from the
above-mentioned species and apparently themselves represent isolated
members of other evolutionary series.

11. E. albicaulis Kar. and Kir. in Bull. Soc. Nat. Mosc. XV (1842)
387; Ldb. Fl. Ross. II, 654; O. and B. Fedtsch. Perech. Rast. Turkm.
IV, 223.—E. jaxarticus Bge. in Beitr. z. Kenntn. d. Fl. Russl. u. Stepp.
Centr. As. (1851) 175; Mém. Sav. Etr. VII, 350; Boiss. Fl. or. III, 428.—
E. kasakorum Pavl. in Vestn. Akad. Nauk KazSSR, 8 (1950) 21, Fig.
16.

Perennial. Stem 40—80 cm high, one or two-branched, sometimes
with short branches and densely white arachnoid-hairy, angular under
pubescence, without glandular hairs. Leaves greenish-gray above due to
sparse arachnoid-hairy pubescence, almost white beneath due to dense
arachnoid-hairy, almost tomentose pubescence; basal leaves petiolate,

19

lanceolate, 20-30 cm long, pinnatifid in upper part, pinnatisect below,
hence petiole as if bearing small prickly lobes; lobes in upper part of
leaf distant, larger, spinose-spatulate, usually over 3-4 cm appearing
decurrent along axes; cauline leaves sessile, smaller. Heads 3-4 cm wide,
terminal on main stem and branches. Capitula 12-14 mm long; bristles
surrounding capitula white, unequal, less than half as long as capitulum.
Involucral bracts glabrous; outer bracts scarcely longer than bristles,
cup-shaped, rhomboidal above, ciliate; middle bracts lanceolate, carinate,
spinescent, scarcely colored above, ciliate along margin; inner bracts
slightly shorter than middle, brownish on inner side, connate. Corolla
whitish, deeply parted, tube sparsely covered with short bristles above.
Achenes covered with yellowish-brown hairs; pappus of short scabrous
bristles fused below. Flowering VI; fruiting VII.

Sandy semidesert.—Soviet Central Asia: Aralo-Caspian Region
(southeastern), Balkhash Region, Dzhungaria-Tarbagatai, Syr-Darya.
Endemic. Described from the Lepsa River. Type in Leningrad.

12. E. kafirniganus Bobr. sp. n. in Addenda, XXVI, 714.

Perennial. Root virgate, three to five-headed. Stems two or three,
60-80 cm high, branched above near tip, indistinctly sulcate, white
arachnoid-hairy throughout, without grandular hairs. Lower cauline leaves
15-20 cm long, 6-8 cm wide, with three or four pairs of subopposite
and one terminal lobe; upper lobes 3—4 cm long and 2-3 cm wide,
prickly-toothed, 2—3 cm apart, short-decurrent, lower lobes spinose, 1—
2 cm long, wider apart; basal leaves semiamplexicaul, spinose; upper
leaves lanceolate, deeply pinnatifid into prickly-toothed lobes, light
grayish-green above, without glandular hairs, arachnoid-hairy, almost
white beneath due to dense arachnoid-hairy pubescence, eglandular.
Heads two or three on stem, 3.5-4 cm wide, surrounded by common
involucre of white membranous bristles, 6-8 mm long, reflexed.
Receptacle pyriform or obovoid, 4-6 mm long. Capitula 14-15 mm
long, bristles surrounding them numerous, somewhat flat, colorless, 8—9
mm long. Involucral bracts 16-20, in four or five rows, very finely
scabrous, pale; outer bracts 9-10 mm long, membranous, almost filiform,
cup-shaped above, brownish and toothed, with two or three long cilia
on each side along margin; bracts of next two rows 12-14 mm long,
carinate, acuminate, with appressed cilia along margin, brownish on inner
side; inner bracts connate over more than one-third length, dark brown
or even blackish on inner side, free above, acuminate and fimbriate;
corolla tube with few glandular hairs below throat. Achene fusiform,
densely covered with yellowish appressed hairs enclosing pappus.
Flowering VIII-IX.

20

20

Pistachio plantations of the semidesert zone of foothills.—Soviet
Central Asia: Pamiro-Alai (southern Tadzhikistan). Endemic. Described
from Kafirnigan, from the slopes between the Aikhatyn well and
Chilchik. Type in Leningrad.

13. E. dubjanskyi Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada, III
(1922) 176.

Perennial. Stem 25-60 cm high, solitary or two, simple or sparsely
branched above, densely white-tomentose throughout, without glandular
hairs. Leaves lanceolate, coriaceous, similarly colored on both sides,
grayish, finely white-tomentose, without glandular hairs, veins prominent
beneath; basal leaves 15—25 cm long, 2-6 cm wide, pinnatipartite, with
oblong prickly-toothed lobes, petiole broadened near base; cauline leaves
reducing above, distant, sessile, auriculate; upper leaves linear-lanceolate,
acuminate, spinescent and prickly-toothed. Heads 2—3.5 cm wide.
Capitula 12-14 mm long; bristles surrounding them half as long as
capitulum, unequal, scabrous. Involucre of 20—22 free and smooth bracts;
outer bracts linear, about 7 mm long, almost as long as bristles, broadened
above, setaceous-ciliate, acuminate; middle bracts lanceolate, carinate,
widest, membranous and setose along margin, with flexuous cilia, long
acuminate; inner bracts lanceolate, dark brown, bluish above, setaceous-
ciliate from midpoint, with scabrous cilia, acuminate. Corolla bluish,
deeply parted; tube glandular. Achene with dense yellow hairs; pappus
of yellowish bristles fused at base. Flowering VI; fruiting VII.

Sandy and gravelly deserts of the Aralo-Caspian region, on consolidated
hummocky sands and in lowland basin.—Soviet Central Asia: Aralo-
Caspian (Ust-Urt), Kyzyl-Kum (Karakalpakia), Kara-Kum (Karabugaz).
Described from the Sam sands. Endemic. Type in Leningrad.

14. E. subglaber Schrenk in Bull. Acad. Sc. Pétersb. III (1845)
110; O. and B. Fedtsch. Perech. Rast. Turkm. IV, 223.

Perennial. Stems two to four, arising from villous-tomentose caudex
at base, with remains of previous year’s leaves, weakly white arachnoid-
hairy mainly in lower part and with numerous short-stalked glandular
hairs throughout. Leaves green on both sides, rough above, densely
glandular beneath along petiole and veins, lanceolate, pinnatilobate in
upper and pinnatipartite in lower part of lamina, with lobes wide apart,
sharply prickly-toothed due to spinules into which prominent veins
terminate; lower cauline leaves 20-25 cm long and 3-4 cm wide in
upper broadest part, cauline leaves reducing upward; upper leaves linear-
lanceolate, prickly-toothed. Heads about 3 cm wide. Capitula 14-16 mm
long; bristles surrounding them 4—5 mm long, somewhat flat and white.
Involucre of 25 bracts; outer bracts scarious, 6-7 mm long, in upper

21

21

part rhomboidal, carinate, ciliate; middle bracts free, 12-13 mm long,
bluish and finely scabrous above, appressed ciliate; inner bracts connate
at base, brownish below, scarious, longitudinally carinate, ciliate-fimbriate
above but not acute, slightly shorter than middle bracts. Corolla
whitish(?), deeply parted; tube pubescent on outer side around throat;
stamen tube exserted from throat (and involucral bracts) by 7-8 mm,
almost equaling corolla lobes.

Sandy-gravelly desert——Soviet Central Asia: Aralo-Caspian (extreme
southeast), Balkhash Region (south). Endemic. Described from Balkhash.
Type in Leningrad.

15. E. pubisquameus Ijin in Bot. Mat. Gerb. Bot. Sada, III (1922)
Mise

Perennial. Root two or three-headed. Stems one or two, 25—30 cm high,
white-tomentose at base due to pubescence of leaf remains, weakly
branched, with glandular hairs on very short stalks, sometimes indistinct
due to white arachnoid-hairy pubescence preserved below along edges.
Lower cauline leaves 10—15 cm long, 3-4 cm wide, amplexicaul along
petiole; cauline leaves shorter and narrower, decurrent, lower pinnatipartite,
upper pinnatifid into deltoid lobes, sharp prickly toothed; leaves grayish-
green above, arachnoid-hairy and scabrous white-tomentose beneath with
prominent veins. Heads 2.2—2.5 cm wide. Capitula 12-13 mm long,
bristles surrounding them white, somewhat flat, unequal, scarcely
reaching 4 mm. Involucral bracts 16-20, arachnoid-hairy on outer side
in nonoverlapping part; outer bracts rhomboidal and acuminate, about 5
mm long, spiny-toothed; middle bracts enlarged to 12 mm, spinescent,
prickly-ciliate along margin, pale green above, brownish below; inner
bracts connate in lower one-third, dark brown, free and prickly-serrate-
ciliate above. Corolla bluish, parted over 4 mm, with glabrous tube.
Achene densely covered with appressed yellowish bristles; pappus bristles
free above. Flowering VII; fruiting VIII.

Gravelly slopes of lower mountain zone.—Soviet Central Asia: Tien
Shan (foothills of Karatau). Endemic. Described from the Ulkun-Buruk
Mountains. Type in Leningrad.

Series 3. Dasyanthi Bobr.—Corolla tube glandular and spreading-
hairy above. Plants of mountain ranges bordering the Fergana valley.

16. E. dasyanthus Rgl. and Schm. in Izv. Obshch. Lyubit. Estestv.
Antrop. Etnogr. 34, 2 (1882) 47; O. and B. Fedtsch. Perech. Rast. Turkm.
IVig225:

Perennial. Root two or three-headed. Stems one to three, 20—-40(60)
cm high, covered at base with remains of previous year’s leaves, white-
tomentose, partly glabrescent toward end of vegetative period, with short

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22

branches. Leaves coriaceous, grayish-green, sparsely arachnoid-hairy,
without glandular hairs and grayish-tomentose ‘beneath; basal eaves
petiolate; pinnatipartite, with prickly, crenate-toothed, narrow (0.5 cm)
leaf axis, lanceolate, up to 20 cm long and 3-4 cm wide; cauline leaves
sessile, amplexicaul-auriculate; upper leaves 3-5 cm long, 1—-1.5 cm
wide, with short, deltoidly acuminate, prickly lobes. Heads solitary,
terminal on branches, 2.5—-3 cm wide, light blue. Capitula about 15 mm
long, receptacular bristles numerous, one-third to one-fourth as long as
capitulum. Involucral bracts 20, free, bluish; outer bracts scarcely
exceeding bristles, broadened above, broadly ciliate; middle bracts
lanceolate, carinate, setose-ciliate along margin, acuminate, spinescent;
inner bracts long, attenuate-acuminate, almost entire. Corolla light bluish,
deeply divided, with glandular tube, upper part of tube densely pubescent
with long and spreading white hairs. Achenes with dense yellow hairs
above; pappus of scabrous bristles, fused over two-thirds length.
Flowering VIII; fruiting IX.

Gravelly slopes of middle mountain zone—Soviet Central Asia:
Syr-Darya (northern border of the Pamiro-Alai Region). Endemic.
Described from Kokand Region, Shahimardan. Type in Leningrad. ;

17. E. tjanschanicus Bobr. sp. n. in Addenda, XXVI, 714.

Perennial. Root two or three-headed. Stems one to three, 25—50 cm
high, covered at base with remains of previous year’s ledves, entirely
densely arachnoid-hairy or loosely white-tomentose with one, less often
two, branches in upper one-third. Leaves coriaceous, grayish or grayish-
green above, sparsely arachnoid-hairy and glandular; glandular hairs
small, sessile, short-stalked on petioles; almost white-tomentose beneath
due to light-colored dense arachnoid-hairy pubescence; basal leaves
petiolate, lanceolate, up to 20 cm long and 3-4 cm wide in upper one-
third part, deeply pinnatipartite, with prickly-toothed lanceolate lobes
and narrow crenate rachis, lobes reduced and spiny toward lower part
of leaf; cauline leaves sessile, amplexicaul, spinose towards base; upper
leaves lanceolate, sharp prickly-toothed. Heads 3-4 cm wide, bluish.
Capitula 14-15 mm long, receptacular bristles few, unequal, one-fourth
to one-fifth as long as capitulum. Involucral bracts 20-21, free, bluish,
scabrous, slightly recurved in upper part; outer bracts smallest, two times
as long as bristles, lanceolate, acute, carinate, ciliate, more attenuate
and gradually elongating toward inner bracts; innermost bracts long ciliate
in upper part, connate and dark brown in lower half. Corolla light bluish,
deeply parted, with glandular tube and fewer short hairs. Achene densely
appressed hairy; pappus of scabrous unequal bristles, free above.
Flowering VIJ-IX. (Plate I, Fig. 2.)

Mountain-steppes and juniper forest zone.—Soviet Central Asia: Tien

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23

Shan (southern range of Kuramin, Uzun-Akhmat, Airtash, Uyunkur,
Naryn). Endemic. Described from the Kuramin Range.

Type in Leningrad.

Note. This species is distinguished from S. dasyanthus Rgl. and
Schm. by the following characters: leaves sparsely arachnoid-hairy above
and with fine sessile glandular hairs which are short-stalked on petioles;
corolla tube puberulent.

Series 4. Karatavici Bobr.—Leaves grayish-green above, usually
with arachnoid-hairy pubescence or glandular, less often scabrous, without
arachnoid-hairy pubescence, leaves white- or grayish-tomentose beneath.
Plants of Tien Shan and Pamiro-Alai.

18. E. chantavicus Trautv. in Bull. Soc. Nat. Mosc. XXXIX, 2
(1866) 365; O. and B. Fedtsch. Perech. Rast. Turkm. IV, 224.—E.
hypoleucus Rupr. in Mém. Acad. Pétersb. sér. VII, XIV (1869) 4.

Perennial. Stems 50-80 and up to 100 cm high, simple or with few
branches at extreme tip, without glandular hairs, almost glabrous or
weakly arachnoid-hairy in lower part, densely arachnoid-hairy to almost
white-tomentose above. Leaves green and rough above, white-tomentose
beneath,without glandular hairs on both sides; lower leaves 30-40 cm
long, reduced above to 10 cm; all leaves deeply pinnatifid, sometimes
narrowly decurrent along leaf axes, with large and distant ovate lobes,
with unequal teeth terminating in short spinules, spinescent. Heads
solitary, 34 cm wide, with glabrous receptacle. Capitula 2—2.5 cm long,
turbinate in lower part, acutely angled-prismatic in upper part; bristles
surrounding them one-fourth to one-fifth as long as capitulum, unequal,
slightly broadened above. Involucral bracts almost entirely free, up to
25, imbricate, without glandular hairs, glabrous, smooth, carinate,
acuminate-spinescent, short- and hard-prickly from middle, bluish above;
outer bracts shortest, spatulately broadened and ciliate above; middle
and inner bracts brownish below, bluish above. Corolla blue, exserted
from scales. Appendages of anthers acute, short, apical appendages
scarcely barbate. Achene densely hairy; pappus setose, bristles free above,
ciliate. Flowering V—VI; fruiting VI—VII.

Gravelly semidesert.—Soviet Central Asia: Balkhash Region (in the
south from Khantau to Chu-Ili mountains and in the Dzharkent Region
in southeast), Tien Shan (central Tien Shan). Endemic. Described from
the Khantau Mountains. Type in Leningrad.

19. E. karatavicus Rgl. and Schm. in Tr. Peterb. Bot. Sada, V
(1877) 256; O. and B. Fedtsch. Perech. Rast. Turkm. IV, 225.—Exs.:
H.F.A.M. No. 39.

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24

Perennial. Stems 60-80 cm or more high, with remains of dead
leaves at base, arachnoid-hairy pubescent and glandular below, finely
arachnoid-hairy or loosely white-tomentose above, branched from middle.
Leaves almost coriaceous; lower leaves lanceolate, 15—25 cm long, short-
petiolate, reduced upward, amplexicaul; all leaves pinnatifid, with broad,
pinnatilobate, prickly-toothed lobes, green above and sparsely arachnoid-
hairy, with scattered stalked glandular hairs, gray-tomentose beneath.
Heads solitary, terminal on branches, 5—6 cm wide. Capitula 2—2.5 cm
long; bristles surrounding them somewhat flat, 5—6 mm long. Involucre
almost as long as florets, free, glabrous, slightly colored; outer bracts
spatulate above, with scarious border, toothed; middle bracts lanceolate,
deltoidly broadened, ciliate; inner bracts gradually narrowed above,
greenish, short-ciliate. Corolla pale blue, about 1.5 cm long, with
glandular tube. Achenes densely hispid, narrow, clavate; pappus as crown
of scabrous bristles fused over two-thirds their length. Flowering VI-—
VII; fruiting VII—VIII.

— Gravelly slopes in middle mountain zone.—Soviet Central Asia:
Tien Shan, Syr-Darya (Fergana Valley), Pamiro-Alai (Alai Range,
Lyangar). Endemic. Described from the Chirchik Valley. Type in
Leningrad.

20. E. obliquilobus Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada, III
(1922) 180.

Perennial. Stem 40—60—80 cm high, shallow-sulcate, almost entirely
floccose-loosely tomentose, also with short-stalked glandular hairs,
branched above, white-tomentose and without glandular hairs below heads.
Leaves thin-coriaceous, green, very sparsely white arachnoid-hairy along
veins, with scattered glandular hairs and scabrous above, white-tomentose
beneath; basal leaves 30-40 cm long, with prickly flattened petioles;
cauline leaves sessile, narrowed toward base; upper leaves not narrowed,
smallest, very short-decurrent; all leaves pinnatifid, with more or less
oblong-deltoid, antrorse lobes, prickly along margin, and scarcely prickly
along broad incisions. Heads solitary or a few, 3.5—4 cm wide. Capitula
16-18 mm long; bristles surrounding them few, 4-6 mm long, unevenly
scabrous and somewhat flat. Involucre of 25—30 free and smooth bracts;
outer bracts linear, spatulately broadened above, setose along margin;
middle bracts lanceolate, carinate, cristate-ciliate along margin, acuminate-
aristate; inner bracts thin coriaceous, aristate. Anther appendages barbate
above. Pappus bristles fused at base. Flowering VII; fruiting VIII-IX.

Limestone outcrops of middle mountain zone.—Soviet Central Asia:
Pamiro-Alai (southwest; Gissar Range and Surkhan-Darya Province).
Described from the Khakimi Mountains near Karataga River in the Gissar
Range. Type in Leningrad.

aS

25

21. E. babatagensis Tschern. in Fl. Uzb. VI (1962) Addenda.

Perennial. Stem 20-30 cm high, shallow-sulcate, finely tomentose
and glandular, with few short branches above. Leaves with sessile
glandular hairs above and weakly arachnoid-hairy at isolated points,
grayish-tomentose beneath, with prominent midrib, basal leaves petiolate,
pinnatipartite, with lanceolate prickly-toothed lobes and narrow and
flat prickly-small toothed rachis; middle and upper leaves sessile,
semiamplexicaul, auriculate; upper leaves pinnatilobate, short-decurrent.
Heads terminal on branches, about 3.5 cm wide. Capitula 10-20 mm
long; bristles surrounding them few, flat, 4-5 mm long. Involucral bracts
28—30; outer bracts linear, scarcely longer than bristles, broadened above,
ciliate; middle bracts 7-15 mm long, lanceolate, carinate, setose-ciliate
along margin, acuminate-aristate; inner bracts lanceolate, 18-20 mm long,
brownish on inner side over more than half length, ciliate along margin,
aristate. Corolla pale blue. Achenes deep yellow; pappus bristles coronate
in lower part, fused, split. Flowering VII-VIII; fruiting VIII-IX.

Limestone and gypsic deposits of middle mountain zone.—Soviet
Central Asia: Pamiro-Alai (southwest—northern part of the Babatag
Range). Endemic. Described from the place indicated. Type in Tashkent.

22. E. fedtschenkoi Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada, III
(1922) 177.

Perennial. Stems two or three, 40-50 cm high, covered with remains
of previous year’s leaves and angular-ribbed below, white pubescent
along ribs, yellowish-brown along furrows, with stalked glandular hairs
in upper and middle part, stems as well as their branches cylindrical,
white-tomentose, without glandular hairs. Lower cauline leaves 25-30
cm long, 4-6 cm wide, with prickly petiole, amplexicaul, lanceolate,
deeply pinnatifid into broad deltoid, large prickly-toothed lobes with
narrow sinuses much smaller than lobes, spinulose along margin; leaves
gradually reduced upward and broad-lanceolate, large prickly-toothed,
4-6 cm long, 1—2 cm wide, densely punctate, grayish-green, arachnoid-
hairy, and without glandular hairs above, grayish-tomentose, with
prominent veins, less densely pubescent-hairy and without glandular hairs
beneath. Heads 4-5 cm wide. Capitula 18-20 mm long; bristles
surrounding them about 6 mm long, white, flat, sometimes ciliate-truncate
at tip. Involucral bracts 18-20; outer bracts scarcely exceeding bristles,
rhomboid above, brownish, carinate, ciliate; middle bracts (9-10)
roundish-carinate, finely scabrous on outer side, acuminate, reflexed,
ciliate-setose along margin, brownish in lower part, bluish and punctate
above; inner bracts free, glabrous, shorter than middle, incised-ciliate-
toothed from middle. Corolla deeply divided, bluish, glandular on outer

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26

side. Achenes hispid-strigose; pappus bristles rusty, fused at base.
Flowering VII; fruiting VIII.

Middle mountain zone —Soviet Central Asia: Pamiro-Alai (Darvaz).
Endemic. Described from Kamchirak Pass. Type in Leningrad.

Series 5. Orientales Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR,
VIII (1950) 70 p. p.—Outer involucral bracts broad-rhomboidal or
spatulately broadened above, with short cusp; middle bracts lanceolate,
attenuate-acuminate. Plants of Transcaucasia and Dagestan.

23. E. orientalis Trautv. Diss. Echin. (1833) 22; DC. Prodr. VI,
526; Iljin in Grossh. Fl. Kavk. IV, 161; Mulk. in Tr. Bot. Inst. Akad.
Nauk ArmSSR, VII, 72.—E. horridus acut. non Desf. (1804); Ldb. FI.
Ross. IJ, 656; Bge. in Bull. Acad. Pétersb. VI, 407; Boiss. Fl. or. III,
438 p. p.; Iljin, op. cit—Ie.: Trautv. op. cit., f. 8.

Perennial. Stem about 70 cm high, solitary (?), simple, sulcate, white-
pubescent, almost white-tomentose above, also with numerous glandular
hairs throughout surrounding brownish, flat, and curly bristles, imparting
reddish-brown tinge to stem. Leaves lanceolate; basal leaves 25-35 cm
long; cauline leaves shorter, bipinnatipartite, with lanceolate lobes,
divided into long-deltoid lobes, prickly-toothed, attenuate into 1—1.5 cm
long spine; upper cauline leaves simple pinnatifid to long prickly-toothed;
leaves scabrous above due to dense cover of stalked glandular hairs,
also somewhat arachnoid-hairy, white-tomentose beneath with few,
brown, stalked glandular hairs along veins. Heads 4.5—6 cm wide.
Capitula 30-35 mm wide, cuneately narrowed toward base; bristles
surrounding them about 12 mm long, almost half as long as young
capitula, numerous, somewhat flat. Involucral bracts scabrous, 20; outer
bracts not longer than bristles, spatulately broadened, mucronate and
serrate, brownish on outer side; middle bracts about 25 mm long,
broadened in middle, long acuminate, appressed serrate-ciliate along
margin; inner bracts connate at base, shorter than middle, acute and
fimbriate-incised above. Corolla pale blue, deeply divided. Achenes
densely appressed-setose; pappus bristles fused and toothed near base.
Flowering VI; fruiting VII.

Foothills of eastern Caucasus in the semidesert zone.—Caucasus:
Dagestan, eastern Transcaucasia. General distribution: Iran (northwest).
Described from Derbent. Type in Leningrad.

Note. The classical authors De Candolle, Ledebour and Boissier
considered E. horridus Desf. as the prior name of this species. However,
we are not confident that they were correct. This is because E. horridus
Desf. was described in a few words and does not characterize the plant
at all. The specimens that could be considered as type specimens

27

27

apparently belong to different species. We had an opportunity to examine
the E. horridus specimens from Schrader’s herbarium, sent to him by
Pearson with a reference to the Paris Garden (“H.P.”). These specimens,
in our opinion, should be included under £. orientalis Trautv. The
specimen from the herbarium of Mertens (“11 Nov. 1807, Hort. Paris”)
is only a portion of the upper part of a stem with terminal leaves. This
plant does not resemble E£. orientalis Trautv.

For a final solution of the problem concerning the name of this
species, it is essential to study the specimens of the plant cultivated in
Paris in 1804, which are the type specimens of E. horridus Desf.

24. E. opacifolius [ljin ex Grossh. Fl. Kavk. IV (1834) 159; Bot.
Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, VII, 1, 60; Mulk. in Tr. Bot.
Inst. Akad. Nauk ArmSSR, VIII, 75.—E. bipinnatus auct. non Freyn
1894; Iljin in Grossh. Fl. Kavk. IV, 160; Grossh. Opred. Rast. Kavk.
471.

Perennial. Stem 30—50 cm high, simple, sulcate, arachnoid-hairy or
almost tomentose and also glandular in lower part, leafy in lower one-
third part. Leaves oblong, pinnatipartite or pinnatisect, with lanceolate
or oblong, prickly-toothed lobes; basal and lower cauline leaves petiolate,
with spinule at base, often trifid; upper leaves sessile, semi-amplexicaul;
leaves finely and very loosely tomentose above, white-tomentose beneath.
Heads 4—5 cm wide. Capitula 20-24 mm long; bristles surrounding them
unequal, white, half to one-thir¢ as long as capitulum. Involucre of 20—
25 glabrous bracts, five or six outer bracts not longer than bristles,
spatulately broadened above, short acuminate and ciliate, next five or
six bracts with broader cuneate base, long acuminate; middle six or
seven bracts linear-lanceolate, dark in lower part, long acuminate and
ciliate-toothed above middle; inner bracts lanceolate, connate in lower
half, dark, pale green above, acuminate and fimbriate-ciliate. Corolla
pale, very deeply divided. Achenes densely appressed-setose; pappus of
yellowish bristles, free above. Flowering VII; fruiting VIII.

Foothill zone.—Caucasus: Southern Transcaucasia (Nakhichevan
ASSR), Talysh. Endemic. Described from the Nakhichevan ASSR. Type
in Baku.

25. E. connatus C. Koch in Linnaea, XXIV (1851) 381; Iljin in
Grossh. Fl. Kavk. IV, 161; Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR,
VIE. 76.

Perennial. Stem about 1 m high, ribbed, arachnoid-hairy, with
numerous thick-stalked glandular hairs, branched above, almost white-
tomentose. Leaves lanceolate, 15-25 cm long, bipinnatifid into distant
deltoid or lanceolate toothed lobes, spinescent; leaves not coriaceous,

28

28

glabrous above, but with thick-stalked glandular hairs, almost white-
tomentose beneath, with few stalked glandular hairs along veins.
Receptacular bristles half as long as capitulum, somewhat flat, whitish.
Involucral bracts glabrous; outer bracts rhomboidal above, with dark
brown spot in broader part, toothed, short acuminate; middle bracts
lanceolate, ciliate, acuminate-cuspidate; inner bracts shorter than middle,
oblong-linear, ciliate, connate in lower half; pappus of reddish barbed
bristles.

Caucasus: Southern Transcaucasia (former Olty District of Kars
Region). General distribution: Asia Minor. Described from the former
Olty District. Type specimen lost.

Note. This is a poorly known and extremely problematic species.
Boissier included it among the synonyms of the East Mediterranean
species E. viscosus DC., distributed from Crete to Syria and Kurdistan.
Ya.I. Mulkijanyan saw the plants preserved in Tbilisi, collected by D.I.
Sosnowsky in the former Olty District, which can be considered as having
originated from the classic locality. These specimens are also inadequate
and do not permit complete characterization of this plant.

M.M. Iljin (op. cit.) suggested that the species of Koch is “one of
the forms of the widespread species E. pungens Trautv. in Trans-
caucasia.”

Ya.I. Mulkiyanyan included E. macrophyllus var. glabricaulis Bornm.
among the synonyms of E. connatus C. Koch. The former name was
proposed by Bornmiiller for the specimens collected by Yu.N. Woronow
(VII, 1911) from the vicinity of Artvin. These specimens are in the
herbarium of Bornmiiller in Weimar. The opinion of I. Bornmiiller, the
very authoritative botanist who specialized in the study of the Iranian
globe-thistles and saw many original specimens, is very important.
Nevertheless, I take the liberty to express doubt concerning the identity
of the East Anatolian species E. connatus C. Koch with the Iranian
species E. macrophyllus Boiss. and Hausskn., distributed only in the
western Kopetdag within the limits of the USSR.

Series 6. Macrophylli Bobr—Lower cauline leaves undivided and
with relatively wide axial part in sinuses between lobes. The only member
of this Iranian group of species in our flora is E. macrophyllus Boiss.
and Hausskn. Apparently, E. jesdianus Boiss. and Buhse belongs to this
series.

26. E. macrophyllus Boiss. and Hausskn. in Boiss. Fl. or. III (1875)
426; Bornm. in Beih. bot. Centralbl. XXXVI, 2, 211; Fl. Turkm. VII,
146.

WA

BeQV,:
VA Ay
(] yk

= 4,

re
EUR

30

30

Perennial. Stem 1—1.5(2) m high, about 2 cm in diameter, simple at
base, branched above, entirely sulcate, sparsely white arachnoid-hairy
along ribs, densely glandular with glandlar hairs on curly bristles. Lower
cauline leaves 25-40 cm long, axial part entire, 1.5—-3 cm wide in sinuses,
lobes up to 10 cm wide, sessile, semiamplexicaul, lanceolate, with
pinnately lanceolate prickly-toothed lobes up to 6 cm long and 1-—1.5
cm wide, prickly-toothed with veins terminating in spinules, with broad
and deep incisions between teeth, and short spines along margin; leaves
gradually reduced upward along stem; upper leaves lanceolate, 10—12
cm long, 2-3 cm wide, not lobed, prickly-toothed; grayish-green above,
densely glandular-hairy with papillate stalks, grayish-tomentose beneath,
with prominent veins and glandular hairs along veins. Heads solitary
terminal on branches, 8-10 cm wide. Capitula 22-24 mm long, with
about 32 mm long florets; bristles surrounding them 12—14 mm long.
Involucre of 16 glabrous bracts; outermost bracts shorter than bristles,
lanceolate, rhomboidal above, brownish, ciliate; middle bracts about 20
mm long, narrowed-subulate from middle, bluish, appressed ciliate along
margin, appearing serrated; inner bracts connate in lower one-third, acute,
not subulate, colored and ciliate. Corolla tube glandular below throat,
deeply parted into linear, about 10 mm long lobes. Pappus bristles fused
only at base. Flowering VI; fruiting VII. (Plate I, Fig. 1).

Found in shady places in ravines, among shrubs.—Soviet Central
Asia: mountainous Turkmenia (southwestern Kopetdag). General
distribution: Iran. Described from southeastern Iran. Type in Weimar.

Note. Bornmiiller, in comparing the plants collected by Schintenish
in the western Kopetdag with the original specimens in the herbarium
of Hausknecht, noted their complete identity. His suggestion (Beih. bot.
Centralbl., XXXVI, 2, 1918, 211, 212) that the plant collected by Yu.N.
Woronow (VII, 1911, No. 6089) along the Chorokh near Artvin should
be included under this species is extremely interesting. These specimens
differ, according to Bornmiiller, only by glabrous stems, which is why,
he named them E. macrophyllus var. glabricaulis Bornm. See also
note to E. connatus Koch.

Section 3. Ritro Endl. Gen. pl. (1836) 467 p. p.; Bge. in Bull.
Acad. Sc. Pétersb. VI (1863) 401, 408; Mulk. in Tr. Bot. Inst. Akad.
Nauk ArmSSR, VIII (1950) 35; Boiss. Fl. or. II] (1875) 425 ex. p—
Involucral bracts 18-25; outer bracts without glandular hairs; leaves
coriaceous. Plants from plains, mountain steppes, and semideserts.

Type of section: E. ritro L.

Series 1. Tenuifolii Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR,
VIII (1950) 65.—Receptacular bristles one-third to one-fourth as long
as capitulum; outer involucral bracts not longer than adjacent bristles.

3]

a1

27. E. ritro L. Sp. pl. (1753) 815, quoad. B.; Trautv. Diss. Echin.
23; Ldb. Fl. alt. IV; Ldb. Fl. Ross. II, 654; Schmalh. Fl. I], 91; Kryl.
Fl. Zap. Sib. XI, 2868; Grossh.. Fl. Kavk. IV, 158; Mulk. in Tr. Bot.
Inst. Akad. Nauk ArmSSR, VIII, 65.—E. ritro B. tenuifolius DC. Prodr.
VI (1837) 524.—E. tauricus Willd. ex Ldb. FI. alt. VI (1831) 44.—E.
tenuifolius Fisch. ex Schkuhr, Handb. III (1808) 181, nom.—E. ruthenicus
auct. non MB. (1819); Vizn. Rosl. UkrSSR, 558; Stank. and Tal. Opred.
(1957) 401.—E. caule... Gmel. Fl. Sib. I] (1749) 102.—Ic.: Gmel. Ibid.,
tab. XLVI: Trautv. op. cit. Fig. 9: Fl. Yugo-Vost. VI, Fig. 693.—Exs.:
GRF No. 2028.

Perennial. Stems solitary or fewer, 20-60 cm high, simple or with
few branches above, surrounded at base by remains of previous year’s
leaves, cylindrical, leafy, white arachnoid hairy-tomentose. Leaves, except
basal, sessile, 10-20 cm long, 3—10 cm wide, semiamplexicaul, deeply
pinnati- or bipinnatisect into lanceolate prickly-toothed lobes, dark green,
glabrous or weakly arachnoid-hairy above, white-tomentose beneath.
Heads one to five, solitary terminal on stems and branches, light bluish,
3.54.5 cm wide; bristles of common involucre broadened above, 5 mm
long. Capitula 12-17 mm long; bristles surrounding them about 4 mm
long. Involucre of 20-22 bracts; outer bracts up to 4 mm long, deltoidly
broadened above, ciliate; middle bracts spatulate; inner up to 15 mm
long, lanceolate, carinate, ciliate along margin, greenish on inner side.
Achenes cylindrical, 6 mm long; pappus bowl shaped, of scarious bristles,
fused in lower half, free above, ciliolate. Flowering V—VI; fruiting VI—
VI.

Predominantly found in steppe zone (less often forest-steppe zone),
steppe meadows on rubbly slopes and outcrops, and on chalk —European
Part: Upper Volga (south), Volga-Don, Trans-Volga, Volga-Kama
(south), middle Dnieper, Bessarabia, Black Sea Region, Lower Don,
Lower Volga, Crimea; Caucasus: Ciscaucasia; Western Siberia: Ob
(south), Upper Tobol, Irtysh, Altai; Soviet Central Asia: Aralo-Caspian
Region, mountainous Turkmenia, Balkhash Region, Dzhungaria-
Tarbagatai. General distribution: Central Europe (introduced), Iran,
western China. The figure cited by me in Gmelin’s “Siberian Flora”
can be considered as the type of this species.

Note I. My proposal to consider the plants shown in the figure in
Gmelin’s Flora as the type of this species is only a preliminary solution
of the nomenclatural problem arising in connection with the selection
of a type specimen. The matter is as follows. While establishing this
species, Linnaeus (op. cit.) cites in the protologue two polynomial names
proposed by Gmelin for the West European plants.

The first of them (Geml. Fil. Sib. II, 100) is E. latifolius Tausch,
and the second, which Linnaeus denoted with the letter 8. (Gmel. op.
cit. 102, tab. 46), Russian investigators have always considered to be

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true E. ritro L. As regards the western European plants, the citations of
Bauhin, Lobelius, and Dalechamp probably refer to some other southern
European plant, possibly to the one designated by De Candolle as E.
ritro a. vulgaris DC. or to the one for which Bieberstein proposed the
name E. ruthenicus MB. It is possible, however, that the names proposed
by De Candolle and Bieberstein pertain to the same southern European
species. To solve the nomenclatural problem it is necessary first to
determine the species to which the plant grown by Linnaeus in the
Uppsala Garden and cited first, i.e., the formal type specimen of E.
ritra. L., belongs.

The Russian name of this plant is common globe-thistle; the Latin
name proposed for it by Linnaeus is “Ritro,” taken by him from the
article by Matthias Lobelius (1538-1616), in which it was used as a
generic epithet. The word “ritro” is the vernacular name of the southern
European globe-thistle.

__ Note 2. The extreme polymorphism of this species was noted already
in the first monograph of the genus by Trautvetter (1833). This tendency
was mentioned also by De Candolle, Ledebour, Korshinsky and Krylov.
Of the forms more or less deserving special study, E£. ritro 6. litwinowii
Korsh. (Tent. Fl. Ross. or., 1898, 231), distinguished by extremely narrow
lobes of the leaves, should be mentioned, which sometimes is an indicator
of the semidesert of western Kazakhstan. In eastern Kazakhstan, var.
latilobus Kryl. (op. cit.), which is characterized by leaves with broad
lobes, shallowly and simply pinnatifid (and not bipinnatisect), the upper
cauline ones lanceolate and sharply toothed, is distributed in eastern
Kazakhstan. Plants with such leaves are predominant in the southern
Altai, the Zaisan Basin, Tarbagatai, and the foothills of the Dzhungarian
Alatau. At the same time, typical specimens of E£. ritro L., with the
lobes of the deeply bipinnatisect leaves narrow, are also found there.
The wide range of the species, occurring in a diversity of habitats, and
its tendency to vary, promote differentiation into races; the latter,
however, have not yet been defined sufficiently clearly. After special
study, probably the western Kopetdag race of E. ritro s.1. will have to
be segregated as an independent race.

28. E. meyeri (DC.) Ijin in Fl. Yugo-Vost. VI (1936) 376 comb.—
E. ritro y. meyeri DC. Prodr. VI (1837) 524; Bge. Reliq. bot. 349.—E.
ritro y. glandulosus Korsh. Tent. Fl. Ross. or. (1898) 231.

Perennial. Stem 20-50 cm high, relatively slender, brownish, not
tomentose, densely glandular-hairy, slightly branched above. Basal leaves
petiolate; cauline leaves sessile, white-tomentose beneath, green above,
glabrous or slightly arachnoid-hairy, simple or bipinnatilobate, with
oblong, prickly-toothed or aristate lobes. Heads light blue, globose,

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33

solitary, 2.5—3.5 cm wide; bristles surrounding them almost half as long
as inner involucral bracts, somewhat flat, toothed. Involucral bracts 14—
20, scaly, bluish; outer bracts linear-spatulate; acuminate, ciliate; middle
bracts lanceolate, spinescent; inner bracts cuspidate; corolla bluish, tube
glandular. Achenes with long hairs; pappus bristles fused over basal
half. Flowering V—VI, fruiting VI-VII.

Zone of dry steppes, semidesert and desert; in the steppe zone on
projecting rocks and along rubbly slopes——European Part: Trans-Volga
Region (south); Volga-Kama (extreme south), Lower Volga; Western
Siberia: Upper Tobol (south); Soviet Central Asia: Aralo-Caspian Region
(up to Ust-Urt in the south), Kyzyl-Kum (Karakalpakia). Endemic. The
locality was not indicated while establishing y. meyeri DC. It may be
assumed that the type material could have originated from southeast of
the European part of the USSR. To a certain extent, the plants of Leman,
collected at Lake Inderskoe, could be considered as type specimens.

29. E. spiniger [ljin ex Bobr. in Tr. Bot. Sada Akad. Nauk SSSR,
XLIV (1931) 75, nomen; Fl. Turkm. VII, 147, descr. ross.

Perennial. Stems 30-50 cm high, covered at base with remains of
previous year’s petioles, strongly branched, densely arachnoid-hairy,
without glandular hairs. Leaves hard, bi- or tri-pinnatisect into narrow,
hard, spinulate, divergent lobes, greenish above with scattered short-
stalked or sessile glandular hairs and white arachnoid-hairy midrib, white-
tomentose beneath with narrow border. Basal and lower cauline leaves
15-20 cm long, petiolate, broadened and ciliate at base; upper leaves
reduced and reaching almost to heads. Heads 3—3.5 cm wide. Capitula
at flowering 20-22 mm long; bristles surrounding them 5—6 mm long,
broadened and toothed above. Involucre of 18—20 free glabrous bracts;
outer bracts not longer than bristles, scarious, spatulate, brownish,
acuminate and long-ciliate; middle bracts oblong-lanceolate, with
prominent veins on outer surface, acuminate, long-ciliate; inner bracts
scarious, free, acuminate above, laterally divided into two or three
narrow-lanceolate parts. Corolla deeply incised, narrowed below throat
and densely hairy. Achenes densely covered with yellowish bristles;
with crown of hairs fused at base. Flowering VI; fruiting VII. Plate II,
Fig. 2).

Steppes of middle mountainous zone.—Soviet Central Asia:
mountainous Turkmenia (The Great Balkhan Range). Endemic. Described
from the western slopes near Dzhebel. Type in Leningrad.

Series 2. Pungentes Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR,
VIII (1950) 59.—Stems glandular or glabrous below (E. polyacanthus
Ijin); leaves bipinnatisect, with linear or lanceolate densely spinulose
lobes.

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34

30. E. pungens Trautv. Diss. Echin. (1833) 18, Fig. 5; Ldb. FI.
Ross. II, 656; Iljin in Grossh. Fl. Kavk. IV, 160; Mulk. in Tr. Bot. Inst.
Akad. Nauk ArmSSR, VIII, 60.—E. persicus Stev. and Fisch. ex Fisch.
Cat. horti Gorenk. IT (1812) 37, nomen; DC. Prodr. VI, 525; Boiss. FI.
or. III, 440.—E. raddeanus Somm. and Lev. in Nuov. Giorn. bot. Ital.
2 (1895) 90; Tr. Peterb. Bot. Sada, XVI, 247; Iljin, op. cit. 158.

Perennial. Stems 40-60 cm high, less often higher, thick, sulcate,
densely arachnoid-hairy, branched above, glandular below, almost white-
tomentose above. Leaves coriaceous, elliptical, bipinnatisect into narrow-
lanceolate distant lobes, with narrow winged 1—2 cm wide axis, and 2—
4 mm long spines along margin; lobes of leaves short, prickly-toothed,
teeth terminating in light colored, 6-10 mm long spine; leaves reduced
upward; lower leaves up to 25 cm long and 12 cm wide; upper leaves
lanceolate, prickly-toothed; leaves arachnoid-hairy and glandular above,
with slightly recurved margin, white-tomentose beneath. Heads 3.5—6
cm wide, solitary, terminal on branches, up to six on single plant;
common involucre of about 15 mm long scarious bristles lanceolately
broadened above. Capitula about 2 cm long, apparently usually sterile;
bristles surrounding them unequal, slightly more than one-third as long
as involucre, colorless, somewhat flat. Involucre of 15—18 scabrous bracts;
outer bracts scarcely exceeding bristles, rhomboidal above, brownish,
acuminate, ciliate along margin, with isolated hairs on outer surface;
middle bracts carinate, attenuate-acuminate, up to 15 cm long, ciliate;
inner bracts not exceeding middle ones, longer and more irregularly
ciliate. Corolla light bluish, deeply parted, with isolated hairs on fused
part. Achenes with yellowish shiny hairs; pappus of barbed bristles fused
up to basal half, unequal. Flowering VII; fruiting VIII. (Plate II, Fig. 1).

Mountain-steppe zone.—Caucasus: Southern and eastern Trans-
caucasia. General distribution: Eastern Anatolia, northeastern Iran.
Described from the Kirovabad Region. Type in Leningrad.

31. E. polyacanthus [jin in Bot. Mat. Gerb. Bot. Sada, IV (1923)
102; Ijin in Grossh. Fl. Kavk. IV, 158; Mulk. in Tr. Bot. Inst. Akad.
Nauk ArmSSR, VIII, 64.

Perennial. Stems 40-60 cm high, ribbed, with two or three branches
above, without glandular hairs, glabrous below, white-tomentose above.
Lower leaves elliptical, 20-30 cm long, 12—20 cm wide, petiolate, with
numerous spines, pinnatipartite into narrow-lanceolate lobes, in turn
divided into large prickly-toothed lobules with numerous spines along
margin, hence appearing densely prickly; leaves reduced above, sessile,
with prickly bases; upper leaves lanceolate, 3-7 cm long, large prickly-
toothed; leaves grayish-green above, slightly arachnoid-hairy with few
short-stalked glandular hairs, grayish-tomentose beneath, with prominent,
sparsely glandular veins. Heads 5—6 cm wide, on 3-4 cm long branches;

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35

common involucre of whitish, flat, 15-20 mm long bristles; bristles
surrounding them slightly less than half as long as involucre, comprising
white, flat unequal scales. Involucral bracts 17—20, finely scabrous; outer
bracts slightly raised above adjoining bristles, i.e., 10-11 mm long,
rhomboidal above and brownish on outer surface; increasingly acuminate
from middle to inner bracts, appressed-ciliate along margin; innermost
bracts sometimes slightly connate at base. Achenes covered with long
golden hairs; pappus bristles unequal, about 2 mm long, fused in lower
half. Flowering VII-—VIII.

Mountain-steppe zone.—Caucasus: Southern Transcaucasia (western
Armenia). General distribution: Eastern Anatolia. Described from the
Kagyzman District of the former Kars Province. Type in Leningrad.

32. E. conrathi Freyn in Bull. Herb. Boiss. III (1895) 356; Ijin in
Grossh. Fl. Kavk. IV, 157; Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR,
VII, 59.

Perennial. Stems up to | m high and about 1.5 cm thick, angular,
branched, white-tomentose, and without glandular hairs above, glandular
below, but not tomentose. Lower leaves oblong, up to 35 cm long,
pinnatisect into oblong-deltoid lobes, with teeth terminating in spines
up to 1 cm long, sinuses between lobes roundish, leaf bases amplexicaul,
spiny; leaves reduced upward; cauline leaves about 15 cm long; upper
leaves pinnatisect into lanceolate-toothed lobes, arachnoid-hairy and
glandular above, weakly pubescent and glandular to grayish-tomentose
beneath with thick glabrous veins bearing few stalked glandular hairs.
Heads 6—6.5 cm wide. Capitula about 25 mm long; bristles surrounding
them scabrous, unequal, almost half as long. Involucre of 16(20) bracts;
outer bracts about 13 mm long, spatulately broadened above, toothed-
ciliate along margin; middle bracts narrowed and pointed, appressed
long-ciliate along margin; inner bracts not strongly acuminate, long-
ciliate, longest, brown below on inner side, scarcely connate at base.
Corolla light blue. Achenes about 8 mm long. densely pubescent; pappus
of short barbed bristles fused at base. Flowering VII-VIII.

Rubbly and shrubby slopes of mountain-steppe zone.—Caucasus:
Southern and eastern Transcaucasia (Somkhetia, northern Armenia,
southeastern Georgia). Described from Akhtala in northern Armenia.
Type in Brno.

Series 3. Transcaucasici Mulk.—Lower cauline leaves bipinnatisect,
more than half or almost as long as stem; outer involucral bracts
spatulately broadened above, brown, scarcely longer than adjoining
bristles.

33. E. foliosus Somm. and Lev. in Tr. Peterb. Bot. Sada, XVI (1900)

36

36

248; Iljin in Grossh. Fl. Kavk. FV, 159; Mulk. in Tr. Bot. Inst. Akad.

- Nauk ArmSSR, III, 57.

Perennial. Stem 40-60 cm high, 1.5 cm thick, with few short
branches above, white-tomentose, glandular below, sometimes with
numerous glandular hairs, on somewhat flat, curly brownish bristles, or
glandular hairs fewer and covered with white tomentum. Lower leaves
up to 40-50 cm long and 10-18 cm wide, on long amplexicaul petioles
prickly along margin, ovate-lanceolate, pinnatipartite with lobes bearing
deltoid falcate teeth terminating in spines up to 1.5 cm long, roundish
sinuses along axes spiny; cauline leaves almost pinnatifid into large
deltoid lobes-teeth, with 1—-1.5 cm long spines along margin and at tip;
upper leaves undivided, large prickly-toothed, with numerous spines;
leaves arachnoid above, with scattered short-stalked glandular hairs,
white-tomentose beneath, with numerous long-stalked brownish glandular
hairs along veins. Heads 4-5 cm wide, 3—3.5 cm on lateral branches.
Capitula 2—2.5 cm long; bristles surrounding them few, unequal, white,
about half as long. Involucre of 20, free, thin scabrous bracts. Outer
bracts slightly longer than bristles, spatulately broadened above, brown,
toothed, and ciliate along margin; middle bracts attenuate-acuminate,
with roundish keel bearing long and appressed cilia; inner bracts longest,
with very long cilia along margin. Corolla light blue. Pappus of very
long bristles, unequal, fused over almost basal half. Flowering VIII-IX.

Shurbby thickets of mountain-steppe zone.—Caucasus: Eastern
Transcaucasia (Borzhomi, Abastumani District, Mtskheta, Dilizhan,
Bzovdal). Endemic. Described from Borzhomi. Type in Helsinki, cotype
in Leningrad.

34. E. transcaucasicus Ijin in Bot. Mat. Gerb. Bot. Sada, IV (1923)
104; Iljin in Grossh. Fl. Kavk. IV, 159; Mulk. in Tr. Bot. Inst. Akad.
Nauk ArmSSR, VIII, 55, Fig. 10.

Perennial. Stem 60-80 cm high, base covered with fibrous remains
of previous year’s leaf bases, up to 1.5 cm thick, white-tomentose,
without glandular hairs, leafless above. Lower leaves lanceolate, up to
45 cm long and 20 cm wide, pinnatisect into lanceolate lobes, latter in
turn divided, prickly-toothed along margin, spines up to 8 mm long,
petioles long, amplexicaul; lower leaves often as. long as stem; cauline
leaves sessile, lanceolate, pinnatifid to undivided, deeply toothed, teeth
terminating in spines up to 1.5 cm long, dark green above and grayish
due to arachnoid-hairy pubescence, without glandular hairs, grayish
beneath due to dense arachnoid-hairy pubescence or loosely tomentose.
Heads 5—6 cm wide. Capitula 22-25 mm long, with cuneate base, almost
pentagonal; bristles surrounding them numerous, somewhat flat, 12-13
mm long. Involucral bracts 17-18; outer bracts scarcely longer than

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37

adjacent bristles, rhomboidly broadened above, acuminate and spatulate-
concave, dark brown in this part, short toothed-ciliate along margin;
middle bracts longest, acuminate and attenuate-accuminate, with
appressed cilia along margin, finely scabrous on outer side; inner bracts
slightly shorter. Corolla pale blue. Achenes ribbed, covered with
yellowish hairs; pappus of long unequal bristles, fused at base. Flowering
VII.

Mountain steppes.—Caucasus: Eastern Transcaucasia (western
Georgia—Bakuryany, Dzhavakhetia). Endemic. Described from the
Akhalkalaki Region. Type specimen lost.

Note. The species is very close to E. foliosus Somm. and Lev.
described from Borzhomi. It differs by the absence of glandular hairs
on veins on the dorsal surface of leaves and lower part of stem, and by
nonfalcate teeth of the lobes of lower leaves.

Series 4. Iljiniani Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR,
VIII (1950) 48.—Stem white-tomentose, without glandular hairs or (E.
szovitsii F. and M.) glandular, with numerous long-stalked glandular
hairs in lower part of stem and along veins on lower leaf surface.

35. E. szovitsii Fisch. and Mey. ex DC. Prodr. VI (1837) 524; Ldb.
Fl. Ross. II, 656; Bge. in Bull. Acad. Sc. Pétersb. VI, 408; Boiss. FI. or.
III, 426; Iljin in Grossh. Fl. Kavk. IV, 160; Mulk. in Tr. Bot. Inst.
Akad. Nauk ArmSSR, VIII, 53.

Perennial. Stem 60-80 cm or higher, thick, branched above, sulcate,
finely white-tomentose, also glandular in middle and lower parts,
glandular hairs sometimes indistinct because of dense pubescence. Leaves
lanceolate, pinnatisect into lanceolate lobes, in turn almost pinnatifid
and sinuses between lobes and lobules occupying not more than one-
third of leaf area, prickly-toothed, spinescent; lower leaves 30-40 ‘cm
long and up to 20—25 cm wide, reduced upward; upper leaves narrow-
lanceolate, sharply prickly-toothed, leaves slightly arachnoid-hairy above,
rugose and scabrous due to abundance of short-stalked glandular hairs,
grayish-tomentose beneath with numerous light brown stalked glandular
hairs along veins. Heads 5—7 cm wide. Capitula 22-25 mm long, with
cuneate base; bristles surrounding them few, yellowish, not more than
half as long as involucre. Involucre of 18-20 scabrous bracts; outer
bracts 12-13 mm long, scarcely exceeding adjacent bristles, spatulate,
broadened, spreading ciliate, brownish on outer side; middle bracts
roundish-carinate, attenuate-acuminate, appressed ciliate, about 2 cm long;
inner bracts usually shorter than middle, fimbriate-ciliate above, connate
below at isolated points. Corolla pale blue. Achenes covered with yellow

38

38

bristles; pappus bristles barbed, unequal, fused below. Flowering VII;
fruiting VIII.

Mountain-steppe zone, at 1,500—2,500 m.—Caucasus: Southern
Transcaucasia (Daralagyoz, Zangezur), eastern Transcaucasia (Karabakh).
General distribution: Northwestern Iran. Described from Karabakh. Type
in Leningard.

36. E. iljinii Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR, VIII
(1950) 49; Bot. Mat. Gerb. Bot. Inst. Akad. Nauk ArmSSR, XVIII,
29k.

Perennial. Stem 1—1.25 m high, usually not solitary, up to 1.5 cm
thick below, cylindrical, sulcate, branched above, finely white-tomentose.
Leaves lanceolate, pinnatipartite into large-toothed lanceolate lobes; lower
leaves pinnatifid, sinuses between lobes and lobules occupying at least
two-thirds of total leaf area; lower leaves 50-70 cm long and up to 30
cm wide, with lobes up to 13 cm long and 5—7 cm wide, lobes short-
prickly-toothed, with teeth terminating in 3-5 cm long spinules, green
above, arachnoid-hairy, grayish-tomentose beneath, with prominent veins;
apical leaves lanceolate, entire, prickly-toothed. Heads 4-7 cm wide,
smaller on lateral branches. Capitula cuneate below, about 3 cm long;
bristles surrounding. capitulum unequal, white, somewhat flat, half as
long as involucre. Involucre of 17—18 glabrous bracts; outer bracts about
13 mm long, carinate and almost bowl-shaped above, irregularly short-
ciliate along margin, brownish on outer side; middle bracts attenuate-
acuminate, about 2.5 cm long, with thin and long appressed cilia along
margin; inner bracts longer, ciliate and entirely free above. Corolla light
blue. Achenes up to 15 mm long, with yellowish hairs; pappus of
brownish bristles fused in basal half. Flowering VII; fruiting VIII.

Zone of mountain meadows and steppes.—Caucasus: Southern
Transcaucasia. Endemic. Described from the Vedi District of Armenian
SSR. Type in Erevan; cotype in Leningrad.

37. E. sevanensis Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR,
VIII (1950) 52; Bot. Mat. Gerb. Bot. Inst. Akad. Nauk ArmSSR, XVIII
(1957) 294.—Ic.: Mulk. op. cit. (1957) figure on page 293.

Perennial. Stem 1 m or more high, cylindrical, poorly branched
above, white-tomentose throughout, without glandular hairs, leafy almost
up to tip. Lower leaves lanceolate, up to 50 cm long and 25 cm wide,
bipinnatisect into oblong-lanceolate lobes, with small lanceolate teeth
in their axils, sinuses between lobes wide, almost rectangular, lobes and
teeth terminating in yellowish, up to 8 mm long spines, petioles of
lower leaves long, broadened at base, spinulose; middle cauline leaves
sessile, oblong-ovate, pinnatifid into lanceolate or deltoid lobes; upper
leaves below heads sometimes entire, large prickly-toothed, with teeth

39

ge)

terminating in up to 15 mm long spines; leaves spinulose along margin,
slightly arachnoid-hairy above, without glandular hairs, grayish-tomentose
beneath, with prominent veins. Heads 5-6 cm wide. Capitula 20—25
mm long, on short peduncles, cuneate below; bristles surrounding
capitulum few, unequal, brownish at base, not more than one-third as
long as involucre. Involucre of 12-16 free bracts; outer bracts exceeding
adjacent bristles, rhomboidal and spatulate-convex above, cristate-ciliate
and glabrous; middle bracts lanceolate, attenuate-acuminate, with
appressed long cilia along margin, arachnoid-hairy on inner side above;
inner bracts shorter, long-ciliate above. Achenes covered with yellowish
bristles, turning dark; pappus bristles unequal, fused at base. Flowering
VII-VIII.

Steppes of middle mountain zone.—Caucasus: Southern
Transcaucasia (Sevan Lake Basin). Endemic. Described from the
Aregyuni District. Type in Erevan.

38. E. karabachensis Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR,
VIII (1950) 50; Dokl. Akad. Nauk ArmSSR, XII, 1 (1950) 27—Ie.:
Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR, Figs. 8, 9.

Perennial. Stem up to 60 cm high, cylindrical, branched above,
white-tomentose, without glandular hairs. Leaves elliptical, pinnatisect,
with yellowish midrib up to 5 mm wide, leaf lobes distant, linear-
lanceolate, prickly-toothed, terminating in spine; lower leaves up to 45
cm long and about 20 cm wide, lobes about 10 cm long and | cm wide,
spinulose along margin, separated by very wide sinuses; middle cauline
leaves much smaller, with linear lobes; apical leaves entire, up to 1 cm
long, green above, weakly arachnoid-hairy, without glandular hairs,
white-tomentose beneath. Heads 4—S cm wide. Capitula 20-25 mm long,
with cuneate base; bristles surrounding them white, numerous, equal to
outer involucral bracts. Involucre of 14-16 bracts; outer bracts four,
spatulately broadened above, with fewer cilia along margin; bracts of
next row rhomboidal above, dome-shaped, cristate-ciliate along margin;
middle bracts narrowed from middle, pointed, appressed long-ciliate;
inner bracts fimbriate-ciliate above, puberulent on outer surface, connate
at isolated points at base; corolla pale. Achenes covered with yellow
bristles; pappus bristles unequal, fused over half their length. Flowering
VU; fruiting VIII.

Caucasus: Eastern Transcaucasia (Nagorno-Karabakh Autonomous
Region). Endemic. Described from the regions between Khanbai and
Stepanokert. Type in Erevan.

Section 4. Pleiacme Bornm. in Beih. bot. Centralbl. XXXVI, 2
(1918) 214.—All involucral bracts free; outer and middle bracts acicular
along margin in upper pointed part appearing almost like branched horns.

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40

Besides our species, E. cervicornis Bornm., described from southern
Iran, also belongs to this section, and is the type species of this section.

39. E. tournefortii Ldb. ex Trautv. Diss. Echin. (1833) 21; DC.
Prodr. VI, 525; Ldb. Fl. Ross. II, 656; Jaub. and Sp. Ill. pl. or. 427;
Boiss. Fl. or. HI, 438; Iljin in Grossh. Fl. Kavk. IV, 159; Mulk. in Tr.
Bot. Inst. Akad. Nauk ArmSSR, VIII, 67.—Ic.: Jaub. and Sp. op. cit.,
tab. 427; Trautv. op. cit., Fig. 7.

Perennial. Stem 40-70 cm high, 5—15 cm thick, sulcate, arachnoid-
hairy, glandular below. Lower cauline leaves 30-40 cm long up to 20-
25 om wide, lanceolate, pinnatisect into lanceolate lobes in turn divided
into 0.5—1 cm wide lobules, prickly-toothed with teeth terminating in
spines up to | cm long; teeth and lobules of leaves often curved, giving
roundish shape to sinuses, latter occupying about half leaf area, leaf
axes narrow, almost winged, with 4—6 mm long spinules along margin,
lower lobes near petiole reduced into spines; leaves smaller and less
incised upward, up to lanceolate, long prickly-toothed, with spines up
to 2 cm long, weakly arachnoid-hairy above, scabrous and short-
glandular, scarcely recurved, white-tomentose beneath, with prominent
midrib and scattered long-stalked brown glandular hairs along veins.
Heads 6-7 cm wide. Capitula 3—3.5 cm long; bristles surrounding
them somewhat flat, unequal, yellowish, slightly longer than one-third
involucral length. Involucral bracts 15-17, membranous, scabrous and
glandular; outer bracts up to 2 cm long, broader in middle, spinescent-
acuminate, with divergent spinules along margin; middle and inner bracts
up to 3 cm long, attenuate-acuminate, scabrous, with fewer glandular
hairs and somewhat divergent spinules along margin. Achenes about 10
cm long, with long yellowish hairs; pappus pentagonal, bristles barbed,
fused at base. Flowering VIII-IX.

In the upper belt of steppe zone at 2,000-2,500 m.—Caucasia:
Southern Transcaucasia (Idzhevan and Akhuryan in Armenia). General
distribution: Eastern Anatolia, Armenia-Kurdistan, western Iran.
Described on the basis of cultivated specimens grown from seeds
collected in the Ararat. Type in Leningrad.

Section 5. Echinops—Sphaerocephalus Bge. in Bull. Acad. Sc.
Pétersb. VI (1863) 401, 409; Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR,
VIII (1950) 35.—Ritro Boiss. Fl. or. III (1875) 425 p. p. non Bge.—
Involucral bracts 16-18; outer bracts usually glandular; leaves
comparatively thin; plants of forest edges and thickets of shrubs.

Type of section: E. sphaerocephalus L.

Series 1. Sphaerocephali Mulk. op. cit. 41.—Leaves thicker, densely

41

glandular above. Outer involucral bracts long-acuminate, with numerous
stalked glandular hairs.

40. E. sphaerocephalus L. Sp. pl. (1753) 814; MB. FI. taur.- cauc.
Ill, 597; Trautv. Diss. Echin. 16; DC. Prodr. VI, 524; Ldb. FI. alt. IV,
45; Ej. Fl. Ross. II, 655; Schmalh. Fl. II, 91; Grossh. Fl. Kavk. IV, 157;
Kryl. Fl. Zap. Sib. XI, 2867—E. maximus Sievers ex Pall. Neue Nord.
Beitr. III (1796) 323.—E. cirsiifolius C. Koch in Linnaea, XXIV (1851)
379.—Ic.: Trautv. op. cit., f. 4; Majevski, Fl. edition 8, 594; Hegi, Ill.
Fl. VI, 2 tab. 269.—Exs.: GRF No. 1479; Fl. Ital. No. 184; Fl. Styr.
exs. No. 46, 1272; Fl. exs. reip. Boh.-Slov. No. 960.

Perennial. Stem about 1 m or higher, solitary, branched above,
ribbed, dirty brown due to numerous brownish glandular hairs, besides
with appressed white hairs above hence appearing white arachnoid-hairy,
leafy to tip. Leaves, except lowermost, sessile, amplexicaul, 15-25 cm
long and 5-10 cm wide, green and scabrous above due to glandular
hairs, grayish-tomentose beneath; lamina deeply incised into acute lobes,
ovate to almost deltoid, spiny or prickly-toothed along margin; upper
leaves small, usually not spatulate, prickly-toothed. Heads solitary
terminal on branches, 4—5.5 cm wide, common involucre of fewer bristles
connate at base. Capitula about 2 cm long; bristles surrounding them
grayish, much shorter than outer involucral bracts. Outer bracts oblong-
rhomboidal, attenuate above into cusp sometimes curved, margin ciliate,
glandular on outer side; inner bracts longest, fimbriate-ciliate above,
greenish below. Corolla pale blue or white, tube smooth. Anthers blue.
Achenes up to 10 mm long, turbinate, truncate above; pappus bristles
free in upper one-third. Flowering VI; fruiting VII.

Steppe meadows, shrubby growths, and along forest edges.—
European Part: Ladoga-I|men (vicinity of Leningrad, introduced), Upper
Dnieper (south), Upper Volga (south), Middle Dnieper, Volga-Don,
Trans-Volga Region, Volga-Kama (south), Bessarabia, Black Sea Region,
Crimea, Lower Don, Lower Volga; Caucasus: In all regions except high

41 mountains; Western Siberia: Upper Tobol (south), Irtysh (south), Altai;
Eastern Siberia: Angara-Sayans (only western in the Minusinsk District);
Soviet Central Asia: Aralo-Caspian (north), Balkhash Region (north),
Dzhungaria-Tarbagatai. General distribution: Southern and Central
Europe, Balkans-Asia Minor. Described from Italy. Type in London.

41. E. erevanensis Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR,
VIII (1950) 43.

Perennial. Stems 1-2 m high, 0.7—2 cm thick at base, usually five
or six, highly branched above and therefore single plant with dozens of
heads, with numerous, brown glandular hairs along stem, somewhat

42

grayish-tomentose above. Leaves thin, ovate, glandular pubescent, pale
green beneath, mildly grayish arachnoid-hairy, slightly prickly and ciliolate
along margin; basal leaves about 40 cm long, long-petiolate, pinnatisect
into deltoid-ovate distant lobes; cauline leaves semiamplexicaul, auriculate;
upper cauline leaves sessile, lanceolate, undivided, toothed. Heads
numerous; bristles surrounding them numerous, somewhat flat, unequal,
half as long as involucre. Involucral bracts 15—18; outer bracts weakly
acuminate, ciliate, glandular on outer side; middle bracts longer; inner
somewhat connate below, obtuse, shorter than adjacent middle bracts.
Corolla slightly swollen in throat; stamen tube sometimes rupturing at
end of flowering. Achenes densely covered with long hairs; pappus
bristles barbed, fused over three-fourths their length. Flowering VII;
fruiting IX.

In ruderal habitats—Caucasus: Southern Transcaucasia. Endemic.
Described from’ the vicinity of village of Garni. Type in Erevan.

Note. An enigmatic plant, known from two localities in Armenia,
distinguished by vigorous growth (may be due to favorable conditions
of the habitats) and shorter pubescence. | am not sure about the racial
independence of this globe-thistle from E. sphaerocephalus L.

42. E. dagestanicus Iljin in Grossh. Fl. Kavk. IV (1934) 157; Bot.
Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, VII, 1, 63; Mulk. in Tr. Bot.
Inst. Akad. Nauk ArmSSR, VIII, 45.

Perennial. Stem 60-100 cm high, branched above and almost
white-tomentose, ribbed almost throughout, white arachnoid-hairy
predominantly along ribs, densely glandular-hairy on brownish bristles
along furrows, appearing brownish or reddish. Leaves lanceolate,
pinnatifid, matte green above, scabrous, setose-glandular, sparsely grayish
arachnoid-hairy beneath, besides glandular lamina incised into deltoid
lobes, sharply short toothed and terminating in spinule, smaller upward;
upper leaves almost undivided below heads, sharply toothed, 3-5 cm
long. Heads solitary terminal on branches, four to six on single stem,

42 4-5 cm wide. Capitula about 20 cm long, bristles surrounding them
unequal, yellowish, not more than one-third as long as involucre.
Involucral bracts 18-20; outer bracts 7-8 mm long, broadest in upper
third, with cuneate brownish base, hairy on outer surface, acuminate,
divergent ciliate along margin, middle bracts carinate, up to 15 mm
long, attenuate-acuminate, puberulent along outer surface, ciliate; inner
bracts shorter than middle, ciliate-fimbriate above, scarcely fused in lower
half, greenish brown below on inner side. Corolla pale blue. Achenes
with yellowish hairs; pappus bristles fused over basal half. Flowering
VI; fruiting VII.

Shrubby thickets and forest edges in middle mountain zone.—

43

43

Caucasus: Dagestan, eastern Transcaucasia. Endemic. Described from
the Shemakhi District. Type in Leningrad.

43. E. viridifolius Iljin in Bot. Mat. Gerb. Bot. Sada, IV (1923)
103; Iljin in Grossh. Fl. Kavk. IV, 158; Mulk. in Tr. Bot. Inst. Akad.
Nauk ArmSSR, VIII, 47.

Perennial. Stem high, branched above, sulcate, brownish due to
numerous stalked glandular hairs, almost white-tomentose above. Upper
leaves sessile at base of inflorescence, auriculate, lanceolate, pinnatifid
into deltoid prickly-toothed lobes or simply prickly-toothed, 3-8 cm
long, 1-3 cm wide, grayish-green above, densely glandular-hairy, slightly
lighter colored beneath, sparsely arachnoid-hairy, and glandular along
veins. Heads 5—6 cm wide. Capitula 20-23 mm long; bristles surrounding
them about 7 mm long, unequal, somewhat flat, white. Involucre of 20—
23 thin, scabrous lustrous bracts, only outer ones puberulent but without
glandular hairs; outer bracts rhomboidal above, sparsely ciliate; middle
bracts lanceolate, acuminate, appressed-ciliate along margin; inner bracts
sometimes fused at some points, acuminate and incised above. Corolla
light blue. Achenes long-hairy; pappus of yellowish bristles fused up to
half their length. Flowering VII.

Steppe slopes.—Caucasus: Ciscaucasia (Mt. Mashuk near
Pyatigorsk). Endemic. Described from vicinity of Pyatigorsk. Type in
Leningrad.

Note. The type specimen of this species is the upper portion of a
single plant. It differs from E. sphaerocephalus L. in having eglandular
outer involucral bracts and leaves less pubescent below (only apical
leaves in the authentic specimen). The herbarium of the Academy of
Sciences of the USSR has typical specimens of E. sphaerocephalus L.,
collected at Mashuk. I am not sure that E. viridifolius Tin is an
independent species, and not just a weakly pubescent form of the
Linnaean species. Mulkijanyan saw specimens in the Baku Herbarium
collected from Anapa.

Series 2. Galatici Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR,
VIII (1950) 36.—Leaves comparatively thin, with few glandular hairs.
Outer involucral bracts short acuminate, with numerous glandular hairs.

44. E. galaticus Freyn in Bull. Herb. Boiss. II (1895) 355; Iljin in
Grossh. Fl. Kavk. IV, 157; Mulk. in Tr. Bot. Inst. Akad. Nauk ArmSSR,
VU, 36.

Perennial. Stem 40-60 cm ‘high, weakly sulcate, branched above,
more or less grayish-woolly throughout, with stalked glandular hairs
along furrows below. Leaves thin, brownish-green above, densely

44

44

glandular-hairy, light gray beneath, almost tomentose; lower leaves 1.5
cm long, petiolate, lyrate, lower lobes small, widely spaced, terminal
lobe largest, elliptical, lateral lobes ovate, short prickly-toothed along
margin; leaves smaller upward, amplexicaul-auriculate, oblong or
pinnatifid above middle into broad deltoid or ovate lobes, sparsely ciliate-
prickly along margin. Heads 3-4 cm wide, bluish. Capitula about 20
mm long; bristles surrounding them almost one-third as long as
capitulum. Involucral bracts 18-20, free, light green, darker in lower
half; outer bracts rhomboidal or spatulate, cristate-fimbriate; middle bracts
lanceolate, attenuate-spiescent, carinate, ciliate-fimbriate from middle,
finely scabrous, with isolated small glandular hairs; inner bracts slightly
shorter, obtuse, slightly incised or almost prickly. Achenes with numerous
yellowish hairs; pappus bristles barbed; free in upper half. Flowering
VII; fruiting VIII.

Shrubby slopes and forest edges—Caucasus: Ciscaucasia (western
part of the main range), western Transcaucasia. General distribution:
Asia Minor. Described from Amasya in eastern Anatolia. Type in Brno;
cotype in Leningrad.

45. E. colchicus Sosn. ex Kem.-Nat. in Bull. Mus. Géorgie, IV
(1928) 138, nomen; Sosn. in Zhurn. Russk. Bot. Obshch. XIV, 87; Mulk.
in Tr. Bot. Inst. Akad. Nauk ArmSSR, VIII, 37.—£. bannaticus ssp.
quercifolius Freyn in Bull. Herb. Boiss. III (1895) 356.

Perennial. Stem 60—100 cm high, relatively slender, angular, loosely
white-tomentose and glandular below, slightly branched above; leaves
almost lyrate, crenate-pinnatipartite, dark green above and glabrous or
slightly arachnoid-hairy, loosely white-tomentose beneath; lower leaves
15-20 cm long, with broad deltoid lobes, slightly crenate, short-spiny at
tip and along margins, smaller and sparse toward lower part; middle
cauline leaves deeply amplexicaul; upper leaves sessile, oblanceolate,
toothed. Heads 3—4.5 cm wide, light bluish; capitula 15—20 mm long,
bristles surrounding them up to one-third their length, fewer and unequal.
Involucral bracts 20—23, free; outer bracts lanceolate, acuminate, carinate,
brownish and hairy on outer surface, slightly longer than bristles; middle
bracts oblong-lanceolate, attenuate-acuminate, scabrous on outside, long-
ciliate along margin; inner bracts slightly shorter, incised-ciliate or
fimbriate above, dark colored below on inner side. Corolla pale blue.
Achenes pubescent; pappus of brownish, free, scabrous bristles fused
only in upper one-fourth. Flowering VII; fruiting VIII.

Edges and swales of mountain-forest zone.—Caucasus: Western
Transcaucasia. General distribution: Eastern Anatolia. Described from
Kutaisi District. Type in Tbilisi.

Note. The absence of the original specimens of Freyn and shortage
of material from western Transcaucasia with me compels me to doubt

45

45

the independent status of this race. The fact that it apparently does not
have an independent distribution area and the plants that should have
been included under it appear to be simply more mesophilous plants,
strengthens my doubt.

46. E. ossicus C. Koch in Linnaea, XXIV (1851) 380; Boiss. FI. or.
suppl. (1888) 304; Iljin in Grossh. Fl. Kavk. IV, 159; Mulk. in Tr. Bot.
Inst. Akad. Nauk ArmSSR, VIII, 39, Fig. 6a.

Perennial. Stem 60-100 cm high, slender, sulcate; loosely white-
tomentose, glandular below, slightly branched above. Lower leaves
oblanceolate, somewhat lyrate, 12-20 cm long, 4-8 cm wide, deeply
crenate-toothed to pinnatilobate with deltoid short-prickly lobes and large
terminal lobe; cauline leaves small, less deeply incised, amplexicaul;
uppermost leaves lanceolate, toothed, leaves dark green above, slightly
arachnoid-hairy, hispid, white-tomentose beneath. Heads 3-4 cm wide.
Capitula 15-18 mm long; bristles surrounding them white, somewhat
flat, slightly less than half length of involucre. Involucral bracts 18-20,
free, scabrous, somewhat pubescent on outer surface; outer bracts
spatulate, long-ciliate, up to 7-8 mm long; middle bracts attenuate-
acuminate, carinate, appressed ciliate; inner bracts slightly longer,
somewhat obtuse, fimbriate-ciliate. Corolla light blue. Pappus bristles
fused almost to tip. Fowering VII; fruiting VIII.

Forest edges and grasslands of mountain-forest zone.—Caucasia:
Western Transcaucasia, eastern Transcaucasia (South Ossetia). Endemic.
Described from South Ossetia. Type lost.

Note. Mulkijanyan (op. cit.) traveled in 1948 along the path followed
by C. Koch and found a globe-thistle in South Ossetia that in his opinion
corresponded to the description of Koch. Taking into account the
evidence of Mulkijanyan, who studied the plants in nature, | retain this
fairly doubtful species among the globe-thistles of the Caucasus.

Series 3. Latifolii Bobr—Leaves with wide lobes and small sinuses.
Outer involucral bracts without glandular hairs, glabrous in East Siberian
species, and pubescent in Mediterranean species. Plants of Eastern Siberia
and east of Soviet Central Asia.

47. E. latifolius Tausch in Flora, XI (1828) 486; Kitag. in Rep.
first Sc. exp. Manch. IV, 2, 119.—£E. dahuricus Fisch. Cat. Hort. Gorenk.
ed. 2 (1812) 37, nom. nud.; DC. Prodr. VI, 523; Turcz. Fl. baic.-dahur.
II, 28; Popov, Fl. Sredn. Sib. I, 750.—E. dahuricus a. latilobus DC.
Prodr. VI (1837) 523.—E. gmelini auct. non Turcz. 1832, Trautv. Diss.
Echin. 14; Ldb. Fl. alt. IV (1833) 45; Ej. Fl. Ross. II, 653; Kom. FI.
Manchzh. III, 713 p. p.; Kom. and Alis. Opred. Rast. Dalnevost, Kr. II,
158.—Sphaerocephalus dauricus O. Ktze. ex Kom. Fl. Mansh. III (1907)

46

46

713.—E. caule ‘subunifloro. . .Gmel. Fl. Sib. I (1749) 100.—Ie.: Trautv.
opacitydivB:

Perennial. Stems 30-60 cm high, simple or weakly branched, slightly
arachnoid-hairy below, almost tomentose above. Leaves bipinnatipartite
or compound, weakly arachnoid-hairy above, less often subglabrous,
almost tomentose below, prickly-toothed or prickly-lobed along margin;
basal leaves on thick 10-15 cm long petioles, pinnatipartite; lower cauline
leaves amplexicaul, pinnatipartite, upper ovate, pinnatifid; sometimes
leaves with finely divided lobes (var. B. Trautv. I.c.). Heads 4-6 cm
wide, bluish; capitula up to 2 cm long, almost ovoid with cuneate base,
broadest in middle; bristles surrounding them dense, scabrous, slightly
longer than involucre. Involucral bracts glabrous; outer bracts spatulate,
acuminate, long-ciliate above; middle bracts oblong, broadest in middle,
ciliate above and attenuate into recurved tip. Corolla longer than
capitulum. Ovary densely hairy. Pappus setose, bristles linear, fused at
base, free and weakly ciliate above. Flowering V; fruiting VI.

Steppe and stony slopes.—Eastern Siberia: Anagara-Sayans
(southeast), Dauria (south). General distribution: Mongolia. Described
from Baikal. Type lost.

48. E. dissectus Kitag. in Rep. first Sc. exp. Manch. IV, 2 (1935)
118.—E. gmelini auct. p. p. non Turcz. 1832; Kom. Fl. Manchzh. II],
TASS pep:

Perennial. Root cylindrical, root collar covered with brown of
previous year’s leaves. Stem solitary, simple, less often with fewer
branches above, almost glabrous below, but with isolated brownish
bristles, densely white arachnoid-hairy or woolly above. Basal leaves
withering before flowering; cauline leaves smaller upward, somewhat
amplexicaul, broadly oblong, pinnatifid to deeply pinnatipartite, usually
glabrous and dark green above, densely, white arachnoid-hairy to woolly
beneath but sparsely along veins, up to 20 cm long and 8 cm wide,
with lobes deeply pinnatisect, prickly along margin and pointed and
prickly at tip. Heads 4-6 cm wide, slightly shorter than outer involucral
bracts, somewhat broadened above, ciliate along margin. Outer involucral
bracts spatulate, scaly, sparsely ciliate along margin, broadened upward;
middle bracts rhomboid to linear-subulate, setose-ciliate along margin,
coriaceous and gradually subulately acuminate above, carinate along outer
surface; inner bracts almost linear, papery, almost cristate-ciliate above,
with longer terminal cilia. Florets pale blue, slightly exserted from
involucral bracts; ovary hairy. Pappus bristles 1 mm long, free above
middle. Anthers up to 5 mm long, lower appendages scarcely 0.5 mm
long, deltoid; style very deeply furcate. Flowering VII; fruiting VIII.

Open mixed forests and grassy southern slopes of hills—Far East:

47

47

Zeya-Bureya (south), Ussuri. Genera’ distribution: Northeastern China.
Described from “Manchuria.” Type in Tokyo.

49. E. tricholepis Schrenk, Emun. pl. I (1841) 47; Ldb. Fl. Ross.
Il, 655; Kryl.. Fl. Zap. Sib. XI, 2868.—E. exaltatus auct. non Schrad.;
O. and B. Fedtsch. Perech. Rast. Turkm. IV, 226. _

Perennial. Stem 1—1.5 m high, branched, ribbed, reddish or greenish,
with short glandular hairs and arachnoid-hairy at tip. below head and
white-tomentose along ribs. Lower leaves petiolate, upper sessile,
amplexicaul, 10-20 cm long and 5-12 cm wide, spatulate, to deeply
pinnatisect; almost glabrous or slightly arachnoid-hairy and green above,
finely tomentose, glandular, and grayish beneath; leaf lobes ovate to
triangular, with teeth along margins terminating in spinules. Heads 4-6
cm wide, one or two terminal on branches, pale blue. Capitula 2—2.3
cm long; bristles surrounding them not longer than half length of
capitulum. Involucral bracts free; outer bracts oblong, rhomboidally
broadened from middle, acuminate, glabrous below, finely pubescent
above, with scabrous cilia along margin; inner bracts oblong-ovate,
fimbriate-ciliate, without terminal spinules. Corolla tube glandular; petals
white, 6-8 mm long. Anthers bluish, with ciliate lobes; pappus free
almost up to half.

Thickets of shrubs along river valleys.—Soviet Central Asia:
Balkhash Region (north), Dzhungaria-Tarbagatai. General distribution:
Dzhungaria-Kashgaria. Described from the foothills of the Dzhungarian
Alatau. Type in Leningrad.

50. E. abstersibilis Iljin in Bot. Mat. Gerb. Bot. Sada, III (1922)
Ve.

Perennial. Stem 50-70 cm high, with two or three branches, sulcate
below and white-tomentose in middle floccose, glabrescent at some
points, without glandular hairs. Lower cauline leaves 25-30 cm long,
7-8 cm wide with broad prickly base, petiolate, pinnatipartite into wideset
oblong-deltoid, prickly lobes, terminating in spinules, separated by broad
round sinuses, spinulose along margin; upper leaves pinnately lobed,
terminating in spines; cauline leaves sessile, semiamplexicaul, smaller
upward and less deeply incised, green, finely arachnoid-hairy, without
glandular hairs above; grayish tomentose, with prominent veins beneath.
Heads 2.5-3 cm wide. Capitula 16-18 mm long; bristles surrounding
them yellowish, somewhat flat, about 6 mm long. Involucral bracts 20,
densely puberulent, visible under high maginfication; outer bracts (six)
membranous, rhomboidal above, about 7 mm long, ciliate; middle bracts
(nine) about 15 mm long, subulate, carinate, slightly divergent above,
long-ciliate; inner bracts free, slightly fused only at base, ciliate or

48

48

fimbriate along margin above. Corolla deeply parted, short-hairy below
throat. Achenes covered with yellowish long bristles longer than pappus
of fused bristles. Flowering VIII.

Upper mountain zone, at altitude of 2000-3000 m.—Soviet Central
Asia: Pamiro-Alai (Central and Southern Tadzhikistan). Endemic.
Described from the Vakhsh-Kyzylsu interfluvium. Type in Leningrad.

Section 6. Terma Endl. Gen. pl. (1836) 467; Bge. in Bull. Akad.
Sc. Pétersb. VI (1863) 400, 410.—Pappus cup-shaped, divided only above.
Lectotype of section: E. exaltatus Schrad.

51. E. exaltatus Schrad, Hort. Goett. (1809) 15, tab. 9; Rchb. FI.
Germ. exc. 292; Trautv. Diss. Echin. 12; DC. Prodr. VI, 523; Rchb. Ic.
Fl. Germ. XV, 2, tab. 735——E. commutatus Juratzka in Verh. Zool.-
Bot. Ges. Wien, VIII (1858) 17; Hegi, Ill. Fl. VI, 2, 811; Szaf. Kulcz.
Pavl. B. Rosl. Polskie, 708; Vizn. Rosl. URSR, 559.—E. strictus Fisch.
ex Sims. Bot. Mag. (1824) tab. 2457.—Ic.: Schrad. |.c.: Trautv. op. cit.
2; Bot. Mag. I.c.; Rchb. Ic. Fl. Germ. XV, tab. 735; Hegi, op. cit., tab.
811.—Exs.: Fl. exs. reip. Boh.-Slov. No. 1185; Fl. austro-hung. No.
3437.

Perennial. Stem 50—150 cm high, usually profusely branched above;
branches terminating in heads, less often stem simple, sulcate-ribbed,
white woolly-tomentose sparsely below and densely above, brownish or
pale. Lower leaves petiolate; cauline leaves few, distant, with prickly
amplexicaul base, 20-30 cm long and up to 15 cm wide, deeply
pinnatipartite, with oblong to deltoid lobes and prickly along margin
terminating in spine, dark green and sparsely brown-strigose above, white-
tomentose beneath, sometimes grayish. Head globose, on long peduncle,
4—6 cm wide. Capitula numerous, up to 2 cm long; bristles surrounding
them unequal but one-third to one-fourth as long as capitulum, colorless,
somewhat flat above. Involucral bracts gradually enlarged and more acute
from outer to inner, with slightly recurved tip, without glandular hairs,
margins ciliate; innermost involucral bracts, however, shorter, scarcely
one-third as long as middle bracts, and incised at tip; all bracts silvery-
gray to greenish. Corolla white; stamen tube grayish. Achene about 7
mm long, densely silky with yellowish hairs; pappus cup-shaped, divided
above. Flowering VI; fruiting VII.

Forest edges and scrub, along river and stream banks——European
Part: Upper Dniester (Transcarpathian Province). General distribution:
Southern and Central Europe. Described from cultivated specimens grown
in the G6ttingen Garden, wrongly attributed to Siberia as the place of
origin. Cotype in Leningrad.

49

Aan yi Re

si
ay “i

~%

a} my Rat} N
Sis

Plate III.

49

1, 2—Echinops integrifolius Kar. and Kir., lower and upper part of plants; 3—E. humilis MB.

50

50

Note. Because Siberia was erroneously cited as the native locality
in the description of this species, further errors were also committed,
although De Candolle expressed doubt about the correctness of this report.

In connection with this report, the plants collected by Carrelin in
Ayagyz (Enum. 1841, No. 469) were identified as E. exaltatus Schrad.
On the basis of the specimens of Carrelin, this species was included in
our “Flora.” This citation was repeated by B.A. Fedtschenko (Perech.
Rast. Turk. [List of Plants of Turkmenia], IV, 226), while expressing, it
is true, some doubt about its correctness. Assuming that E. exaltatus
Schrad. was described “from Siberia,” Juratzka (1.c.) described the
European plant on the basis of specimens originating from the environs
of Trieste as a closely related plant replacing the European species. The
study of the original descriptions and cited figures of E. exaltatus Schrad.,
as well as the plants of Schrader raised by him in the Gottingen Garden
(type specimen!), leads to the conclusion that E. exaltatus Schrad. is
not from Siberia but is a West European plant, and the name proposed
by Juratzka—E. commutatus Juratzka—is a simple synonym. As regards
the plant collected by Carrelin (Enum. 1841, No. 469), it was wrongly
identified, and is actually E. tricholepis Schrenk. Thus, E. exaltatus
Schrad. is excluded from the flora of Siberia, more precisely, from the
flora of Soviet Central Asia.

Section 7. Chamaechinops Bge. in Bull. Acad. St. Pétersb. VI (1863)
411; Keller in Tr. Obshch. Estestv. Kaz. Univ. XLIV, 5 (1912) 96.—
Pappus bristles subulate, spreading above, toothed, not barbed. Plants
from the northeast of Soviet Central Asia and mountainous Siberia—
from Tien Shan to Sayans.

Type of section: E. humilis MB.

52. E. integrifolius Kar. and Kir. in Bull. Soc. Nat. Mosc. XIV
(1841) 446; Ldb. FI. Ross. II, 656; Kryl. Fl. Zap. Sib. XI, 2872.

Perennial. Stem 30-80 cm high, usually simple, with numerous
remains of previous year’s leaves at base, densely leafy to tip, with
short-stalked glandular hairs, white-tomentose above. Leaves 5-10 cm
long and up to 1.5 cm wide, numerous, oblong or lanceolate, with broad
amplexicaul base, smaller upward and less dense, undivided, only lower
cauline and basal leaves sometimes with one to three teeth on each
side; upper leaves with two or three spines along margin; leaves green
above and densely covered with short-glandular hairs, white-tomentose
beneath. Heads solitary, whitish, 2-4 cm wide; bristles surrounding
capitula not longer than outer involucral bracts; latter linear, rhomboidal
above, long-ciliate, glandular above on inner side, acuminate; middle
bracts densely pubescent with spinescent tip and long cilia along margin;

51

ay

inner bracts oblong, spiny, brownish on outer surface. Corolla white,
tube glabrous. Pappus bristles unequal, free above over not more than
one-third length. Flowering VII; fruiting VIII. (Plate III, Figs. 1, 2).

Sandy and rubbly valleys and slopes of foothills—Western Siberia:
Altai (extreme south—Naryim Range, Kurchum Mountains, Saur); Soviet
Central Asia: Balkhash Region (Zaisan Depression, north). General
distribution: Dzhungaria-Kashgaria (extreme north of Chinese Dzhungaria),
Mongolia (extreme west). Described from the Kurchum Mountains. Type
in Leningrad.

53. E. humilis MB. FI. taur.-cauc. III (1819) 598; Trautv. Diss.
Echin. 27; Ldb. Fl. alt. IV, 45; DC. Prodr. VI, 523; Ldb. FI. Ross. II,
653; Kryl. Fl. Zap. Sib. XI, 2871.—£E. davuricus Trevir. Ind. Sem. h.
Wratisl. p. a. 1819 sec. Ldb. Fl. Ross—Ie.: Trautv. op. cit., f. 11.

Perennial. Stem 7—15, less often up to 30 cm (var. elatus Kryl.)
high, covered with remains of numerous leaves in lower part, solitary,
usually simple, sometimes branched and then with several heads, densely
white-tomentose throughout. Basal leaves numerous, short-petiolate,
rosulate, usually lyrate-pinnatilobate, sometimes almost undivided, 2—7
cm long and about 1 cm wide, white-tomentose, sparsely above and
more densely below, with ovate lobes, obtuse, less often spinescent;
cauline leaves pinnatifid, divergent, with oblong lobes, spinescent; upper
leaves prickly-toothed, more divergent and even recurved. Heads 3-4
cm wide, solitary, smaller on branched stems. Capitula 1.5—1.7 cm long;
bristles surrounding them almost as long as capitulum. Outer involucral
bracts lanceolate; inner bracts long-ciliate, exceeding outer bracts. Corolla
light blue, with smooth tube and light bluish anthers. Achenes long-
hairy; pappus bristles unequal, fused at base. Flowering VII; fruiting
VII. (Plate III, Fig. 3).

Gravelly slopes and valleys, often mildly alkaline —Western Siberia:
Altai (eastern Altai); Eastern Siberia: Angara-Sayans (Tuva Autonomous
Republic). Endemic. Described from Chu Valley. Type in Leningrad.

54. E. tschimganicus B. Fedtsch. in O. and B. Fedtsch. Perech.
Rast. Turk. IV (1911) 224.—Exs.: H.F.A.M. No. 40.

Perennial. Stems fewer, 10-30 cm high, white-tomentose at base,
densely covered with remains of previous year’s leaves, densely leafy
particularly in lower part, usually simple, with single head, stalked-
glandular hairy, finely white-tomentose above. Basal leaves short-
petiolate; cauline leaves sessile, all deeply pinnatipartite into linear,
entire, short-acuminate lobes; leaves oblong, green above, densely
stalked-glandular hairy, involute, 3-4 mm wide along midrib. Heads
about 3 cm wide, solitary. Capitula about 1.5 cm long; bristles

52

52

surrounding them somewhat flat above, almost as long as outer involucral
bracts. Outer bracts deltoid above, ciliate, pubescent on outer surface;
middle bracts linear-lanceolate, with whitish border, long ciliate above,
cuspidate, pubescent on outer surface; inner bracts lanceolate, with wide
white border, hairy on outer side, margin ciliate upward. Corolla light
blue, glandular-hairy on outer side below throat. Achenes white-
pubescent; pappus bristles white, fused at base. Flowering VI; fruiting
VU.

Gravelly slopes of middle mountain zone.—Soviet Central Asia:
Tien Shan (western Tien Shan—Chimgan, Karatau, Talass Alatau).
Endemic. Described from the Kumyshtag Valley. Type in Leningrad.

55. E. saissanicus (Keller) Bobr. comb. n.—E. gordjaginii saissanicus
Keller in Tr. Obshch. Estestv Kaz. Univ. XLIV, 5 (1912) 95.—E.
gordjaginii Keller ex Kryl. Fl. Zap. Sib. XI (1949) 2870.—Ie.: Keller,
op. cit.

Perennial. Root virgate, two or three-headed, sometimes bearing,
besides scape, basal rosettes of leaves of daughter plants. Stem 20-40
cm high, with remains of numerous old leaves, arachnoid-hairy at base,
short-glandular hairy above, erect, branched in upper part, sometimes
white arachnoid-hairy below solitary terminal heads. Basal leaves
numerous, petiolate; middle and upper leaves sessile, with broad spiny
base; all leaves lanceolate, convolute, pinnatisect, with narrow (f.
angustisecta Keller) or wider, deltoid and decurrent (f. latisecta Keller)
lobes, S—15 cm long; upper leaves 2—3 cm long; leaves green and densely
glandular hairy above, white-tomentose beneath, with prominent midrib,
bearing stalked glandular hairs; lobes of leaves distant, entire, lanceolate
to linear; terminating in spines. Head globose, 2-4 cm wide; bristles
surrounding them as long as outer involucral bracts. Outer bracts linear,
ciliate, rhomboidal and short-ciliate above; middle and inner bracts
lanceolate, subulate, short-ciliate above, scabrous and slightly pigmented
on outer surface. Corolla white, glabrous on outer side below throat.
Anther appendages very short, filiform. Pappus bristles subulate, toothed,
fused over half their length. Flowering VII.

Stony slopes and in crevices of rocks in semidesert.—Western
Siberia: Altai (extreme south); Soviet Central Asia: Balkhash Region
(Zaisan Depression). Endemic. Possibly, of course, also grows in
Dzhungaria-Kashgaria. Described from Kaldzhir. Type in Leningrad.

Note. In establishing E. gordjaginii Keller the author, contrary to
the rules of nomenclature, included under this name the species E.
tschimganicus B. Fedtsch., validly described earlier by B.A. Fedtschenko
as a subspecies or variety while distinguishing his own Zaisan plants at
this rank (8). Given that the description of B.A. Fedtschenko was fully

%

53

53

valid, B.A. Keller should have described the Zaisan plants either as a
separate species or under the species established by Fedtschenko. Keller
did neither; therefore, the name proposed by him is invalid and should
be rejected. I propose to name this Zaisan globe-thistle as E. saissanicus
(Keller) Bobr.

Section 8. Nanechinops Bge. in Bull. Acad. Sc. Pétersb. VI (1863)
401, 411.—Appendages of anthers subulate below, barbed above. Annuals
of sandy and halophytic semidesert of Central and Soviet Central Asia.

Type of section: E. gmelini Turcz.

56. E. gmelini Turcz. in Bull. Soc. Nat. Mosc. V (1832) 195; DC.
Prodr. VI, 527.—E. turczaninovii Ldb. ex Trautv. Diss. Echin. (1833)
28; Ldb. FI. Ross. II, 657; Iljin in Bot. Mat. Gerb. Bot. Sada, IV, 181;
Kryl. Fl. Zap. Sib. XJ, 2872.—Echinops foliis integris Gmel. Fl. Sib. II
(1749) 103.—Ie.: Gmel. I.c. tab. XIV, fig. 3; Trautv. op. cit. fig. 12.

Annual. Stem 15-30 cm high, simple or divaricately branched from
base, with numerous long-stalked glandular hairs. Leaves numerous,
undivided or slightly lobed, sessile, oblong, semiamplexicaul, 2—5 cm
long, 0.5—2 cm wide, with glabrous, acicular, up to 0.5 cm long spines
along margin; usually lower leaves somewhat tomentose above due to
long entangled hairs; middle and upper leaves green above and glandular-
hairy on both sides. Heads whitish, globose, up to 3 cm wide, solitary
terminal on branches; bristles surrounding them almost as long as
involucral bracts. Involucral bracts imbricate, oblong-linear; outer bracts
aristate; inner fimbriate above; all bracts plumose-ciliate, pubescent on
outer surface. Basal appendages of anther subulate, apical barbate. Pappus
bristles fused at base, spreading and subulately acuminate. Achenes
densely hairy. Flowering VI; fruiting VIII.

Sandy deserts of Central Asia, in Mongolia and China; Western
Siberia: Altai (Naryim Range, near Altaiskaya sta.). General distribution:
Mongolia, China. Described from northwestern Mongolia. Type in
Leningrad.

57. E. nanus Bge. in Bull. Acad. Sc. Pétersb. VI (1863) 411; O.
and B. Fedtsch. Perech. Rast. Turkm. IV, 226.

Annual, less often biennial. Stem 20-30 (50) cm high, solitary, white-
tomentose, often profusely branched; branches divaricate, leafy, with
solitary terminal heads. Basal and lower cauline leaves short-petiolate;
upper leaves sessile, slightly decurrent; leaves lanceolate, pinnatipartite
into acuminate lobes, or undivided, toothed; upper leaves crenate-toothed,
undivided; leaves densely arachnoid-hairy above, grayish, glandular,
grayish-tomentose beneath. Heads 2.5-3 cm wide, pale, somewhat light

54

bluish. Capitula about 1.5 cm long; bristles surrounding them white
scabrous, not more than half as long as capitulum: Involucral bracts 22,
free; outer bracts 6-8 mm long, not longer than bristles surrounding
capitulum, pubescent on outer side and villous at base, ciliate along
margin; middle bracts 12—13 mm long, lanceolate, cuspidate, ciliate along
margin, pubescent on outer surface; inner bracts slightly shorter than
middle bracts, strigose, hairy below. Corolla light bluish, about 1 cm
long, tube glandular below throat. Achenes densely glandular-hairy;
pappus bristles short and scabrous, fused at base. Flowering VI; fruiting
VII.

Stony-sandy semidesert in plains and on gravelly slopes of
foothills—Western Siberia: Irtysh (extreme south—Karkarala Region);
Soviet Central Asia: Balkhash Region, Dzhungaria-Tarbagatai (foothills
of the Dzhungarian Alatau), Tien Shan, Syr-Darya (Fergana), Pamiro-
Alai (Zeravshan and western Pamir). General distribution: Dzhungaria-
Kashgaria. Described from Balkhash. Type in Leningrad.

58. E. knorringianus Ijin in Bot. Mat. Gerb. Glavn. Bot. Sada, III
(1922) 173.

Annual. Stem 10-40 cm high, solitary, erect, sometimes with one
or two divergent but not divaricate branches, densely covered with long,
yellowish, stalked glandular hairs, sometimes sparsely tomentose-
arachnoid-hairy above. Leaves green or yellowish green above with
numerous stalked glandular hairs, white-tomentose beneath, with very
sparse yellowish glandular hairs along midrib; basal leaves petiolate,
largest leaves 4-10-cm long, lanceolate to ovate, crenate to prickly-
toothed along margin, less often pinnatilobate; cauline leaves smaller
upward, lanceolate, crenate-toothed, terminating in spines. Heads usually
solitary terminal, sometimes two or three on branches, 2-3 cm wide.
Capitula about 1.5 cm long; bristles surrounding them about | cm long,
white, scabrous. Involucral bracts 20, free; outer bracts not longer than
bristles, long-hairy on inner side, scabrous on outer side, strigose above;
middle bracts about 1.5 cm long, cuspidate, divergent hairy on outer
surface, coarsely ciliate along margin; inner bracts 12-13 mm long,
lanceolate, obtuse, short-strigose along margin, long-hairy on outer side.
Corolla whitish, about 1 cm long, with hairy limb. Achenes densely
hispid; pappus bristles yellowish, scabrous, fused below. Flowering VI;
fruiting VII.

Wormwood-saltwort semidesert, on gravelly and rubbly slopes of
foothills ——Soviet Central Asia: Syr-Darya (Fergana Valley); Pamiro-Alai
(Upper Zeravshan, Madm). Described from Shaidan in the vicinity of
Namangan. Endemic. Type in Leningrad.

54

55

GENUS 1568. Acantholepis Less.':?

Less. in Linnaea, VI (1831) 88; Mulk. in Izv. Akad. Nauk ArmSSR,
II, 9 (1950) 839.

Capitula single-flowered, numerous, aggregated into dense hemi-
spherical or subsequently almost globose heads, surrounded by well
devgloped many-rowed involucre of usually large bracts, inner bracts
smaller and compressed, outer bracts squarrose, leafy, much longer than
head. Individual involucres (i.e. involucres of individual single-flowered
capitulum) many-rowed, imbricate, oblong; involucral bracts free; outer
bracts forming so-called brush, setaceous, inner bracts longer, <ristate,
long ciliate. Common receptacle very small, spherical-conical, irregularly
tuberculate, glabrous, smooth. Flowers uniform: all bisexual, tubular,
regular; corolla tube short and cylindrical with 5-lobed limb, lobes linear.
Anthers sagittate at base, with short ciliate-fimbriate appendages. Style
with rather thick, short, subsequently divergent branches, surrounded by
ring of hairs at base. Achene oblong-obovoid, irregularly globose-4-
angled, densely pubescent. Pappus one-rowed, coronate, very short, finely
dentate.

Annual. Usually small, white-pubescent plant with alternate, entire,
and linear-lanceolate leaves. |

A monotypic genus.

1. A orientalis Less. in Linnaea, VI (1831) 88; DC. Prodr. VI, 527;
Ldb. FI. Ross. II, 2, 657; Boiss. Fl. or. III, 442; O. and B. Fedtsch.
Perech. Rast. Turkm. IV, 226; Grossh. Fl. Kavk. IV, 161; Mulk. op. cit.
839; Nikitin in Fl. Turkm. VII, 148; Gadzhiev in Fl. Azerb. VIII, 354.—
A. orientalis var. songorica (Trautv.) O. and B. Fedtsch. in Perech.
Rast. Turkm. IV (1911) 226.—Echinops acantholepis Jaub. and sp. Ill.
pl. or. Ill (1847-1850) 99; Bge. in Bull. Acad. Pétersb. VI, 412.—E.
acantholepis var. songorica Trautv. in Bull. Soc. Nat. Mosc. XXXIX
(1866) 368.—E. olivieri Jaub. and Sp. op. cit. 101.—TIe.: Jaub. and Sp.
op. cit. tab. 273 and 274; Nikitin, op. cit. Plat XXIII, Fig. 1.—Exs.: Pl.
or. exs. No. 72; Aucher, Herb. or. No. 4812; Sintenis, It. transcasp.-
pers. 1900-1901, No. 1886; Bornm. It. pers. alt. 1902, No. 7295.

Annual. Plants 5—25(40) cm high; stem white-tomentose and finely
glandular hairy, erect, simple or cymosely branched above (less often
almost from base). Leaves white arachnoid-hairy tomentose, linear-
lanceolate, involute, up to 5—6(8) cm long and I—1.5 cm wide, acuminate,
terminating in stiff bristles, narrowed toward base, sessile and more or
less amplexicaul, usually with sparse irregularly spaced, stiff, whitish,

'Treatment by I.A. Linczevsky.
*From the Greek words acantha—spine, and /epis—scale.

35

56

2-4 mm long bristles above and along margin. Heads up to 20-25 mm
wide and about 15 mm long, comprising 10-25 one-flowered capitula;
common involucre two or three-rowed, comprising 12-20 bracts; outer
bracts squarrose, almost leafy, linear-lanceolate, up to 20-30 mm long,
much longer than head, stiff and somewhat sparsely ciliate-setose along
margin, white arachnoid-hairy tomentose on outer side, densely pubescent
on inner side less and often glabrous and glossy at base; inner bracts
appressed, much shorter, 5-10 mm long and 1-1.5(2) mm wide, stiff
and rather densely ciliate-setose along margin, acuminate with up to 2—
3 mm long awn, densely arachnoid-hairy tomentose on outer side,
glabrous and glaucous on inner side. Capitula 10-12 mm long and about:
5 mm wide, sessile often two or three capitula fused together at base;
individual involucre dense, many-rowed, outer rows (brush) of white,
split, flexuous above and slightly plumose, 5—7 mm long bristles; middle
and inner involucral bracts stiff-leafy, densely covered with long curly
hairs on outer side, glabrous on inner side, glossy and green; innermost
bracts black-brown; middle bracts shorter, 5-7 mm long, linear-spatulate,
long plumose-ciliate in upper half; inner bracts longer, up to 10—12
mm, lanceolate, long plumose-ciliate above, with long, weakly recurved,
often reddish-reddish brown, up to 4—6 mm long setaceous awn. Corolla
reddish-violet (when dry) or light blue (according to Mulkyanyan, op.
cit., p. 841), about 5 mm long, limb about 3 mm and tube about 2 mm
long. Achenes 4—5 mm long and about 1.5—2 mm wide, densely
pubescent with appressed, upward directed, very short-plumose, long
hairs covering pappus; pappus comprising short, up to 0.5—0.7 mm long,
unequal, flat bristles irregularly connate at base in ring. Flowering I[V—
VII; fruiting VI—-VII. (Plate IV, Fig. 1).

Desert plains, foothills, and lower mountains; fixed hummocky sands
and sandy takyrs, in clayey and stony submountainous plains, dry clayey
and stony slopes. Caucasus: Eastern and southern Transcaucasia; Soviet
Central Asia: Aralo-Caspian Region (south), Balkhash Region (south),
Dzhungaria, Tarbagatai (south), Kyzyl-Kum, Kara-Kum, mountainous
Turkmenia, Amu-Darya, Syr-Darya, Pamiro-Alai (southern Tadzhikistan).
General distribution: Eastern Mediterranean (Syria), Armenia-Kurdistan,
Iran (in the southeast up to Baluchistan). Described from Syria or Iraq
(from the collections of Olivier and Brugier between Baghdad and Haleb
[Aleppo]). The type specimen was in Berlin (in Kunth Herbarium);
possibly, duplicates [isotypes] are available in Paris, where the main
collection of Olivier and Brugier is preserved.

Note. Soon after the establishment of Acantholepis Less., Jauber
and Spach (op. cit.) reduced it to a subgenus under Echinops L., and
included the two above-mentioned species in this subgenus. A little
later, Bunge (op. cit.), having seen Acantholepis Less. as only a
monotypic section of Echinops L. in Boissier (op. cit.), recognized

56

57

Acantholepis Less., however, as a separate monotypic genus without
accepting its other species described by Jauber and Spach; the probable
baselessness of the second species was quite convincing from the
excellent plates of the authors mentioned, which, it seems, very distinctly
demonstrate the absence of significant differences between the two
species depicted. Subsequent authors shared the opinion of Boissier,
indicating close affinity of Echinops L. and Acantholepis Less.
Mulkijanyan (op. cit., p. 840), who studied this question in great detail
recently, hypothesized that the “genus Acantholepis Less. is derived from
the genus Echinops L. and, probably originated from an ancestor of the
E. integrifolius Kar. and Kir. or E. humilis type.”

Tribe 12. CYNAREAE Less. in Linnaea, V (1830) 128 p. p. and
Synops. Comp. (1832) 4 p. p.; DC. Prodr. VI (1837) 499 p. p.; Boiss.
Fl. or. III (1875) 422 p. p.; O. Hoffm. in Pflanzenfam. IV, 5 (1894) 312
p. p.—Cinarocephalae Juss. Gen. pl. (1789) 171.—Acarnaceae Link,
Handb. I (1829) 684.—Fam. Cynaraceae Lindl. Nat. Syst. ed. 2 (1836)
151.—Tnb. Cynaroideae Benth. in Benth. and Hook. f. Gen. pl. II (1873)
211 p. p.—Trib. Carduées Baill. Hist. des Plantes, VIII (1882) 69
p. p.—Fam. Carduaceae Bessey in Ann. Miss. Bot. Gard. II, 2 (1915)
164.—Capitula homogamous or heterogamous, usually many-flowered,
homochromous or heterochromous. Involucres many-rowed; involucral
bracts imbricate, usually coriaceous or leafy, cuspidate, with diverse
type of spinules or scales, less often without appendages. All florets
tubular, bisexual (unisexual, in some Cirsium species, and then plants
dioecious), sometimes outer florets sterile, enlarged or not, funnel-shaped,
less often pistillate (their corolla bilobed). Receptacle covered with scales
or bristles, less often glabrous. Anthers sagittate at base, with diverse
(cuspidate, strigose, ciliate, fimbriate, split or entire) tips (anthers without
basal appendages only in Anura (Juz.) Tschern). Style usually with crown
(collar) of pollen collecting hairs below branching or thickened; its
branches free or fused, partly or up to tip. Corolla variously colored.
Leaves alternate.

Herbs and semishrubs, distributed throughout Europe, northern
Africa, and a large part of Asia; the largest number of genera of this
tribe are found in the Mediterranean Region, Northwest and Soviet
Central Asia and only a few genera are represented in America, Australia,
and Tropical Africa.

KEY TO SUBTRIBES OF CYNAREAE

1. Achenes with straight (occasionally slightly oblique) areole.... 2.
+ Achenes; with oblique or lateral areole .....-....9e¢ esti tees. weet <--e-
BAIR eee ence Anan vate ced Subtribe 3. Centaureinae O. Hoffm.

57

58

2. Achenes appressed-hairy or villous, without distinct margin at apex
epee. ald! bres melee! ageol Subtribe 1. Carlininae O. Hoffm.
+ Achenes glabrous with scattered divergent hairs (only in Polytaxis
Bge.); apex mostly with dentate margin... lees eseeeeeeeeeees
Ab paartadaubsebg AUER. eel owes Subtribe 2. Carduinae O. Hoffm.

Subtribe 1. CARLININAE O. Hoffm. in Pflanzenfam. IV, 5 (1894)
314.—Trib. Carlineae Cass. in Bull. Soc. Philom. (1815) 173 p. p.; Ej.
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. 1
(1826) 292 p. p.; I] (1826) 202 p. p—Subtrib. Carduineae Less. in
Linnaea, VI (1831) 86 p. p. and Synops.-Comp. (1832) 8 p. p.—Subtnb.
Cardopateae DC. Prodr. VI (1837) 527.—Subtrib. Xeranthemeae
DC. Ibid. 528.—Subtrib. Carilineae DC. Ibid. 531 p. p.; Benth. in
Benth. and Hook. f. Gen. pl. II (1873) 212; Boiss. Fl. or. II (1875)
442.—Achenes with straight areole, without distinct margin at apex,
villous. Pappus with one or two rows of scales, less often of bristles
(Atractylodes DC., Carlina L.).

KEY TO GENERA OF SUBTRIBE CARLININAE!

1. Pappus of more or less narrow aristate scales, usually very short-

barbate: but mot plUmOSe, cae be aca te a tech aeons case heeese eee a 2,
+ Pappus of thin plumose bristles or narrow scales, free almost to
basesing plumose SetaceOuS 1OBES sa.cc8t Artasis ort eye ee ee eee 6.

2+. All florets similansbisexuadl <fertil 624... shaste) adeteee ett aoe
Se ee ee eee ee ee ee 1569. Cousiniopsis Nevski
+ Ray florets pistillate, sterile or fertile; disk florets bisexual, fertile

Re ee en ey. A ER ae en Seen rere, cee ee 3
3. Corolla of ray (pistillate) florets bilabiate ............0.. ccc 4.
Corolla, ofyallyflorets weegubar dissec eee. Deane eee cen 5.
4. Corolla of bisexual florets very shallow 5-fid. Involucral bracts
not prickly. Capitula on peduncles........ 1570. Xeranthemum L.
+ Corolla of bisexual florets deeply 5-fid. Involucral bracts prickly.
Capituladsessiles... Aue. aha hehans 1571. Siebera J. Gay
5. Filaments united. Annuals ........0..... ee 1572. Chardinia Desf.
Filaments free. Perennials.................. 1573. Amphoricarpus Vis.
6. Innermost involucral bracts long, much longer than florets, radially
divergent orirecurvedt 50". esti Peete ee 1574. Carlina L.

'By I.A. Linczevsky.

59

+ Innermost involucral bracts shorter, not longer than florets, erect
SRN ONS CE 1 EEE, SABES 1575. Atractylodes DC.

GENUS 1569. Cousiniopsis Nevski? *

Nevski in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 4 (1937) 288.
Capitula many-flowered, small (about 10 mm wide), spherical-ovoid,
58 solitary. Involucre many-rowed, imbricate; involucral bracts stiff-leafy,

prickly; outer bracts with recurved, stiff, whitish appendage with three
to seven spinules; inner bracts with thick, less recurved cusp. Receptacle
slightly convex, punctate, densely covered with large almost transparent
scales closely enclosing florets; scales shallow split at apex into thin
bristles. Florets similar: all bisexual, tubular, regular; corolla with short
tube and deeply 5-partite limb having linear lobes. Anthers with long,
fimbriate-hairy, tail-like basal appendages; filaments glabrous. Style
gradually thickened in upper :part, with very short, flat, cylindrical
branches and with ring of short hairs at their base. Achenes obovoid,
sericeous-villous, obtuse, with straight areole. Pappus in two rows: outer
row of (8)10 lanceolate or narrow-lanceolate scales, approximate at base,
finely divided at apex, with short spinose continuation of midrib; inner
row as five-toothed crown, one-fourth to one-third as long as outer scales,
teeth split at tip into bristles; middle bristle aristate, long, almost equal
or as long as scales of outer row.

A short annual; leaves alternate, pinnatisect or toothed with prickly
lobes or teeth.

A monotypic genus.

1. C. atractyloides (Winkl.) Nevski in Tr. Bot. Inst. Akad. Nauk
SSSR, Ser. 1, 4 (1937) 288; Nikitin in Fl. Turkm. VII, 156.—Cardopatium
atractyloide Winkl. in Tr. Peterb. Bot. Sada, IX (1886) 517; O. and B.
Fedtsch. Perech. Rast. Turkm. IV, 227; Popov in Tr. Turk. Nauchn.
Obshch. I, 33; Vved. in Tr. Sredneaz. Gos. Univ., Ser. VIIIb, 102.—Ie.:
Nikitin, op. cit., Plate XXIII, Fig. 2—Exs.: H.F.A.M. No. 477; Sintenis,
It. transcasp.-pers. 1900-1901, No. 250.

Annual. Plants 5—15(25) cm high, sparsely white arachnoid-hairy;
stem branched from base, usually with fewer slender, whitish branches,
simple or slightly branched. Leaves 3—7 cm long, lower (basal) leaves

?Treatment by I.A. Linczevsky.

‘From the word Cousinia—name of a genus of plants of the Compositae, and the
Greek word opsis—external appearance; .based on the similarity with some species of this
genus.

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60

linear-lanceolate, prickly-toothed or pinnatisect, with spinescent lobes,
middle (very sparse on branches) and upper (usually crowded forming
rosettes below capitula) leaves similar to basal, but often pinnatisect,
with five to seven (nine) distant, prickly-toothed lobes. Capitula 8-10
mm wide, fewer, (3)5—10(15) on plant, borne singly in stem forks
(beginning right from base, almost at soil surface) and at apices of
branches. Involucral bracts more densely pubescent than other parts;
outer bracts four or five-rowed, gradually reduced towards outer side,
narrow-ovate to narrow-lanceolate, S—3.5 mm long and 2—1.5 mm wide,
with up to 3-4 mm long process, and spinules about 2 mm long; inner
bracts one or two-rowed, longer than outer, oblong-ovate, 6—7 mm long
and 2.5—3 mm wide, dark purple above, with about 1-2 mm long cusp;
receptacular scales 5-6 mm long and 2.5-3 mm wide, split at apex up
to 0.5-1 mm. Corolla centaury or lilac-blue, about 8 mm long, with
about 4 mm long limb and about 4 mm long tube. Achenes 1.5—2 mm
long and about 1 mm wide; pappus 3.5—4 mm long, light straw-yellow,
inner row (crown) about 1 mm long, aristate tips of pappus scales and
teeth of crown barbellate. Flowering I[V—V; fruiting VI—VIII. (Plate IV,
Bigep2)):

Submountainous plains, foothills and lower mountains on loamy and
loamy-stony soils, usually in regions of variegated gypsum rocks.—Soviet
Central Asia: Kyzyl-Kum (extreme south: Kenimekh Region), Kara-Kum
(along Middle Murgab), mountainous Turkmenia (Ashkhabad and Dushak
regions), Amu-Darya, Syr-Darya (Fergana Valley, Namangan Region),
Pamiro-Alai (southwest and western lower mountains). General
distribution: Possibly occurs in northern Iran and northern Afghanistan.
Described from the foothills of the interfluve of the Kafirnigan-Vakhsh
rivers between Kabadian and Dzhilikul. Type in Leningrad.

Note. In describing this species, Winkler (op. cit.) indicated its
intermediate position between Cardopatium Juss. and Atractylis L., but
still included it under the former genus, placing it under the monotypic
subgenus Distemma Spach [Monographia generis Cardopatium, Ann. Sc.
Nat. sér. 3, V (1846) 233-247], which until then included the only
species, C. boryi Spach from Laconia, which differs from other species
of the genus by its two-rowed pappus, the second row being coronate,
five-toothed, and very short. Boissier (Boiss. Fl. or. III. 1875, 444) in
the note to C. boryi Spach, noted that the flowers in the herbarium
specimens of this species preserved in Paris are damaged by insects,
and, therefore, the description of the pappus is given only from Spach.
Popov (op. cit.) doubted the correctness of including this species in
Cardopatium, and Vvedenskii (op. cit.) pointed out that it should be
considered as a separate genus. Nevski (op. cit.), in describing the genus
Cousiniopsis, writes that Cardopatium Juss. differs from Cousiniopsis

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61

by having a one-rowed pappus, and the two-rowed pappus in
Cardopatium boryi Spach is doubtful (“pappus biseriatus in C. boryi
Spach dubius est!”’).

It can be seen from what is stated above that the taxonomic position
of Cousiniopsis is not yet entirely clear; to resolve this, probably first
of a. a critical revision of the genus Cardopatium Juss., apparently not
entirely homogeneous, will be necessary.

GENUS 1570. Xeranthemum L.!: 2

L. Sp. pl. (1753) 857; DC. Prodr. VI (1837) 528; Benth. and Hook.
f. Gen. pl. VII (1873) 463; O. Hoffm. in Pflanzenfam. IV, 5 (1892)
315.—Harrisonia Neck. Elem. I (1790) 84.—Xeroloma Cass. in Dict.
sc. nat. LIX (1829) 120.

Capitula many-flowered, small (up to 15—20 mm wide), campanulately
or hemispherically ovoid or oblong-cylindrical, solitary, terminal on long
leafless branches. Involucre many-rowed; imbricate; involucral bracts
scarious, not prickly; outer bracts appressed, gradually reduced towards
outer side; innermost bracts oblong, colored (pink, brown to purple or
violet) or white, erect or reflexed. Receptacle flat, densely covered with
stiff, narrow, concave, undivided or divided scales. Florets variable; ray
florets pistillate, very few, sterile, with a short bilabiate corolla; lips
small, erect, outer lip entire or three-toothed, inner smaller and bifid;
disk florets bisexual, fertile, with regular tubular corolla, and very short
five-fid limb. Anthers sagittate at base, with short, fimbriate or cuspidate
appendages or obtuse; filaments free, glabrous. Style of bisexual florets
with very short erect or subsequently recurved branches. Achenes of
pistillate florets thin, empty, with or without pappus of fine scales;
achenes of bisexual florets oblong-cuneate, narrow-obdeltoid or almost
turbinate, with short appressed pubescence, cylindrical or angular on
inner side, truncate at apex and with one-rowed pappus or with more or
less numerous (5—15) aristate-acuminate scales.

Annuals. Nonprickly, white pubescent-hairy plants with alternate,
narrow, entire leaves.

The genus includes five or six species growing in the Mediterranean
Region.

Type species: Xeranthemum annuum L.

Economic Importance: The species of this genus, in particular X.
annuum L. and X. squarrosum Boiss., have long been used in the southern

'Treatment by I.A. Linczevsky.
*From the Greek words Xeros—dry, and anthemon—flower.

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USSR as ornamental plants for dry winter bouquets, wreaths; and
garlands. The seeds of X. cylindraceum Sm. contain oil with the iodine
number 105.8 (after A.A. Grossheim).

Note. The basic taxonomic studies on the genus Xeranthemum L.
have been cited at the corresponding places of the present treatment.
The brief but interesting review of the Czechoslovakian botanist J. Chrtek
(“The relation between genus Xeroloma Cass. and genus Xeranthemum
L.,” Novitates bot. Hort. bot. Universitatis Carolinae Pragensis, 1961,
pp. 3-5), published recently, should also be mentioned; it considers
Xeroloma Cass. as a separate genus, and proposes two subgenera
(Xeranthemum and Xeranthelia Chrtek) for the remaining four species
of Xeranthemum L. and two sections (Xeranthemum and Longipapposa
Chrtek), i.e., it proposes a rather detailed infrageneric division. For
technical reasons (late receipt of the above mentioned issue of Novitates)
I could not cite the new taxa proposed by J. Chrtek in the main text.

1. Capitula more or less narrow-cylindrical at flowering, 5—8(10) mm

Wide, enlarged. after GON CNS (i... t cece tytisc oe. Sn eee eae 2
+ Capitula ovoid or hemispherically ovoid at flowering 10—15(20)
TIN WAC Cs rc sett edaatt wade lade acai obit acoas mineral eines aad Ba ade elas 3.

2. Outer involucral bracts glabrous, with short cusp at tip. Pappus
scales) five. almost ,cotia lis: ete haat chat Oe acetate ate dee
Ke aSil a W Miei ila ver act aula tale 4. X. longipapposum Fisch. and Mey.
+ Outer involucral bracts finely tomentose on outer surface, obtuse.
Pappus scales. (7)10—15,, highly umequall pic, .2.<-9¢-ccdescceck ate eenucPS- atndd
eS ee ee ees eee rey re 5. X. cylindraceum Sm.
3. Inner involucral bracts erect, brown to purple or violet; outer bracts
usually with wide, longitudinal brown stripe ..............:eeeeeeeeeeeeees
ee SO Se ee eee | Aer 3. X. inapertum (L.) Mill.
+ Inner involucral bracts radially recurved, pinkish or bright pink,
occasionally white; outer bracts whitish or yellow, glossy, sometimes
with narrow longitudinal, brown to purple stripe ..............-:0ee 4.
4. Pappus scales deltoid at base, very gradually attenuate into short
subulate awn, much shorter than corolla and receptacular scales;
capitula often with 100-120 florets................ 1. X. annuum L.
+ Pappus scales short-deltoid at base, abruptly attenuate into long
subulate awn, almost as long as corolla and receptacular scales;
capitula often with 30-40(60) florets ... 2. X. squarrosum Boiss.

Section 1. Xeranthemum—Euxeranthemum DC. Prodr. VI (1837)
528, pro sect. nom. illeg—Pappus scales five.

1. X. annuum L. Sp. pl. (1753) 857; M.B. FI. taur.-cauc. IT, 302; J.
Gay in Mém Soc. Hist. Nat. Paris, III, 358; Kalenicz. in Bull. Soc. Nat.

63

Plate IV.

1—Gymnaster savatieri (Makino) Kitamura, habit, disk floret with achene, ligulate peripheral
floret, stamens, involucral bract; 2—Conyzanthos [sic;—thus] graminifolius (Spreng.)
Tamamsch., habit, achene with pappus, stamen, involucral bract, tubular floret, tubular-ligulate
floret, style; 3—Tripolium vulgare Nees, habit, achene with pappus, disk floret, peripheral
ligulate floret, style, involucral bract, stamen.

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Mosc. VII, 188; Boiss. Fl. or. III, 444 : Schmalh. Fl. Il, 92; Hegi. Ill.
Fl. VI, 2, 814; Grossh. Fl. Kavk. IV (1934) 162; Iljin in Fl. Yugo-Vost.
VI, 377.—X. annuum f. gracilius Fisch. and Mey. in Nouv. Mém. Soc.
Nat. Mosc. IV (1835) 336.—X. radiatum Lam. FI. Fr. II (1793) 48; DC.
Prodr. VI, 528.—xX. radiatum var. a. macranthum C. Koch in Linnaea,
XVII (1843) 38.—X. radiatum var. a. caule stricto etc. Ldb. Fl. Ross.
II, 2 (1845-1846) 658.— YX. annettae Kalenicz. in Bull. Soc. Nat. Mosc.
VII (1834) 194.—Centaurea dubia S.G. Gmel. Reise, I (1770) 126.—
Ic.: Gmel, Ibid. tab. 20; J. Gay, op. cit. tab. 7, fig. 1; Kalenicz. op. cit.
tab. VI, fig. 1 and 3; Fisch. and Mey. op. cit. tab. XII; Rchb. Iconogr.
bot. VII, fig. 864; O. Hoffm. op. cit. fig. 144, B-D; Hegi, op. cit. fig.
517, d-h, fig. 518; Iljin, op. cit. Fig. 694.—Exs.: Novopokr. Gerb. Donsk.
Fl. No. 50; Callier, It. taur. 1900, No. 642; Pl. bulg. exs. No. 99; Orph.
Fl. Graec. exs. No. 754; Fl. hung. exs. No. 886; Fl. exs. austro-hung.
No. 976; Dorfl. Herb. norm., No. 4137.

— Annual. Plants 10-40(60) cm high, appressed white arachnoid-hairy
tomentose. Stem erect, usually flexuous, virgate, with solitary terminal
capitula on leafless branches. Leaves up to 3-4 cm long and 2—8(12)
mm wide, almost sessile; lower leaves narrow-lanceolate, upper highly
reduced, linear. Capitula with 100—120 florets, hemispherical-ovoid at
flowering, (10)15—20 mm wide. Involucral bracts scarious, glabrous; outer
bracts whitish or yellowish, sometimes with longitudinal brown to purple
stripe, ovate, abruptly short acuminate, more or less appressed; inner
bracts two times as long as outer, up to 5 mm wide, elliptical-linear,
radially recurved, pinkish or bright pink (occasionally white). Anther
appendages fimbriate. Achenes dark brown, oblong-deltoid, cylindrical-
angular, 3-5 mm long and about | mm wide; pappus scales five, unequal,
2-3 mm long, deltoid at base, very gradually attenuate upward into
short subulate awn, much shorter than corolla and receptacular scales.
Flowering VI—X; fruiting VII—X.

In steppes, on chalk outcrops, sands, dry slopes, among shrubs in
lower mountains, sometimes as a weed.—European Part: Middle Dnieper
(south), Volga-Don (south), Trans- Volga Region (southwest), Bessarabia,
Black Sea Region, Crimea, Lower Don, Lower Volga (west); Caucasus:
Ciscaucasia, Dagestan. General distribution: Central Europe (south),
western Mediterranean, Balkans-Asia Minor (west). Described from
Western Europe (Austria). Type in London.

2. X. squarrosum Boiss. Diagn. pl. or. ser. I, VI (1845) 101, incl.
a. pictum Boiss. and B. unicolor Boiss.; Boiss. Fl. or. Il], 444; Grossh.
Fl. Kavk. IV, 162; Nikitin in Fl. Turkm. VII, 154; Gadzhiev in FI.
Azerb. VIII, 356.—NX. radiatum var. c. micranthum C. Koch in Linnaea,

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XVII, (1843) 38.—X. radiatum var. b. caule subsquarroso etc. Ldb. FI.
Ross. II, 2 (1845-1846) 658.—Ic.: Gadzhiev, op. cit., Plate XXXIX,
Fig. 2.—Exs.: Herb. Fl. Cauc. No. 248; Aucher, Herb. or. No.
4777; Balansa, Pl. or. 1854, No. 269; 1855, No. 635; 1856, No. 882;
Kotschy, It. cilic.-kurd. 1859, No. 308 and Pl. Syr. bor. 1862, No. 75;
Bornm. Pl. exs. Anatol. or. 1889, No. 546; Sintenis, It. or. 1892, No.
4914.

Annual. Plants 10—-40(60) cm high, appressed white arachnoid-hairy
tomentose. Stem erect, usually slender-virgate profusely branched with
solitary terminal capitula on leafless branches. Leaves up to 3-4 cm
long and 3-4 mm wide, oblong-linear, sessile; lower leaves decurrent,
upper highly reduced. Capitula with 30—-40(60) florets, hemispherical-
ovoid at flowering, 10—15 mm wide. Involucral bracts scarious, glabrous;
outer bracts whitish, sometimes with longitudinal brown to purple stripe,
glossy, ovate, abruptly short-acuminate, weakly squarrose; inner bracts
two to three times as long as outer bracts, up to 2.5—-3.5 mm wide,
linear-elliptical, radially recurved, glossy, pinkish or bright pink. Anther
appendages fimbriate. Achenes dark brown, oblong-obdeltoid, cylindrical-
angular, 3-4 mm long and about 1 mm wide; pappus scales five, unequal,
3—6 mm long, short-deltoid at base, abruptly attenuate into long subulate
awn, almost as long as corolla and receptacular scales. Flowering VI—
VU; fruiting VII-VIII.

Foothill plains to middle mountain zone (in Azerbaidzhan up to
1,900 m), on dry, shallow, stony and rocky slopes, and among shrubs.—
Caucasus: Ciscaucasia, Dagestan, western, eastern and southern
Transcaucasia, Talysh; Soviet Central Asia: mountainous Turkmenia
(Kopetdag Range). General distribution: Eastern Mediterranean
(northeast), Balkans-Asia Minor (east), Armenia-Kurdistan, Iran
(northwest). Described from the Caspian Sea, from Gilan and the vicinity
of Diyarbakir in northern Mesopotamia. Type in Geneva; isotype in
Leningrad.

Note. Boissier, who described this species (Boiss. Fi. or. p. 445),
included it here as a separate species, but later questioned whether
it should be considered as a variety of X. annuum. However, subsequent
work did not confirm the doubts, and present Caucasian plant
taxonomists have clearly distinguished X. squarrosum Boiss. from xX.
annuum L.

Boissier (Boiss. Fi. or. |.c.) included under the synonyms of X.
squarrosum §. unicolor Boiss. the species X. annettae Kalenicz.,
described by Kharkov botanist I].0. Kaleniczenko, from the vicinity of
Stavropol in the northern Caucasus. If these species should prove
convincingly to be the same, the name X. annettae Kalenicz. would take
priority (published 11 years earlier) over X. squarrosum Boiss. However,

66

the study of the specimens of X. annettae preserved in Leningrad shows
that they should be referred to X. annuum L. ;
Among the herbarium specimens cited by Boissier in describing
this species, it appears to me more appropriate to select as type specimens
those collected by Aucher from Gilan (Aucher, Herb. or. No. 4777), as
these specimens correspond well to the diagnosis of Boissier, are precisely
identified thanks to the presence of the number, and, besides, were
distributed as duplicates to the largest herbaria of the world.

3. X. inapertum (L.) Mill. Gard. Dict. ed. 8 (1768) No. 2; Willd.
Sp. pl. Ill, 3, 1902; J. Gay in Mém. Soc. Hist. Nat. Paris, HI, 360;
Kalenicz. in Bull. Soc. Nat. Mosc. VII, 191; Ldb. FI. Ross. II, 2, 658;
Boiss. Fl. or. III, 445; O. and B. Fedtsch. Perech. Rast. Turkm. IV, 227;
Hegi, Ill. Fl. VI, 2, 815; Grossh. Fl. Kavk. IV, 161; Karjagin, FI.
Apsherona (1952) 382; Gadzhiev in Fl. Azerb. VIII, 357—X. annuum
B. inapertum L. Sp. pl. (1753) 858, p. p. excl. ic. Moris.—xX. erectum
Presl, Delic. Prag. I (1822) 106; Fisch. and Mey. in Nouv. Mém. Soc.
Nat. Mosc. IV, 338; DC. Prodr. VI (1837) 529; cum. B: meyeri DC.—
X. foetidum Moench, Meth. pl. (1794) 582, p. p. nom. superfl.—Ic.: J.

5 Gay, op. cit. tab. 7, fig. 2; Kalenicz. op. cit. tab. VI, fig. 2; Fisch. and

Mey. op. cit. tab. XII; Rchb. Iconogr. bot. VII, fig. 863; Hegi, op. cit.
fig. 517 i-l; Karjagin, op. cit. Plate XX A-—B; Gadzhiev, op. cit. Plate
XXXIV, Fig. 4, Exs.: Fiori, Bég. Pamp. FI. Ital. exs. No. 988; Todaro,
Fl. Sic. exs. No. 600; FI. Gall. et Germ. exs. Nos. 686, 1905 and 1905
bis; Fl. exs. austro-hung. No. 977.

Annual. Plants 10-40 cm high, appressed white arachnoid-hairy
tomentose. Stem erect, usually flexuous, virgate, with solitary terminal
capitula on leafless branches. Leaves up to 5—6(8) cm long and 4—7(10)
mm wide, almost sessile, decurrent; lower leaves oblong-obovate, upper
leaves linear-oblong. Capitula with 30—40 florets, ovoid at flowering,
10-15 mm wide. Involucral bracts scarious, glabrous, whitish, and usually
with brown, longitudinal, broad stripes; outer bracts ovate, abruptly short-
acuminate; inner bracts about one and one-half times as long as outer,
up to 2.5-3 mm wide, lanceolate, acute, erect (not radially recurved),
brown to purple or violet. Anther appendages setaceous. Achenes dark
gray, narrow-obdeltoid, cylindrical-angular, about 5 mm long, and
1—-1.5 mm wide; pappus scales five, more or less equal, about 5 mm
long, deltoid or deltoid-lanceolate, usually gradually attenuate into thin
finely barbed awn. Flowering V—VIII; fruiting VII—VIII. (Plate IV,
Fie: %3)!

Dry, shallow, and stony slopes, up to middle mountain zone,
sometimes as weed.—European Part: ?Crimea; Caucasus: Ciscaucasia,
Dagestan, eastern Transcaucasia, Talysh. Soviet Central Asia: (Reported

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by O. and B. Fedtsch. op. cit. from “Northern Turkmenia,” according to
Karelin, but not confirmed by subsequent collections). General
distribution: Central Europe (south), Atlantic Europe (southeast), western
Mediterranean, eastern Mediterranean, Balkans-Asia Minor, ? Armenia-
Kurdistan. Described from Western Europe (Italy, Switzerland, France).
Type in London.

4. X. longipapposum Fisch. and Mey. in Nouv. Mém. Soc. Nat.
Mosc. IV (1835) 337 (“longepapposum’’); DC. Prodr. VI, 529; Ldb. FI.
Ross. II, 2, 659; Boiss. Fl. or. III, 445; O. and B. Fedtsch. Perech. Rast.
Turkm. IV, 227; Grossh. Fl. Kavk. IV, 162; Nikitin in Fl. Turkm. VII,
153; Gadzhiev in Fl. Azerb. VIII, 357.—Ic.: Fisch and Mey. op. cit.
tab. XII; Gadzhiev, op. cit., Plate XXXIV, Fig. 1—Exs.: Pl. or. exs.
No. 95; Sintenis, It. transcasp.-pers. 1900-1901, Nos. 681, 1852; Kotschy,
Pl.-Pers. austr. No. 542.

Annual. Plants 10—30(50) cm high, appressed white arachnoid-hairy
tomentose. Stem erect, usually divaricate and rather densely branched,
slender-virgate, with solitary terminal capitula on leafless branches.
Leaves up to 3-4 cm long and 2—3(5) mm wide, almost sessile, elliptical-
linear to linear. Capitula with 10-15 florets, narrow-oblong to cylindrical
at flowering, 5-6 mm wide. Involucral bracts scarious, glabrous; outer
bracts whitish, small, deltoid to ovate, abruptly short-acuminate; inner
bracts three times as long as outer, up to 2—2.5 mm wide, linear to
linear-lanceolate, acuminate, erect (not radially recurved), lilac-pink, later
turning white. Anther appendages acute. Achenes silvery white, narrow-
cuneate, almost linear, cylindrical-angular, 5—6 mm long and about 1
mm wide; pappus scales five, almost equal, 9-10 mm long, narrow-
lanceolate, finely ciliate-barbate, very gradually acuminate, on average
about one and one-half times as long as achenes. Flowering V—VII;
fruiting VI—VIII.

Lower and middle mountain zone, on dry, shallow, and stony slopes,
sometimes in steppe phytocenoses.—Caucasus: Southern Transcaucasia,
Talysh; Soviet Central Asia: Dzhungaria-Tarbagatai (Dzhungarian
Alatau), mountainous Turkmenia, Pamiro-Alai, Tien Shan. General
distribution: Balkans-Asia Minor (east), Armenia-Kurdistan, Iran.
Described from northwestern Iran (vicinity of the village of Seidkhoji
in the region of Khoi). Type in Leningrad.

Section 2. Xeroloma (Cass.) DC. Prodr. VI (1837) 529.—Xeroloma
Cass. in Dict. sc. nat. LIX (1829) 120 p. gen.—Pappus scales (7)10—15.

5. X. cylindraceum Sm. Prodr. fl. Graec. II (1813) 172; J. Gay in
Mém. Soc. Hist. Nat. Paris, III, 362; Kalenicz. in Bull. Soc. Nat. Mosc.

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VII, 197; DC. Prodr. VI, 529; Ldb. Fl. Ross. II, 2, 659; Boiss. FI. or.
III, 446; Schmalh. FI. II, 92; Hegi, Ill. Fl. VI, 2, 813; Grossh. Fl. Kavk.
IV, 162; Gadzhiev in Fl. Azerb. VIII, 356—X. annuum B. inapertum L.
Sp. pl. (1753) 858, p. p. quoad ic. Moris.—xX. foetidum Moench, Meth.
pl. (1794) 582, p. p. nom. superfl—yX. inapertum auct. non Mill. : M.B.
Fl. taur.-cauc. II (1808) 302 and III (1819) 568; Fisch. and Mey. in
Nouv. Mém. Soc. Nat. Mosc. IV, 340.—X. incanum foetens, flore minore
Moris, Plant. hist. univers. III (1699) sect. 6, tab. 12, fig. 1. excl.
descript.—Ie.: J. Gay, op. cit. tab. 7, fig. 3, Kalenicz. op. cit. tab. VI,
fig. 4; Fisch. and Mey. op. cit. tab. XII; Rchb. Iconogr. bot. VII, fig.
862; Hegi, op. cit. fig. 517a—c; Gadzhiev, op. cit. Plate XXXIV, Fig.
3.—Exs.: GRF No. 1781; Herb. Fl. cauc. No. 247; Aucher, Herb.
or. No. 3242; Bornm. It. pers.-turc. 1892-1893, No. 1410; Orph. FI.
Graec. exs. No. 755; Fl. exs. austro-hung. No. 978; Fl. hung. exs. No.
886; Pl. Bulg. exs. No. 98; Fl. Gall. et Germ. exs. Nos. 884, 1024,
1024 bis.

Annual. Plants 10—40(60) cm high, appressed white arachnoid-hairy
tomentose. Stem erect, usually straight and slender-virgate, with solitary
terminal capitula on leafless branches. Leaves up to 3—4(6) cm long and
2—4(7) mm wide, nearly sessile, decurrent; lower leaves lanceolate, upper
linear. Capitula with 10-15 florets, narrow oblong-cylindrical at
flowering, 5-10 mm wide, distinctly enlarged after flowering, turbinate
or campanulate. Involucral bracts appressed; outer bracts ovate, obtuse,
stiff-leafy and finely tomentose on outer surface, with wide light-colored
membranous border along margin; inner bracts slightly longer than outer,
up to 2.5—3 mm wide, lanceolate, subacute, almost erect (only weakly
radially divergent), pale pinkish or pink, later turning dark brown. Anther
appendages aristate. Achenes dark gray, narrow-obdeltoid, compressed.
5—6 mm long and about 2 mm wide; pappus scales (7)10—15, unequal,
(1.5-—5)2.5—5 mm long, finely barbed along margin, narrow-deltoid below,
abruptly attenuate into subulate awn, on average, about half as long as
achene. Flowering V—VIII; fruiting VI—VIII(1X).

Foothill plains to middle mountain zone; on dry shallow and stony
slopes, in shrubby areas and grassy phytocenoses, sometimes as a weed.—
European Part: Bessarabia, Crimea; Caucasus: Ciscaucasia, Talysh.
General distribution: Central Europe (south), Atlantic Europe (south),
western and eastern Mediterranean, Balkans-Asia Minor, Armenia-
Kurdistan. Described from the Mt. Bithynian Olympus (Turkey). Type
in London (?).

Note. In accepting X. cylindraceum Sm. as the name for this species
I am following Gay, the author of the first monograph of the genus
Xeranthemum L. in almost its present circumscription (Gay, op. cit. 1827).
Gay provided quite precise delimitation of the two species, which earlier

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69

were combined under the name X. inapertum (L.) Mill. (1768). Besides,
Gay acted exactly in accordance with the modern rules of botanical
nomenclature, retaining the initial name, 1.e., X. inapertum (L.) Mill., for
one of these species, and adopting the earliest valid name X. cylindraceum
Sm. (1813) for the other (the circumscription of which can easily be
determined from Morison’s polynomial and figure cited above and cited
by both Linnaeus and Smith). The name X. foetidum Moench (1794),
adopted by some authors for this species, cannot be considered valid, as
it is nomenclaturally superfluous (nomen superfluum), given that
Xeranthemum inapertum Linn. is listed as a synonym in his publication.

GENUS 1571. Siebera J. Gay':?

J. Gay in Mém. Soc. Hist. Nat. Paris, III (1827) 344, in adnot,
nom. conserv.; DC. Prodr. VI (1837) 531; Benth..and Hook. f. Gen.
pl. II (1873) 464; O. Hoffm. in Pflanzenfam. IV, 5 (1892) 315.—
Sieberia Spreng. Anleit. 2, 1 (1817) 282.—Siebera Hoppe, Flora 2 (1819)
24.

Capitula many-flowered, small (up to 12-15 mm wide), campanulate.
Involucre many-rowed, imbricate; involucral bracts wider below, abruptly
attenuate into long, stiff, later spiny cusp; inner bracts long, much longer
than florets, gradually reduced outward. Receptacle flat, densely covered
with somewhat deeply incised scales. Florets variable: peripheral florets
pistillate, fewer, sterile; their corolla bilabiate with short tube and erect
lips; outer lip entire or three-toothed, inner smaller, bifid; disk florets
bisexual, fertile, with tubular regular corolla, thin corolla tube, and
slightly wider, deeply five-fid limb with narrow lobes. Anthers sagittate
at base, with minute sharp cusp; filaments free, glabrous, flattened. Style
of bisexual floret with or without very short branches. Achene of bisexual
floret oblong-cuneate, sparsely pubescent, narrow at base, almost
cylindrical, with numerous longitudinal grooves, and truncate apex;
pappus cornate, of one row of many (8—10) transparent, subulately
acuminate scales.

Annual, short, prickly (except involucre), whitish pubescent-hairy
plant with alternate, narrow, entire leaves.

A monotypic genus.

1. S. pungens (Lam.) DC. Prodr. VI (1837) 531, incl. B. nana DC.;
Boiss. Fl. or. II, 447; Nikitin in Fl. Turkm. VII, 154.—X. gayana Cass.

'Treatment by I.A. Linczevsky.
*In honor of the botanist and traveler from Prague, Franz Wilhelm Sieber (1785-1844).

69

70

in Dict. sc. nat. LIX (1829) 126.—Xeranthemum pungens Lam. Dict. III
(1789) 236.—Ie.: Lam. Ibid. III, [V (1797) tab. 692, fig. 2.—Exs.:
Aucher, Pl. or. No. 3351 (sec. Boiss. op. cit.); Kotschy, Pl. Pers. austr.
1845, No. 385; Bornm. It. pers. alt. 1902, No. 7296.

Annual. Plants (5)10—20(40) cm high, appressed white arachnoid-
hairy tomentose. Stem simple or often branched from base, with more
or less curved, ascending or almost erect branches. Leaves up to 2—3(5)
cm long and 4—6(8) mm wide, sessile, elliptical to linear, rather broadly
rolled down along margin. Capitula with 12—20 florets, up to 12—15
mm wide, solitary terminal on branches and surrounded by crowded,
almost rosulate apical leaves, sometimes, with almost or totally abortive
stems nearly at soil surface. Involucral bracts ovate or broad-lanceolate
in lower part, finely white-tomentose, with wide membranous border,
abruptly attenuate into long (up to 25-30 mm) awn, more spiny in fruit,
and pinkish or purple in inner bracts; scales of receptacle shorter than
florets, 11-14 mm long, free almost to base in linear, sinuate, subulate,
1 mm wide lobes. Achenes of bisexual florets whitish, 7-9 mm
long and about 1 mm wide; pappus about 9-12 mm long, distinctly
longer than corolla; pappus scales almost equal, linear-lanceolate or
lanceolate, serrulate, gradually acuminate above, usually with more or
less distinct midrib, terminating into finely subulate, finely plumose-
barbate, 3-4 mm long awn. Flowering IV(VI); fruiting V(VII). (Plate V,
Fig. 1).

Dry slopes in foothills and lower mountains sometimes on cultivated
lands (according to Boissier, op. cit.).—Soviet Central Asia: mountainous
Turkmenia (Pul-i-Khatum). General distribution: Eastern Mediterranean,
Balkans-Asia Minor (eastern), Iran. Described from eastern Mediterranean
(“Levant”). Type possibly in Paris.

Note. S. pungens (Lam.) DC. has so far been collected in our country
only once, in 1895, by S.I. Korshinsky from southern Turkmenia,
near Pul-i-Khatum. The specimens were in the herbarium under an
incorrect name until they were correctly identified by M.M. Iljin in
1935.

The variety B. nana DC. mentioned above was described on the
basis of dwarf specimens, apparently stunted because of water stress;
such specimens are known from several places in its distribution
area.

J. Gay, who described the genus Siebera is, however, not the author
of the specific combination, because he did not publish such a
combination (cf. p. 32 of International Code of Botanical Nomenclature);
only 10 years later, De Candolle (DC. op. cit.) published an effective
and actually valid specific combination.

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71
GENUS 1572. Chardinia Desf.':?

Desf. in Mém. Mus. Paris, III (1817) 455; DC. Prodr. VI (1837)
530; Benth. and Hook. f. Gen. pl. II (1873) 464; O. Hoffm. in
Pflanzenfam. IV, 5 (1892) 315.

Capitula many-flowered, small (up to 20-25 mm wide), campanulate
or turbinate, solitary, terminal on long leafless peduncles. Involucre
many-rowed, imbricate; involucral bracts appressed, foliaceous-scaly or
scaly, greenish-white, nonprickly; outer bracts appreciably reduced toward
outer side; innermost bracts slightly longer, with slight pigmentation at
base. Receptacle flat, densely covered with stiff and narrow scales. Florets
variable; outer florets pistillate, very few (one to three), fertile; disk
florets bisexual, fertile. Corolla of all florets tubular, regular, three to
five-fid in the pistillate florets, five-fid in bisexual flowers, limbs with
narrow and stiff lobes, fused above in some pistillate florets or free
right from base in all florets. Achenes in pistillate florets oblong-obovoid,
almost flat, with two-toothed lateral wings and ventral keel, corniculate
above; pappus without or with 1—4 unequal stiff scales. Achenes of
bisexual florets narrow-cuneate or cuneate, cylindrical, with numerous
(15-20) fine oblong grooves, and truncate apex; pappus coronate, of
one row of many (8-10) large aristate-acuminate scales.

Annual, often short, nonprickly, white-pubescent plants with
alternate, narrow, entire leaves.

So far two species have been described in this genus.

Type species: Chardinia orientalis (L.) Ktze.

1. Achenes of bisexual florets narrow-cuneate, densely covered below
(over one-third to half length) with long appressed bristles, sparsely
short-setose above or almost glabrous (with few short bristles)...
BN A ce Meat ae Bh al ce 1. C. orientalis (L.) Ktze.

+ Achenes of bisexual florets cuneate (or oblong-turbinate), almost
entirely covered with dense, long, appressed bristles....................
SA aiid ie Mt CORA erent Oe I aaa cheek 2. C. macrocarpa C. Koch

1. C. orientalis (L.) Ktze. in Tr. Peterb. Bot. Sada, X (1887) 201;
O. and B. Fedtsch. Perech. Rast. Turkm. IV, 227; Grossh. Fl. Kavk. IV,
163; Papava in Zam. Sist. Geogr. Rast. [Tbilisi] 13, 80 and Fl. Gruzii,
VIII, 432; Nikitin in Fl. Turkm. VII, 155; Gaddzhiev in Fl. Azerb. VIII,
358.—C. orientalis var. erivanica Tamamsch. in Fedde, Report. XXX VIII

(1935) 171 (sphalm. “verivanica’’)—Xeranthemum annuum y. orientale

'Treatment by I.A. Linczevsky.
*In honor of Jean Chardin (1643-1713), the French traveler of the Orient.

71

q2

L. Sp. pl. (1753) 957.— X. orientale Mill. Gard. Dict. ed. 8 (1768) No.
3; Willd. Sp. pl. HI, 1902.—C. xeranthemoides Desf. in Mém. Mus.
Paris, III] (1817) 456; J. Gay in Mém. Soc. Hist. Nat. Paris, III, 366;
Fisch. and Mey. in Nouv. Mém. Soc. Nat. Mosc. IV, 341; DC. Prodr.
VI, 530; Ldb. Fl. Ross. II, 2, 659; Boiss. Fl. or. III, 446.—Ie.: Desf. op.
cit. tab. 21; J. Gay, op. cit. tab. 8; Fisch. and Mey. op. cit. tab. XIII; O.
Hoffm. op. cit. fig. 71, G, H; Papava, op. cit. 84, Fig. I]—Exs.: GRF
No. 883; Ed. H.B.P. No. 98; Aucher, Herb, or. No. 3350; Kotschy, Pl.
Pers. austr. Nos. 217, 807 and It. syr. 1855, No. 471; Balansa, Pl. or.
1855, Nos. 636, 1857, No. 1280.

Annual. Plants 10—30(50) cm high, appressed white arachnoid-hairy .
tomentose. Stem erect, simple or often more or less strongly branched,
less often from base, usually with weakly curved and somewhat thick
branches. Leaves up to 4—5(7) cm long, and 10-15 mm wide; narrow-
elliptical to narrow-lanceolate, almost linear, obtuse or subacute; lower
leaves basally narrowed into petiole; upper leaves sessile or almost
sessile. Capitula with 8—10 florets, almost cylindrical when young, greatly
enlarged at fruiting, broad-turbinate, up to 10—20(25) mm wide. Involucral
bracts glabrous; outermost very small, broad-ovate; middle bracts much
larger, broad-obovate, acuminate and mucronate, herbaceous in middle
with wide membranous border; inner bracts narrower, broad-lanceolate
or lanceolate, more gradually long acuminate; receptacular scales much
shorter than florets, narrow-linear, 10-12 mm long and 0.5—2 mm wide.
Achenes of pistillate florets glabrous, 10-12 mm long and 4-6 mm
wide; achenes of bisexual florets narrow-cuneate, 8-10 mm long and 1-
2 mm wide, densely covered below (over half to one-third length) with
long bristles, in middle one-third (or often almost to tip) sparsely short-
setose, upper one-third (sometimes upper two-thirds) almost glabrous
(with very few short bristles). Pappus (10)12—17 mm long, much longer
than corolla; pappus scales lanceolate, (1.5)2—3 mm wide, very gradually
attenuate into aristate cusp, thicker at base, with distinct midrib,
puberulent, margin very finely ciliate. Flowering IV—VI; fruiting V—
VII. (Plate V, Fig. 2).

Dry stony and loamy slopes in foothills, lower mountains and middle
mountain zone,.sometimes as weed in cultivated fields.—Caucasus:
Eastern and southern Transcaucasia, Talysh; Soviet Central Asia:
mountainous Turkmenia, Syr-Darya (Mogoltau), Pamiro-Alai, Tien Shan
(west, up to Karatau). General distribution: Eastern Mediterranean (east),
Balkans-Asia Minor (east), Armenia-Kurdistan, Iran. Described from the
eastern Mediterranean (Syria ?). Type in London or Paris.

Note. C. orientalis var. erivanica Tamamsch., mentioned above, is
distinguished by achenes with reddish pubescence and reddish inside
pappus scales.

Vz

73

2. C. maerocarpa C. Koch in Linnaea, XXIV (1851) 382; Papava
in Zam. Sist. Geogr. Rast. [Tbilisi] 13, 80; Grossh. Opred. Rast. Kavk.
473.—lIe.: Papava, op. cit. 84, Fig. 1—Exs.: Fl. or. ed. W. Siehe, No.
198 (“C. xeranthemoides Desf.”).

Annual. Differs from preceding species in shape and pubescence of
achene of bisexual florets; achenes cuneate (or oblong-turbinate) and
densely covered with long appressed bristles almost entirely, as well as
by shape and size of pappus scales: slightly broader, shorter and abruptly
narrowed (and not very gradually) into aristate cusp. Flowering IV—V];
fruiting V—VII.

Dry slopes in lower mountains and middle mountain zone (in Asia
Minor to 1,600 m).—Caucasus: Southern Transcaucasia (the only locality
between the villages of Shorbulag and Avdalar near Erevan). General
distribution: Balkans-Asia Minor (two localities in Asia Minor; in
Lycaonia and Cappadocia), ? Armenia-Kurdistan. Described from
Armenia or Asia Minor (place of collection not indicated by the
author, apparently unknown to him) from the specimens collected by
Hundelsheimer, a companion of Tournefort in his famous trip to the
Orient in 1700-1702. The type specimen was in Berlin (?).

Note. This species was not recognized by Boissier (Boiss. Fl. or.
III, 446), in his time nor by other botanists subsequently, who considered
it simply a synonym of C. orientailis (L.) Ktze. (C. xeranthemoides
Desf.).

Recently (1947), however, V.I. Papava (op. cit.) reported C.
macrocarpa C. Koch from the USSR and presented a series of arguments
in favor of restoring it as a separate species. Papava studied two
herbarium specimens from: 1) the vicinity of Erevan (S. Tamamschian,
1931); and 2) Asia Minor (the above cited No. 198, W. Siehe, 1912).

I saw a duplicate of the second of these specimens (duplicates of it
apparently are widely distributed), and based on a study of the achenes,
the differences between the two species, given above in the key, as well
as in the description of C. macrocarpa C. Koch, were formulated. As
can be seen, these differences are generally similar to those indicated
by Papava. Further study, especially of these characters, in all of the
abundant herbarium material of the genus Chardinia Desf. preserved in
the Herbarium of V.L. Komarov Botanical Institute, Academy of Sciences
of the USSR yielded very interesting results.

First, it was found that among the large number of specimens
examined from Asia Minor, many of which (or more precisely their
duplicates) are cited by Bossier (op. cit.), none was found besides the
already mentioned No. 198, W. S:ehe with the characters of C.
macrocarpa C. Koch; thus, the conclusion of V.I. Papava concerning
the distribution of the latter species mainly in Asia Minor has not yet

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74

been confirmed. Typical C. macrocarpa C. Koch (as might be visualized
on the basis of Koch’s diagnosis and W. Siehe’s ‘Herbarium specimens)
also was not found among the abundant material from the Caucasus and
Soviet Central Asia. On the contrary, the mass examination of this
material revealed that the pubescence of the achenes of C. orientalis
(L.) Ktze. varies almost to the degree (cf. description of the achene of
the bisexual floret in the description of this species) that the long
appressed pubescence of the achene of C. macrocarpa C. Koch, nearly
throughout its length, possibly could be considered as one of the extreme
manifestations of the intraspecific variation of C. orientalis (L.) Ktze.
and that, perhaps, Boissier, indeed, was not very correct in combining
these species. Incidentally, it seems to me still somewhat early to make
a final decision now: the collection and critical study of new material
are necessary, and for this reason it is more convenient, practically, to
consider the forms of interest to us as separate species, for now.

Correct typification is of great importance; so far it appears to be
inadequate in both cases. Because after the Linnaean basionym C.
orientalis (L.) Ktze., that is to say Xeranthemum y. orientale L., the
Tournefort polynomial “Xeranthemum orientale, fructu maximi,” was
listed in the synonymy, there is reason to believe that the specimen so
named (i.e., the type) from Tournefort’s Oriental collection may be
located in London, in the Linnaen Herbarium, or, besides that, in Paris,
where the main Herbarium of Tournefort is preserved.

As indicated by Koch at the end of the diagnosis of C. macrocarpa
C. Koch, the specimen from the Hundelsheimer Herbarium (which should
be treated as the type specimen) was designated as “Xeranthemum
armenum, fructu maximo,” which speaks to the probability of this
specimen originating in particular from Armenia and to its similarity
(which also could be suspect) with the type of the previous species. In
the event this specimen is lost or impossible to find in Hundelsheimer’s
Herbarium, it would be appropriate to select a neotype, which could be
collected (together with a sufficient number of duplicates for distribution
to the largest herbaria of the world) from the vicinity of Erevan.

GENUS 1573. Amphoricarpus Vis." *

Vis Fl. dalmat. II (1847) 27 (“Amphoricarpos”); Benth. and Hook.
f. Gen. pl. II (1873) 465; O. Hoffm. in Pflanzenfam. IV, 5 (1892) 316.—
Amphorocarpus Post and Ktze. Lexic. gen. Phaner. (1903) 26.

'Treatment by I.A. Linczevsky.
2From the Greek words amphoreus—an obovate pot with two handles above, and
carpus—fruits; based on the shape of achene of the pistillate florets of the type species.

73 Plate V.

1—Siebera pungens (Lam.) DC., whole plant; achene with pappus; 2—Chardinia orientalis
(L.) Ktze., whole plant; achene of pistillate floret; achene of bisexual floret (with pappus).

iS

76

Capitula many-flowered, small (up to 15-20 mm wide), initially
oblong, later campanulate or hemispherical, solitary, terminal. Involucre
many-rowed, imbricate;.involucral bracts appressed, stiff-herbaceous, with
narrow membranous border; outer bracts gradually reduced outward;
inner-most bracts slightly longer, erect. Receptacle flat, densely covered
with stiff and narrow, undivided or more or less deeply split scales.
Florets variable; outer florets pistillate, few (in one row), fertile; disk
florets bisexual, fertile. Corolla tubular, regular, with more or less long
five-fid limb cleft up to half its length. Anthers sagittate at base, long
caudate-fimbriate; filaments free, glabrous. Style branches in bisexual
florets very short. Achenes of pistillate florets oblong-obovoid, strongly
compressed, with numerous veins and two entire, narrow, lateral wings,
rounded or corniculate above, somewhat flat on outer side, convex and
carinate on inner side. Achenes of bisexual florets obconical, cylindrical-
angular, slightly compressed and curved. Pappus in both types of achenes
one-rowed, usually of numerous, narrow, almost setaceous or narrow-
linear and very gradually almost setose-acuminate, finely serrate-ciliate
or barbed scales, fused at base (for varying distance).

Perennial, not spiny, white pubescent-hairy herbs with alternate,
lanceolate or linear-lanceolate, entire leaves.

The genus comprises two species, one of which grows in the
Caucasus and the other in the Balkan Peninsula.

Type species: Amphoricarpus neumayeranus (Vis.) Vis. (Jurinea
neumayeriana Vis.)

Section 1. Chodatella Alb. in Bull. Herb. Boiss. II (1894) 249.—
Barbeya Alb. in Zap. Kavk. Otd. RGO, XVI (1894) 143 p. gen., non
Schweinfurth (1891).—Alboviodoxa Woron. ex Grossh. in Opred. Rast.
Kavk. (1949) 473 p. gen.—Achenes of pistillate florets with wings
rounded above (without corniculate projections). Pappus scales narrow,
almost setaceous, numerous (30-40). Stem leafy almost to tip.

1. A. elegans Alb. in Bull. Herb. Boiss. II (1894) 247 and Prodr. fl.
Colch. (1895) 141; Grossh. Fl. Kavk. IV, 163.—A. kuznetzowi Winkl.
ex Lipsky in Tr. Peterb. Bot. Sada, XIV (1898) 284 and in Fl. Kavk.
(1899) 355, nomen.—Kusnetzovia caucasica Winkl. in sched. and ex
Lipsky, op. cit. (1898) 284, nomen, p. syn.—Alboviodoxa elegans (A\lb.)
Woron. in sched. (1931) and ex Grossh. Opred. Rast. Kavk. 473; Kolak.
Fl. Abkhazii, IV, 259 (“Albowiodoxa’’); Papava in Fl. Gruzii, VIII, 435.—
Ic.: Alb. in Bull. Herb. Boiss. II (1894) tab. 7; Kolak. op. cit. Plate
XXVII; Papava, op. cit. 433, Fig. 406.—Exs.: GRF No. 3248.

Perennial. Plants 15—40 cm high; rhizome rather thick, woody,
modified into more or less multicapitate caudex above, stems 5—10,
almost erect, simple or sometimes with fewer short branches above,

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V7.

densely feafy almost to tip, white-tomentose. Leaves light green above,
almost glabrous (slightly arachnoid-hairy), silvery-white tomentose
beneath, sessile, oblong-lanceolate, or lanceolate, 4-7 cm long and 8—
12(15) mm wide, very gradually acuminate above; middle leaves often
subobtuse at base; uppermost leaves greatly reduced. Capitula with 30—
60 florets, oblong when young, enlarged at fruiting, campanulate or
hemispherical, up to 15—20 mm wide. Involucral bracts light green or
purple above; outermost bracts small, white-tomentose, deltoid-lanceolate
or oblong-elliptical; middle bracts much larger, slightly arachnoid-
hairy and crimped-ciliate along margin to almost glabrous, broad
ovate-elliptical, mucronate; inner bracts glabrous, glossy, with wide
membranous margin, broad-lanceolate to linear, acuminate; receptacular
scales as long as florets, linear, undivided or slightly lacerate, 10—12
mm long and 0.5—0.8 mm wide. Corolla pale pink. Achenes of pistillate
florets with short and sparse appressed pubescence, about 5 mm long
and 2 mm wide, with 8-9 mm long pappus; achenes of bisexual florets
with short and dense appressed pubescence, 4-5 mm long and about |
mm wide, with 10-11 mm long pappus. Flowering VII—VIII; fruiting
VIII-IX. (Plate VI, Fig. 1).

Limestone rocks and stony slopes in alpine zone, at 1,900—2,300
m.—Caucasus: Western Transcaucasia (limestone ranges of Abkhazia
and western Georgia, from the Bzyb River Basin to the Inguri River
Basin). Endemic. Described from the Khechigvar Valley near the
Chipshiram Mountain in the Bzyb Range and from the Migariya
Mountain above the village of Kurdtsu in Megrelia. Type in Geneva,
isotype and paratypes preserved in Leningrad.

Note. A. elegans Alb. belongs to the group of “limestone endemics”
described in rather large numbers by Albov and other botanists from
the mountains of the western Caucasus.

As can be seen from the list of synonyms given above, different
viewpoints have existed in relation to the taxonomic position of A.
elegans.

Albov, having described this plant at first as a separate genus,
Barbeya Alb., in the same year, 1894, reduced it to the rank of a section
of genus Amphoricarpus Vis. A.A. Grossheim, in agreement with Albov,
accepted this species in Amphoricarpus, but subsequently (op. cit. 1949)
considered it necessary to validate the generic name Alboviodoxa Woron.,
until then unpublished and appearing on an herbarium label (Herb. FI.
Cauc. No. 499, ined.), and the combination of A. elegans (Alb.) Woron.
A.A. Kolakovsky (op. cit. 1949) agreed with Grossheim, but a little
later, at the end of 1949, issued the species in question in the “Herbarium
of the Russian Flora” (under No. 3248 cited above) as Amphoricarpus
elegans Alb. Papava (op. cit.) included A. elegans in Albovidoxa Woron.

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78

It may also be noted that this plant was already known to Winkler in
1890 from collections of Kuznetsov and was named by the former (in
sched. ad herb. et ic.) as a new genus and species—Kusnetzowia
caucasica Winkl.

My critical consideration of this question, taken up in view of the
obvious need, led to the following conclusions.

1. In publishing the generic name Alboviodoxa Woron. ex Grossh.
(which, as was mentioned above, should have replaced the homonym
Barbeya Alb.), A.A. Grossheim did not indicate any difference from
the genus Amphoricarpus Vis.

2. U.N. Woronow, ignoring the Latin aeaetintin of the genus
Alboviodoxa cited above on the herbarium label prepared for issue,
distinguished it from Amphericarpus by essentially the same characters
for which Albov established section Chodatella Alb. (see above), namely:
a) all scales of receptacle entire (not mixed, entire and often lacerate);
b) achenes of pistillate florets with apically rounded lateral wings (and
not corniculate); and c) pappus scales setaceous, finely toothed-scabrous
(and not linear, indistinctly finely serrate-ciliate).

3. Further study of the herbarium material showed that: a) the
receptacular scales in A. elegans are definitely not always entire, and
slightly lacerated scales are very common among them; b) the achene
of pistillate florets in A. elegans actually always has rounded (not
corniculate) wings, but it is extremely similar in its other morphological
details to the analogous achene of A. neumayeranus; and c) the pappus
scales on the achenes of A. elegans are actually always narrow, almost
setaceous, about 0.1 mm wide, numerous (30-40) and almost in a single
row in the achenes of pistillate and bisexual florets, whereas the pappus
scales of pistillate florets in A. neumayeranus are always broader,
narrowly linear, 0.2-0.3 mm wide, and much less numerous (four to
eight), while those of bisexual florets vary (in different specimens) in
width from 0.1 to 0.3 mm, i.e., from almost setaceous to narrowly linear
and somewhat more numerous (12-20), but still many fewer (one-third
to half) in number than in A. elegans; I could not find significant
differences in the nature of pubescence of the pappus scales between
the two types, except for only a slight difference in the density of this
pubescence.

Thus, as the most significant differences between A. elegans and A.
neumayeranus, only the following can be listed: the shape of the upper
part of achenes of pistillate florets (absence or presence of small
corniculate projections, a continuation of lateral wings), the shape (more
precisely, the width) and relative number of the pappus scales, as well
as (in addition to the differences listed earlier) the nature of the foliage
on the stems.

78

56}

All these characters, it appears to me, can hardly be considered as
more than sectional differences, especially considering the extremely
similar achene morphology of the bisexual florets, involucral bracts,
capitula, leaves, and also the entirely similar nature of pubescence on
the whole plant in both species.

Therefore, I can only agree, obviously, with Albov, who established
(op. cit. 1894, 249) two monotypic sections in the genus Amphoricarpus
Vis., one of which includes the Caucasian “limestone endemic” A.
elegans Alb., and the other A. neumayeranus (Vis.) Vis., which grows
on the limestone mountains of the southwestern Balkan Peninsula
(Dalmatia). The unique range discontinuity and ecological specialization
(limestone in both cases!) provided the basis for Albov (ibid. 250) to
include this pair of species among the relicts of the Tertiary Flora. It
may be added that quite a few of the ranges of the Tertiary relict species
listed in the detailed review of V.P. Maleev (Tertiary Relicts in the
Flora of Western Caucasus and Main Stages in the Quaternary History
of the Flora and Vegetation. Materials on the History of the Flora and
Vegetation of the USSR, I, 1941, pp. 61-144) are extremely similar in
the nature of their discontinuity from the range of Amphoricarpus Vis.
species.

GENUS 1574. Carlina L.!': 2

L. Sp. pl. (1753) 828; DC. Prodr. VI (1837) 545 p. p. excl. sect. 3;
Benth. and Hook. f. Gen. pl. II (1873) 465; O. Hoffm. in Pflanzenfam.
IV, 5 (1892) 316.—WMitina Adans. Fam. II (1763) 116.—Athamus Neck.
Elem. I (1790) 85.—Carlowitzia Monech. Meth. Suppl. (1802) 225.—
Chromatolepis Dulac, Fl. Hautes-Pyrén. (1867) 526.

Capitula many-flowered, rather larger to very large (up to 150-200
mm wide), broadly campanulate or almost hemispherical, sessile or
subsessile among basal rosette or leaves or terminal, solitary or nearly
corymbose. Involucre many-rowed; outer involucral bracts wider, leafy,
prickly-toothed, more or less squarrose; middle bracts narrower and
shorter, appressed and imbricate below, cristate-prickly, with furcate
spines; innermost bracts long, much longer than florets, linear, entire,
stiff-scaly and lustrous, whitish or yellow, sometimes purple, usually
divergent. Receptacle flat, fleshy, densely covered with long stiff scales, .
almost as long as florets, more or less deeply incised into thin bristles.
All florets similar, bisexual. Corolla tubular, regular, with cylindrical,

'Treatment by I.A. Linczevsky.
Origin of the name not known precisely.

2

80

narrowly five-fid limb, purple or yellow. Anthers sagittate at base, with
long and narrow caudate processes ciliate along margin; filaments
glabrous. Style branches very short. Achenes oblong, almost cylindrical,
without ribs, densely pubescent, truncate, with one-rowed pappus of
numerous scales, deeply split into long plumose bristles.

Biennial and perennial plants, sometimes stemless herbs, less often
shrubs (Canary Islands) with basal or alternate, toothed or pinnatifid,
lobed or incised leaves with spiny teeth or lobes, less often leaves
undivided, somewhat sparsely ciliate-spiny.

The species of Carlina L. are distributed in the Mediterranean Region
(including the Canary Islands and northern Africa), Western and Eastern
Europe, and in the temperate zone of Asia (east to the Baikal Region).

Type species: Carlina vulgaris L.

Note. Carlina L. is a complex and confusing taxonomic group, most
species of which (as well as a large number of infraspecific taxa) were
described from Western Europe. It is well known that particularly in
such cases the study of the taxonomic composition and the resolution of
the nomenclatural problems are difficult, and to some extent sometimes
simply impossible without a monographic study of the whole genus.
The situation is further complicated by the extreme morphological
variability of Carlina L. species, which was also reflected in pre-
Linnaean botany (16th century) in the form of the name Camaeleon
(= Chamaeleon, which, as is known, has long been used to designate
properties like variability, inconsistency, instability, etc.), by which, for
example, the species that is now called C. acaulis L. was known (as
“Camaeleon niger vulgaris”). Naturally, the present analysis, not based
on some fresh and complete monographic review of the genus, cannot
be critical, and is more a presentation of the traditional understanding
of Russian taxonomists concerning the endemic species of Carlina L.,
with some nomenclatural and taxonomic additions.

1. Plants with well developed, tall (up to 80-120 cm high) stem often
branched above. Leaves entire, more or less deeply spinose-toothed
along margin or thin-spiny; upper cauline leaves surrounding
capitulum, almost as long as capitulum or much shorter. Capitula
25—O0 eM WADE crrcuaen. careless, actodaerts.. asteetnsst. fore teh een! 2

+ Plants without stem, with single capitulum, among basal rosette of
leaves, or with short (up to 20-40 cm long), usually ascending
simple stems bearing solitary capitula. Leaves pinnatilobate to
pinnatisect; upper cauline leaves surrounding capitulum much
longer than it. Capitula 70-150(200) mm wide...................08 4.

2. Innermost involucral bracts light purple .............. 3. C. lanata L.

80

81

+ Innermost involucral bracts light yellow .............ccccceccceeeseeees 3}
3. Leaves more or less deeply spiny-toothed along margin; middle
and upper leaves ovate-lanceolate or ovate; upper leaves
surrounding capitulum, usually much below it, less often almost
as lOnpe..... oF ..4 ce Se. 2 Oe..omeercamee Swe... Hg 1. C. vulgaris L.
+ Leaves finely ciliate- or serrate-spiny along margin; all leaves
lanceolate or linear-lanceolate; upper leaves, surrounding capitulum,
almost jas long; as capitulumy.is4e.ce trons. tales. tue aythen es ces
La datncalh, Soau lta acc .smdeeeats. farads 2. C. biebersteinii Bernh. ex Hornem.
4. Plants without stem. Leaves pinnatilobate, with almost deltoid,
large-toothed lobes, arachnoid-hairy above, tomentose beneath.
Capitula up to 150-200 mm wide. Pappus bristles connate at base
Inferaqups Ol NO—lBperes... celine ea mman suse knee. meen
Soigeelt hudba 4. C. onopordifolia Bess. ex Szafer, Kulcz., B. Pawl.
+ Plants without stem or with short (up to 20-40 cm tall), usually
ascending, simple stem bearing solitary capitulum. Leaves more
or less deeply (sometimes up to midrib) and often irregularly
pinnatisect or pinnatipartite, glabrous or arachnoid-hairy, but never
tomentose. Capitula 70-150 mm wide. Pappus bristles connate at
base,inveroupsiof, five to eiehite, resets ssssrees! .leisedus. Weaesade eae 5:
5. Leaves deeply (up to midrib) and irregularly pinnatisect, with short,
angular, more or less distant lobes, glabrous or subglabrous above,
somewhat arachnoid-hairy beneath. Usually without stem, often
mountain (Carpathians) plants..............0.cccccce 5. C. acaulis L.
+ Leaves less deeply incised, with long and close-set lobes, prickly
and arachnoid-hairy on both surfaces, more dense along petiole
and beneath along midrib. Plants with short (up to 20-40 cm long)
SECIS fee ta sas 8 oe a 2h tea danse hee eC - 6. C. cirsioides Klok.

1. C. vulgaris L. Sp. pl. (1753) 828; M.B. Fl. taur.-cauc. II, 282;
DC. Prodr. VI, 546; Ldb.. Fl. Ross. I], 2, 675 p. p.; Boiss Fl. or. III,
448; Schmalh. Fl. II, 93 p. p.; Fedtsch. and Fler. Fl. 999; Hegi, Ill. FI.
IV, 2, 821 p.p.; Hayek, Prodr. Fl. Balc. (1931) 693; Grossh. Fl. Kavk.
IV, 163; Kolak. Fl. Abkhazii, IV, 260; Klok. in Vizn. Rosl. URSR,
560; Papava in Fl. Gruzii, VIII, 435; Szafer, Kulcz. B. Pawl. Rosl.
Polskie (1953) 708; Kirp. in Majevski, FI., 8th edition, 595, in note;
Nikitin in Fl. Turkm. VII, 156 p. p. excl. descr.; Gaddzhiev in Fl. Azerb.
VII, 359.—C. vulgaris L. var. araneosa K.-Pol. and Preobr. in Izv.
Peterb. Bot. Sada, XIV, 3 (1914) 318; Grossh. Fl. Kavk. IV, 163.—C.
vulgaris L. B. sublanata DC. Prodr. VI (1837) 546 an p. p.; Ldb. FI.
Ross. II, 2, 675; ? C. Koch in Linnaea, XXIV, 384.—C. caulescens
Gilib. Fl. Lithuan. III (1781) 195 (non vidi)—C. acaulis auct. non L.:
Ldb. Fl. Ross. II, 2, 674 p. p. quoad specim, osil—C. Janata auct. non

82

82

L.: Habl. Pall. Georgi sec. Ldb.: op. cit. 675.—Ic.: Fedtsch. and Fler.
op. cit. Fig. 1000; Hegi, op. cit. fig. 524, 526, 527, tab. color, 269, fig.
4; Dostal, Kli¢. Kvét. CSR (1954) fig. 2515.—Exs.: Fl. cauc. exs. No.
174; Fl. exs. reip. Boh.—Slov. Nos. 285, 962; Fl. exs. austro-hung. No.
3765; Billot, Fl. Gall. et Germ. exs. No. 1508.

Biennial. Plants 20-80 (125) cm high. Stem erect, simple or with
few branches above, more or less densely arachnoid-pubescent. Leaves
entire, oblong-lanceolate and lanceolate (basal and lower cauline leaves)
to ovate-lanceolate and ovate (middle and upper cauline leaves), 2—
10(15) cm long and 0.5—2 cm wide, glabrous above, more or less densely
arachnoid-hairy beneath, usually deeply spinose-toothed along margin;
basal leaves petiolate; cauline leaves sessile, gradually reduced above,
uppermost leaves surrounding capitulum, usually much below it, less
often almost at level. Capitula 30-45(50) mm wide, usually fewer (one
to five), terminal on stem and branches, almost corymbose. Outer
involucral bracts green, leafy, oblong-lanceolate, arachnoid-hairy
(sometimes rather densely), toothed-spinose along margin; middle bracts
brownish or dark purple, along margin cristate- and branched-spinulose
(with branched spinules); innermost bracts light green, stiff-scaly and

-lustrous, linear, much longer than middle bracts, radially divergent; inner

bracts yellowish-purple. Achenes 2-4 mm long; pappus two to four times
as long as achenes, up to 8-9 mm long, bristles connate at base in
groups of two to three. Flowering VI-IX.

In taiga (south), forest and forest-steppe zones, dry meadows, dry
slopes, open forests, on forest glades, in scrub, along forest fringes; in
southern regions (the Caucasus and Soviet Central Asia) found in
mountains, from forest to subalpine zone; sometimes on deposits,
waste lands and along roads.—European Part: Almost all regions,
approximately from 62°N (in the west) and 60°N (in the east, in the
Urals) to Caucasus, Crimea, and foothills of the Carpathians; not found
in semidesert zone; more typical forms found in the west (see note).
Caucasus: All regions. Soviet Central Asia: mountainous Turkmenia
(western Kopetdag). General distribution: Scandinavia (south,
approximately up to 62°N), Central and Atlantic Europe, western
Mediterranean (north), Balkans-Asia Minor (north). Described from
Western Europe. Type in London.

Note. Judging from the sufficiently large collection preserved in the
Herbarium of the Botanical Institute, Academy of Sciences of the USSR,
the more typical form of C. vulgaris L. (with a shorter and wider toothed-
incised and stiff spiny basal and cauline leaves, the uppermost of which
are shorter than the capitulum) within the limits of the European part of
the USSR is more frequently found in the west (in the Baltic Region,
Ladoga-IImen, partly along Upper Dnieper). At the same time, more or

83

ZA ss

)
~ } ©, (1.
.N Ms
\ A N Ae {Ry h J “<4; U2 me
ISNA VEEN CS Y (EE y
<{[ RRA ane
= SS Sind SD ,

81 Plate VI.

i—Amphoricarpus elegans Alb., whole plant; achene of bisexual floret;
2—Carlina acaulis L., whole plant; achene with pappus.

83

84

less abundant collections of the same form are available from almost all
regions of the European part of the USSR, which by its characters (long
basal and lower cauline leaves, but short and stiff-spinose upper leaves)
appears to be intermediate between C. vulgaris L. and its closely related
species C. biebersteinii Bernh., which is also widely distributed in the
European part of the USSR. In the past, this form was described as C.
intermedia Schur (Schur, Enum. pl. Transsilv. 1866, 413), but some
authors subsequently considered it as a subspecies, C. vulgaris L. ssp.
intermedia (Schur) Hayek (op. cit. 694), or variety, C. vulgaris L. var.
intermedia (Schur) Lindb. f. (Haralo Lindberg in Meddel. Soc. pro fauna
et flora fennica, 26, 1900, 70). Identification of this form is not easy;
the material for a precise understanding of its distribution is still
insufficient.

2. C. biebersteinii Bernh. ex Hornem. Hort. Hafn. Suppl. (1819)
94; Kryl. Fl. Zap. Sib. XI, 2873; Kirp. in Majevski, Fl. 8th edition,
595; Popov, FI. Sr. Sib. II, 750.—? C. longicaulis Somm. and Lev. in
Nuov. Giorn. bot. Ital. 2 (1895) 91 and in Tr. Peterb. Bot. Sada, XVI,
250.—C. longifolia Rchb. Pl. Crit. VIII (1830) 25, tab. 761, non. Viv.
(1824); Lindb. f. in Meddel. Soc. pro fauna et flora fennica, 26: (190C)
69; Iljin in Fl. Yugo-Vost. VI, 378; Grossh. Opred. Rast. Kavk., 473;
Popov, Ocherk Rast. i Fl. Karpat., 251; Klok. in Vizn. Rosl. URSR,
560; Szafer, Kulcz. B. Pawl. Rosl. Polskie, 709.—C. longifolia Rchb.

_ var. B. pontica Boiss. FI. or. III (1875) 448; Grossh. Fl. Kavk. IV, 163;

Papava in FI. Gruzi, VII, 436—C. vulgaris L. var. longifolia Korsh.
Tent. Fl. Ross. or. (1898) 234; Fedtsch. and Fler. Fl. Europ. Ross. 999.—
C. vulgaris L. y. microcephala Ldb. Fl. Ross. I, 2 (1845-1846) 675 an
p. p-—C. acaulis auct. non L.; Ldb. Fl. Ross. II, 2, 674 p. p. quoad
specim. altaica——C. nebrodensis auct. non Guss.: Ldb. Fl. Ross. II, 2,
675; Kryl. Fl. Alt. III (1904) 684 (pro C. vulgaris L. var. nebrodensis
Kryl.); Pavlov, Fl. Tsentr. Kazakhst. III, 292.—? C. racemosa auct. non
L.: Gilib. sec. Ldb. Fl. Ross. II, 2 (1845-1846) 675; Georgi, sec. Ldb.
l.c—C. vulgaris auct. non L.: Ldb. Fl. alt. TV (1833) 13; Kar. and Kir,
Enum. pl. alt. No. 507; Ldb. Fl. Ross. II, 2 675 p. p.; Turcz. in Bull.
Soc. Nat. Mosc. XX, 3, 49 (Fl. baic.-dahur. No. 672); Schmalh. FI. II,
93 p. p.; O. and B. Fedtsch. Perech. Rast. Turkm. IV, 237; Pavlov, op.
cit. 292.—Ie.: Ijin, op. cit. Fig. 695; Klok. op. cit. Fig. 296; Kirp. op.
cit. Fig. 224.—Exs.: Wolosz, Fl. polon, exs. No. 541; Fl. exs. austro-
hung. No. 3763.

Biennial. Plants 20—80(120) cm high. Stem erect, simple or slightly
branched above, usually weakly arachnoid-pubescent. Leaves undivided,
lanceolate or linear-lanceolate, 2—10(20) cm long and 0.5—2 cm wide,
glabrous or subglabrous above, arachnoid-hairy beneath, margin finely

85

and quite often (almost ciliate or serrate) spinulose; basal leaves petiolate;
cauline leaves sessile, gradually reduced upward, uppermost leaves,
surrounding capitulum, more or less as long as capitulum. Capitula
(25)35-40(60) mm wide, usually fewer (one to five), terminal on stem
and branches almost corymbose. Outer involucral bracts. green, leafy,
linear-lanceolate, arachnoid-hairy, margin ciliate-spinulose; middle bracts
brownish or dark purple, margin cristate and branched-spinulose (with
branched spinules); uppermost bracts light yellow, stiff-scaly and lustrous,
linear, much longer than middle bracts, divergent. Corolla dark or yellow
purple. Achenes 2-4 mm long; pappus two to four times as long as
achene, up to 8-9 mm long, its bristles connate in groups of two or
three at base. Flowering VI-IX.

In taiga (south), forest and forest-steppe zones, in dry meadows,
dry slopes, open (pine, mixed, birch) forests, forest glades, among
shrubs, along borders; in Western Siberia, often in birch fellings;
in Central Siberia, according to Popov. (op. cit.), “steppe plant, some-
what of a weed”; in western Caucasus—in forest zone; sometimes on
deposits, wastelands, near roads—European Part: Almost all regions,
approximately from 62° to 60°N toward south; not found in semidesert
zones; Caucasus: Western Transcaucasia; Western Siberia: Ob (south,
up to 56—58'N), Upper Tobol, Irtysh, Altai; Eastern Siberia: Angara-
Sayans. General distribution: Scandinavia (south), Central and Atlantic’
Europe, western Mediterranean (north), Balkans-Asia Minor (north).
Described from cultivated specimens in the Copenhagen Botanical
Garden. Type in Copenhagen.

3. C. lanata L. Sp. pl. (1753) 828; DC. Prodr. VI, 546; incl. B.
monocephala DC. and y. prolifera DC.; Ldb. Fl. Ross. II, 2, 676; Vis.
Fl. Dalm. II, 29; Boiss. Fl. or. III, 451; Nym. Consp. Fl. Eur. 400;
Hayek, Prodr. Fl. Balc. 696.—Ic.: Sibth. and Sm. Fl. Graeca, tab. 836.—
Exs.: Balansa, Pl. d’Alger, 1852, No. 502; Billot, Fl. Gall. et Germ.
exs. No. 2290; Bornm. It. syr. No. 883.

Biennial or annual (?). Plants 10—20(40) cm tall. Stem erect, simple
or almost cymosely branched (with central almost sessile capitulum and
lateral capitula surrounding it on peduncles), densely long arachnoid-
lanate. Leaves hard coriaceous, convolute, lanceolate or broad-lanceolate,
3-10 cm long and 1—1.5(2) cm broad, more or less densely arachnoid-
tomentose, margin shallow crenate-dentate, with short teeth, each with
1-2 or 3 spinules; basal leaves petiolate, cauline sessile, semiamplexicaul;
uppermost leaves crowded, subtending capitulum, and usually equal to
it or slightly longer. Capitula 25—40(60) mm across, few (often 1-3),
solitary, terminal on stem and branches. Outer involucral bracts leafy,

84 lanceolate, spinulose-dentate, longer than innermost; middle bracts linear-

86

lanceolate with simple or (on lower side) branched spinules; innermost
bracts bright purple, stiff-scaly and lustrous, linear, acute, much longer
than middle ones, longer than florets by 6-8 mm, outwardly divergent.
Corolla purple. Achene about 3 mm long; pappus two and one-half to
three times as long as achene, up to 9-10 mm long, its bristles connate
below in groups of three to five.

The conditions of its growth in the Caucasus are not known; it
probably grows as a weed, because it is reported by Boissier (Boiss. op.
cit.) for the neighboring territories (Anatolia, Syria, Greece) growing on
fallow or waste lands, and along roads.—Caucasus: Western
Transcaucasia (the only locality in the vicinity of Batumi known since
the times of Ledebour; most probably not collected later). General
distribution: Mediterranean (west and east), Balkans-Asia Minor.
Described from Southern Europe (Italy and southern France). Type in
London.

Note. Some authors consider this species as an annual, others a
biennial plant. It perhaps belongs to the category of the so-called
“perennating” plants (i.e., usually annual, but in some years under suitable
weather conditions behaving as biennial or perennial) that are very
commonly found in the Mediterranean Region.

4. C. onopordifolia Bess. ex Szafer, Kulcz. B. Pawl. Rosl. Polskie
(1924) 641; DC. Prodr. VI, 545, nomen. p. sun.: Nym. Consp. fl. Eur.
II, 400; nomen p. syn.; Popov, Ocherk Rast. i Fl. Karpat., 251, in note;
Klok. in Vizn. Rosl. URSR, 560 and in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVI, 356, 357, in note; A. Jas. 1 B. Pawl. in Fragm. Florist.
et Geobot. [Krakow] II, 2, 12; Shelest in Ukr. Bot. Zhurn., XIII, 4,
45.—C. acanthifolia auct. non All.: Bess. Enum. pl. Volh. (1822) 32
(No. 1057); Boiss. Fl. or. III, 448 p. p. quoad pl. ross—C. acanthifolia
[var.] B. cynara auct. non All. nec. DC.; Ldb. Fl. Ross. II, 2, (1845—
1846) 674; Schmalh, FI. II, 93 p. p. quoad pl. ross.; Fedtsch. and Fler.
Fl. Evrop. Ross. 1000.—Ie.: Kozlowska in Mém. Acad. Polon. Sc. et
Lettr. sér. B. III (1931) tab. 5, fig. 9 (non vidi); A. Jas. and B. Pawl.
op. cit. fig. phot. 1-6.

Perennial. Without stem. Leaves in basal rosette; outer leaves
petiolate, inner sessile; all leaves pinnatilobate, with almost deltoid,
spinose, large-toothed lobes; arachnoid-hairy above, tomentose beneath,
5—15(50) cm long and 1.5—7 cm wide. Capitulum large, up to 150-200
mm wide when fully open, solitary among rosette of leaves. Outer
involucral bracts glabrous above, spatulately broadened and pointed;
middle bracts brown to deep brown, almost linear, both cristate-branched
spinulose along margin; outermost bracts light yellow, stiff-scaly and
lustrous, linear, slightly broadened above base and gradually acuminate

85

87

beyond, much longer than outer bracts, 30-35 mm longer than florets,
erect or radially divergent. Corolla yellow (?). Achenes 5—6 mm long;
pappus three to four times as long as achenes, up to 20 mm long, its
bristles connate at base in groups of 16-18. Flowering VIII-IX.

Dry south slopes, in shrubby outgrowths.—European Part: Upper
Dnieper (extreme southwest Kremenets), Middle Dnieper (western bank
of Dnieper-Vinnitsa, Kamenets-Podolskii, Balta), Upper Dniester (north,
submountainous part—Rogatin, Zolochev, Belyi Kamen), Bessarabia
(Rashkov on Dniester, after Schmalhausen). General distribution: Central
Europe (southern part of Poland-Malopolska and Lubelska hills).
Described from western Ukraine and southern Poland. See Note |
regarding the type specimen.

Note 1. The name C. onopordifolia Bess. was first published by De
Candolle (DC. op. cit.) as a synonym of another species—C. acanthifolia
All. B. cynara DC., with a reference to the herbarium specimen of Besser
1831 (“in Podolia prope Winnica”). This name (as a synonym) was not
valid (according to Article 37 of the International Code of Botanical
Nomenclature). The first description of C. onopordifolia Bess. based on
the material collected in Ukraine was published in 1897 by Schmalhausen
(op. cit.) but again under the same name as in the case of De Candolle.
As far as can be determined, the name C. onopordifolia Bess. was first
accepted and provided with a description by the Polish botanists Szafer,
Kulczynski and B. Pawlowski (I.c.) in 1924, which should, therefore,
also be considered as the date of valid publication of this name. Because
the type specimen of the species was not designated in this publication,
it would be necessary (in accordance with Article 7, Note 3 of the
International Code) to select a lectotype later from the material cited in
the description, i.e., from one of the four localities mentioned here. As
far as is known to me, the Polish botanists still have not published a
lectotype selection for this species.

Note 2. C. onopordifolia Bess. ex Szafer, Kulczynski and B.
Pawlowski, according to all authors, is an extremely rare species,
representing the northeastern (North Cis-Carpathian) race from the
Mediterranean complex (or series) of C. acanthifolia All. In spite of the
very close affinity of these two species, some differences of C.
onopordifolia are quite sharp. We point out, for example, that the pappus
bristles in C. onopordifolia are connate at the base in groups of 16—18,
whereas in C. acanthifolia they are in groups of 9-10; besides of course,
it is desirable to compare these species on more extensive material than
we had.

The range of C. onopordifolia is restricted to the Volyno-Podolsk
hills of southwestern Ukraine and in the west, in Poland, to the adjoining
Lubelska and Malopolsk uplands, but the species is found only at a few

86

88

places and in small numbers. The Polish botanists, who have long been
interested in this species, confirmed by 1956 three localities in Poland,
which were all declared State Reserves; it may be noted that in Poland,
besides C. onopordifolia, C. acaulis L. is also protected by the State.

The above-cited article of A. Jasiewich and B. Pawlowski (l.c.),
reporting the discovery of a new (third) locality C. onopordifolia in
1954, provided much interesting information on the biology and ecology
of this species. It has been established, for example, that C. onopordifolia
should be considered a perennial and not a biennial-triennial, although
isolated specimens may likely grow faster. There have been reports of a
small number of specimens flowering simultaneously, for example, in
the third locality mentioned above (slopes of the limestone-chalky Waly
in Krakow Province), where more than 20,000 specimens have been
found (very fewer in other habitats); only about 60(!) flowering plants
were found in 1954. A very interesting new hybrid has been found at
the same habitat: C. onopordifolia Bess. ex Szafer, Kulcz. and B. Pawl.
x C. acaulis L.: x C. szaferi A. Jas. and B. Pawl. (op. cit., p. 19).

5. C. acaulis L. Sp. pl. (1753) 828; Szafer, Kulcz. B. Pawl. Ros.
Polskie (1924) 641 and (1953) 708; Hegi, Ill. Fl. VI, 2, 817; Klok. in
Vizn. Rosl. URSR, 559 and Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVI, 356, 357, in note.—C. acaulis [var.] B. caulescens (Lam.)
Schmalh. Fl. I (1897) 93 p. p. quoad nomen; Hegi, Ill. Fl. VI, 2, 819;
Popov, Ocherk Rast. 1 Fl. Karpat. 251—C. caulescens Lam. FI. Fr. II
(1795) 7.—Ic.: Fedtsch. and Fler. Fl. Evrop. Ross., Fig. 1001; Hegi, op.
cit. figs. 519-523.—Exs.: Fl. exs. reip. Bohem.—Solv. No. 284; F.
Schultz, Herb. norm. Nos. 87, 87 bis; Wolosz. Fl. polon. exs. No. 956.
Callier, Fl. siles. exs. Nos. 621, 622.

Perennial. Without stem. All leaves in basal rosette petiolate, all
irregularly and deeply (sometimes up to midrib) pinnatisect, with short,
angular, spiny-toothed or distantly lobed, glabrous or subglabrous above,
weakly arachnoid-hairy beneath, 4—12(25) cm long and 1.5—4(6) cm
wide. Capitulum large, 70-150 mm wide when open, solitary among
rosette of leaves. Outer involucral bracts herbaceous, green; middle bracts
brownish to brown, cristate and sinuate-spiny along margin; innermost
bracts whitish above, with wide dark purple stripe below on outerside,
stiff-scaly and lustrous, linear, acuminate, not or slightly broadened
above, much longer than outer bracts, exceeding florets by 25 mm,
erect or radially divergent. Corolla whitish or reddish. Achenes 4-6 mm
long; pappus three to four times as long as achenes, up to 15-18 mm
long, its bristles connate at base in groups of five to eight. Flowering
VIUI-IX (Plate VI, Fig. 2).

Meadows of forest and subalpine zone, on mountain slopes, forest
glades.—European Part: Upper Dnieper (northwest region of Grodno),

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89

upper Dniester (Carpathians and Cis-Carpathian Region). General
distribution: Northern Europe, Atlantic Europe (south), western
Mediterranean (north), Balkans-Asia Minor (west). Described from
Europe (Italy and Germany). Type in London.

Note. Besides the typical acaulescent form of C. acaulis L., the
short-stem form, C. acaulis L. var. caulescens (Lam.) Schmalh., is also
found in the Carpathian and Cis-Carpathian Region. It differs mainly in
the presence of a short, 10—20(40?) cm high, ascending, leafy stem and
slightly larger leaves than in the acaulescent form. According to M.V.
Klokov (op. cit. 1954, 356), “there apparently is no taxonomic
significance” to this form.

6. C. cirsioides Klok. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVI (1954) 355 and Vizn. Rosl. URSR (1950) 560, nomen et
descr. breviss. ucr.—C. acaulis auct. non L. : Ldb. Fl. Ross. II, 2 (1845—
1846) 674 p. p.; Schmalh. FI. II, 93; Fedtsch. and Fler. Fl. Evrop. Ross.,
999.—C. acaulis [var.] B. caulescens auct. non L. nec Schmalh.:
Schmalh. Fl. II, 93; Fedtsch. and Fler. Fl. 1000.—C. simplex auct. non
Waldst. and Kit.: Bess. Enum pl. Volh. (1822) 32 (No. 1052); Ldb. FI.
Ross. Il, 2, 674.—Exs.: GRF No. 1523 (“C. acaulis L. var. simplex
Waldst. and Kit.”).

Biennial. Plants 20-40 cm high. Stem ascending or almost erect,
simple, sparsely floccose-arachnoid-hairy. Leaves petiolate, pinnatisect
or lobed, with oblong, palmati- or short-pinnatifid, spiny-toothed, more
or less close-set, floccose-arachnoid-hairy lobes, 5-45 cm long and 1.5—
12 cm wide. Capitulum large, open, 70-100 mm wide when open,
solitary, terminal surrounded by somewhat more dense cluster of leaves.
Outer involucral bracts leafy; middle bracts brownish, cristate and
branched-spinose along margin; innermost bracts whitish above, with
wide dark purple stripe on outer surface, stiff-scaly and lustrous, linear,
slightly broadened above base and gradually narrowed beyond, much
longer than outer bracts, exceeding florets by 25 mm, erect or radially
divergent. Corolla yellowish brown. Achenes about 3 mm long; pappus
three to four times as long as achene, up to 12-14 mm long, its bristles
connate at base in groups of five to eight. Flowering VII-IX.

Open forests, dry meadows, along steppe slopes.——European Part:
Upper and Middle Dnieper (right bank of Dnieper. Upper Dniester
(Carpathians)). General distribution: Central Europe (southeastern Poland-
Lubelska Hills). Described from the vicinity of Belichi in Kiev Province,
from the above-cited specimen No. 1523 “Herbarium of Russian Flora”
(GRF). Type in Leningrad; isotypes in many places.

Note. Besides on the plains and low hummocky-undulating elevations
along the right bank of the Dnieper, this species quite apparently occurs

88

90

rarely at rather high altitudes in the Carpathians. For example, K.N.
Igoshina collected a unique specimen on September 11, 1948, from the
southern slopes of Mt. Petros in the Carpathians at an altitude of 1600
m, which can be identified as C. cirsioides Klok. without any difficulty
on the basis of the distinct pubescence, as well as the shape and size of
the leaf lobes, and is not at all similar to the specimens of C. acaulis L.
collected from the same region.

GENUS 1574a. Thevenotia DC.

DC. in Guill. Archiv. Bot. II (1883) 331.

Capitula homogamous; florets tubular, bisexual. Outer involucre
leafy, inner imbricate; involucral bracts plumosely spinescent.
Receptacular scales incised. Filaments free; anthers appendages plumose.
Achenes sericeous; pappus one-rowed with sericeous bristles connate at
base. Annual herbs.

Besides our native species there is one more species in Iran.

Type species: 7. persica DC.

1. T. scabra Boiss. Fl. or. III] (1875) 455; Czerm. in FI. Uzb. VI
(1962) 228, Plate XXIII._—Atractylis scabra Boiss. Diagn. pl. or. ser. I,
6 (1845) 109.

Annual. Stem 6—12 cm high, glabrous, with solitary spines, leafy,
dichotomously branched. Leaves coriaceous, finely arachnoid-hairy,
sinuate-spinosely toothed. Capitula almost sessile, solitary or in groups
of twos or threes, ovoid, 10-12 mm wide. Inner involucral bracts with
dark-colored cusp. Florets purple; achenes oblong. Flowering VII-—VIII.

High deserts.—Soviet Central Asia: Tien Shan, mountainous
Turkmenia. General distribution: Iran. Described from Iran. Type in
Geneva.

GENUS 1575. Atractylodes DC.':?

DC. Prodr. VII (1838) 48; Fl. Koidz. Fl. Symb. Or.-As. (1930) 3;
Kitam. Comp. Japon. I (1937) 18.—Giraldia baroni in Nuov. Giorn.
bot. Ital. 4 (1897) 431, tab. 12.

Capitula homogamous, bisexual, with solitary florets, all fertile.
Involucre almost cylindrical, involucral bracts many-rowed; outer bracts

'Treatment by E.G. Bobrov.

*Named on the basis of similarity with the Mediterranean genus Atractylis. The name
originates from the Greek word atractos—spindle; possibly named on the basis of fusiform
root.

89

91

surrounding capitulum very large, long-pinnate; inner bracts—involucral
bracts proper—much smaller, woolly along margin and above. Receptacle
concave, scaly. Corolla tubular, five-fid into elliptical and weakly
divergent lobes. Anthers sagittate at base. Style cylindrical; stigma
bilobed with obtuse lobes, pubescent on outer surface. Achenes
compressed, oblong-ovate iustrous, pubescent, covered with plumose
bristles.

Perennial. Rhizomatous herbs, almost semishrubs in northeastern
China. Leaves alternate, glabrous, usually petiolate, less often sessile;
lower leaves usually trifid with orbicular or ovate lobes, sometimes lower
leaves lobed, lyrately-incised; upper leaves undivided.

Type species: A. lancea (Thunb.) DC.

The genus includes three or four species, distributed in the south
and Soviet Far East, in northeastern China, Korea, and Japan. However,
it must be mentioned that Japanese authors have described a few more
species. Taking them into consideration, the total number of species
reaches dozens. In my opinion, almost all of them are described without
a sound basis. The error in establishing individual species can partly be
explained by the following reasons.

The existing opinion that the native habitat of the type species is in
Japan is erroneous. The type species and consequently also the genus
were established on the basis of plants cultivated in Japan, which
originated in China. These plants, cultivated in Japan as ornamentals
for more than two centuries, have naturally been modified. The
impression of the diversity of Atractylodes species could also be created
from the fact that their leaves vary from simple and undivided to trifid-
compound. They are also lyrately-pinnate. It should be mentioned that
the lower leaves are usually lyrately-lobed or trifid-compound, whereas
the upper leaves are undivided. Undivided lower leaves are usually found
in young plants. The presence of petiolate and sessile leaves is important
for distinguishing Atractylodes species. The former are typical for A.
ovata (Thnb.) DC., distributed in the USSR, and sessile leaves for A.
chinensis (Bge.) Koidz., the species most widely distributed in
northeastern China, which has been repeatedly reported wrongly from
Soviet territory.

1. A. ovata (Thnb.) DC. Prodr. VII (1838) 48; Koidz. Fl. Symb.
Or.-As. (1930) 4; Kom. and Alis. Opred. Rast. Dalnevost. Kr. I], 1058.—
A. lyrata var. ternata (Kom.) Koidz. op. cit. 5——A. japonica Koidz. ex
Kitam. Comp. Japon. I (1937) 19, saltem p. p.—Altractylis ovata Thnb.
ex Murr. Linn. Syst. veg. ed. 14 (1784) 730; Thnb. Fl. Japon. (1784)
306; Kom. Fl. Manchzh. III, 714—Acarna ovata Willd. Sp. pl. Ill, 3

90

92

(1800) 1701.—A. chinensis auct. non Bge. (1836); Maxim. Prim. FI.
amur. (1859) 172, p. max. p—TIc.: Kom. and Alis. op. cit. Plate 316.

Perennial. Rhizome procumbent, 6-10 cm long, about 2 cm thick,
usually producing two or three, 40-60 cm high stems. Stems erect, nbbed,
usually simple, less often branched above, densely leafy. Leaves petiolate;
petioles of lower leaves 3—6 cm long, shorter in upper leaves, lower
leaves trifid, lobes-leaflets ovate or almost orbicular, 5-10 cm wide,
lighter colored beneath, finely serrate and short-setose along margin;
leaves reduced upward, trifid or undivided to broadly lanceolate. Capitula
solitary, terminal on stem, or four or five, one each at apices of branches,
erect; subtended and surrounded by large pinnatisect floral leaves.
Involucral bracts imbricate, five or six-rowed, greenish, woolly above;
outer bracts elliptical, inner bracts linear, both slightly colored above.
Florets 10-12 mm long, with whitish, tubular corolla narrowed to half
and parted into lanceolate lobes in broader part. Achenes oblong, about
4 mm long, ribbed, compressed and stiff white-hairy; pappus bristles
phimose, about 8 mm long. Flowering VIII; fruiting IX.

Oak and mixed forests, along forest edges—Far East: Zeya-Bureya,
Uda (south), Ussuri. General distribution: Northeastern China (north),
Korea (north). Described from cultivated plants in Japan originating from
China. Type in Uppsala.

Note. As was also stated in the description of the genus, we are not
convinced that specimens identical with those distributed in the south
of the Soviet Far East are typical specimens of A. ovata (Thnb.) DC.
The use of the name A. ovata (Thnb.) DC. for our plant without studying
the type specimens of Thunberg, preserved in Uppsala, is only a tentative
solution to the nomenclatural problem of this species.

Subtribe 2. CARDUINAE O. Hoffm. in Pflanzenfam. IV, 5 (1984)
317.—Trib. Carduaceae Cass. in Dict. sc. nat. VII (1817) 94 p. p—
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.—Tnrib.
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. Carduineae
Less. in Linnaea, VI (1831) 86 p. p. and Synops. Comp. (1832) 8 p. p.;
DC. Prodr. VI (1837) 617; Benth. in Benth. and Hook. f. Gen. Pl. II
(1873) 213 p. p.; Boiss. Fl. or. HI (1875) 457.—Subtrib. Centaurieae
Less. Synops. Comp. (1832) 4 p. p.—Subtrib. Carlineae DC. op. cit.
531 p. p.—Subtrib. Silybeae DC. op. cit. 615—Subtrib. Serratuleae DC.
op. cit. 662 p. p—Achene with straight, less often slightly oblique areole,
mostly toothed near apex, glabrous (except in Polytaxis Bge.). Pappus
bristles one- to many-rowed.

9]

+

93

KEY TO GENERA OF SUBTRIBE CARDUINAE

. ilaments hairy joridensely ‘barbate®....: 29 eis. San) eed: ..! 2.
Filaments glabrous, less often glandular or slightly barbate; but if
haityantheny outereflonets) Stemlee ts ite aeh en cck. Bon eeeeh. a aan As. 8 fi.

» WRTAMES SRO ESP Ue es case eo . 0 ce 5 SER bi el 3.
PL ATNGS iS fol eter ce tenses cee har Aer Gat (2a iala uy bh Gas vba todscu Ravlieawes bette 4.

. Receptacle flat, honeycomb-pitted, pits bordered with short scales;

achenes without apical border, with rounded margins; plants without
stem or with short (up to 30 cm high, usually simple stem
KC AMC ASUS) )issice. meee drank. Beate, 1585. Jurinella Jaub. and Spach
Receptacle almost cylindrical, convex, covered with long scales;
achene with a small apical border; stems higher (50—100 cm),
strongly virgate from base, numerous (Central Kopetdag)............
Eh a Pedal Shae eS a OR AN SE 1588. Nikitinia [ljin

Paps Cols SernatCmDrStles aes 35 Sees naked: aleene al Sea home ar hneehs. Ns 5.
Pappusrar plumose“bristlese ose in ee Pete Ren a tea 6.
. Filaments free above, connate below due to dense, short, and slimy

hairs; pappus surrounding crown of very short, thin, simple hairs
forming ring at base; leaves with large white spots...............0.....
BRED. ABS. AAO. VME GDD AO Se SAL MIO OBL, 1600. Silybum Adans.
Filaments wholly free; pappus without crown of short hairs inside;
leaves, without, spots... ...22 V2.0. HIRO: 1589. Carduus L.
Achene with thick pericarp, becoming hard and strong on maturity,
ovoid-hemispherical, or more or less pyriform .............0::cccccees
BS oder chia Bade cco: etek Rick eer eee Ee eee ee 1959°. Laymra Cass.
Achene with thinner pericarp, of different shape (cf. also step 26)
saeae eae dati ete saa in tebe eldc fh vat hare hell 1592. Cirsium Mill.
Receptacle deeply pitted, receptacular pits with membranous,
irregularly incised-toothed border; capitula large (4-7 cm wide;
smaller; only in O. frickii Tamamsh.); plants strongly spiny .......
* soetiice Aces 5 yee bh adherent is iene: manne 1601. Onopordum L.
Receptacle covered with bristles or scales, occasionally (in some

SGUSSUTER. WC ;..SPECICS)\ ClADLOUS, cane ias. .csunedevoaa.. docelstentest-ceee- -eeeee 8.
Pappus bristles scabrous or serulate, less often achenes without
Pappus (n” some” Cousimia (Cass: ispecies), :..0..363...Ae ae 2.
All or only inner bristles of pappus plumose ...................006 ae

Anthers without basal appendages; achenes (mature) usually single
in capitulum (Nurata Mountains in Soviet Central Asia)..............
Seppe duce. Merwinstic, ans Ie 9 RS 1577. Anura (Juz.) Tschern.
Anthers with basal appendages; achenes more numerous in
(SE VoNLA DIU yeaa nediee: rete ela memebenirmnctetny sl © etl ined ntacte it) uel gel sl iN 10.

. Pappus bristles connate at base into ring, readily shedding as a

WAC Eres ata ena dn Aula Mesa tt tin toast orks ails otal cliewatmiltae ae Lt.

92

94

16.

Pappus bristles not connate into ring at base, shedding individually

ldo Ra a Be Oe eS ies £8 aera, Uiaee Ate 4
. Leaves not spinulose ...t.......cccceeeeeeeeeeees Om. NOMA SRR 2, Wen
Leaves and involucral bracts spinulose ........0......::cccesecceseereeeees 13.

. Receptacle with a fewer scales; achenes oblong-turbinate, with

indistinct ribs, without apical border (central Kopetdag) ..............
Bpnai eiseim tit be hdc atlanta mh es. Bing hoa 1586. Perplexia I}jin
Receptacle densely covered with scales over large part; achenes
obpyramidal, four-angled, ribbed, less often somewhat compressed,
more or less oblong with apical border (cf. also step 21)............
wie ti a ic a eae OE den oo ORE Se ees ae 1584. Jurinea Cass.

. Involucral bracts with scaly appendages; corolla tube always shorter

than limb; basal processes short (1—2.5, less often up to 4 mm
long), wide; inner pappus bristles slightly flattened above...........
sh a= as a i taal Sola ore 1590. Alfredia Cass.
Involucral bracts without appendages; corolla tube longer than limb,
less often variable in length; basal processes long and narrow;
inner pappus bristles not flattened above, but longer and desnely

SORRAtE Gh saccade ade gece eens ete wad cat ie 1591. Olgaea Iljin
. Pappus of slightly unequal bristles, usually uniform in thickness
catch Bint aaa an wagner acta aes inane petite cd bolane eto nr gat HSE
Pappus of distinctly unequal bristles 0.0.0.0... ee eeeeeeeeeeeeeeeee 16.

. Involucral bracts coriaceous, with terminal spine, usually not

uncinate; receptacular scales longer than or almost as long as
achene, less often half to one-third as long; pappus bristles readily
Shedding caches wiamdel owen. ral alee eta 1578. Cousinia Cass.
Involucral bracts membranous, spinulose at tip; receptacular bristles
one-fourth to one-fifth as long as achene; pappus bristles persistent,
later. shedding 4 eve. te .. Pik ee ee 1579. Lipskyella Juz.
Involucral bracts with stiff terminal spine, not uncinate; receptacular
bristles serrate-scabrous; plants perennial; leaves spinose (Tien
Shani) it 8.8 Ae SED ee ath 1580. Hypacanthium Juz.
Involucral bracts with terminal, thin, uncinate spine; receptacular
scales scabrous; plants biennial; leaves without spines .................
Ae ae NE: Re Oe me EEE SR ot ccs BCE etc ae oer 1576. Arctium L.
Capitula heterogamous, peripheral florets sterile, central florets
bisexual) 2:8 124, t. Shoaib. aie, Beatle. can ad, 1593. Notobasis Cass.
Capitula homogamous; all florets tubular, bisexual ................. 18.

. Receptacle with scales, less often papillae (glabrous in some

Saussurea DC. ospecies)) ies...) 5 2. Renta aie Beet eee Sh
Receptacle wath bristlesiesi24.. bevss cnt ees beh eee ce 22

. Achenes covered with scattered long hairs, recurved in lower part,

spreading above; bristles of inner row of pappus connate to variable

93

on.

22:

Aah,

27s

95

height into tube (3—-5.5 mm long), has -hairy on outer side
Dis: SANG BPN ORS cot ssh Me sa ace dca whWaalds 1583. Polytaxis Bge.

. Achenes glabrous; pappus bristles not connate into tube........ 20.
20.

Pappus many-rowed; its bristles variable in length, connate at base
into ring or firmly attached to achenes and without ring ....... Zale.
Pappus of one or two rows; bristles of outer row of pappus short,
scabrous or slightly plumose, readily shedding individually; bristles
of inner row equal, always long-plumose, connate at base into
TAME eOCESISLCMIL coccneccsceseocertecsotsbrsasancrneceetass cs 1582. Saussurea DC.
Anthers covered with long, very thin, flexuous, fragile readily
shedding hairs; pappus without 2—5 mm long inner bristles (Soviet

Wenliral inSain eee erect anne: 1587. Pilostemon [ljin
Anthers glabrous; pappus with two to five longer inner bristles
(eit aiso Slemml) e iirivir ess ete tases carcweeans 1584. Jurinea Cass.

Pappus bristles not connate at base into ring, readily shedding
individually, short-plumose. Alpine and subalpine plants with
bipinnatisect leaves (Tien Shan).... 1581. Schmalhausenia Winkl.
Pappus bristles connate at base into ring and shedding together
saab lala I sa a allah ge cb ow Aca ah 8 23:
Capitula surrounded by apical leaves; outer involucral bracts with
plumose and prickly process; plants annual-biennial .....................
Suncast dedi ane ode dla riety salteetial Ame 1594. Picnomon Adans.
Capitula not surrounded by apical leaves; prickly process of

involucral bracts not plumose; plants perennial ...................... 24.
. Receptacle fleshy; leaves very large, wide, bipinnatisect, forming
TOSELC! CUMIVALCE! IDVANIES aos cons. ssuseresc-cnenctse ssa 1599. Cynara L.
Receptacle not fleshy; leaves almost entire, undivided, or pinnatifid
lh gc Aa a cag NR NG Et AR SE EA 25.
. Achenes weakly flattened on both sides, not ribeed................ 2:
Achenes not flattened, ribbed-angular ...................::esceseseesreeees 27.

. Achenes with obliquely truncate apex; pappus many- -rowed (cf.

AISOUStCD WO) a aed eae a Mec 1592. Cirsium Mill.
Achenes flat-tipped, with saucer-shaped depression; pappus in two
rows of light golden bristles, slightly cystoid below and pointed
above, (Soviet -Cemtral cA sia) cscs. casi tt Becht eee be cial: me gunveoy ea
Ee See ee aT 1596. Lamyropappus Knorr. and Tamamsch.
Stem winged; leaves oblong-elliptical or lanceolate; achenes white,
pale or dark brown; pappus many-rowed, of short-plumose bristles,
outer bristles: Shorten... 2:c024.. vs-edsssces, eecdeaatas toaster hee wey aiweneeee ot
‘net, coud nena edaeees ah oct 1598. Modestia Charadze and Tamamsch.
Stem not winged; leaves linear or linear-lanceolate, achenes dark
brown or almost black; pappus usually in two rows of almost equal,
long-plumiose ‘bristles .......1....:.c0ccessssssessesoee-s- 1597. Ancathia DC.

94

96
GENUS 1576. Arctium L.!: 2

L. Sp. pl. (1753) 816.—Bardana Hill. Veg. syst. 1V (1762) 28.—
Lappa Adans. Fam. pl. (1763) 116.

Capitula homogamous. Receptacle flat, somewhat fleshy, covered
with numerous bristles initially flat, later spirally twisted. Involucre
glabrous or arachnoid-hairy, many-rowed; involucral bracts linear or
lanceolate, outer and middle bracts narrowed into distinct uncinate tip;
inner bracts more or less membranous, erect. Florets bisexual, similar,
all fertile, with regular, tubular, five-lobed corolla. Stamens with free
glabrous filaments; anthers sagittate below, acuminate above. Style
surrounded with suprapistillate disk at base persisting in fruit, with linear
and basally curved hairy branches. Achenes oblong, laterally compressed,
truncate at apex, ribbed, usually somewhat rugose between ribs
(especially at base and near tip), less often smooth; pappus short, its
bristles unequal, scabrous, in many rows, free to base, shedding; areole
basal, straight. Biennial, almost nonprickly large herbs with alternate,
large, simple, .petiolate leaves and branched inflorescence-corymbose or
paniculate.

The genus is of Mediterranean origin, some of its 11 species are
widely distributed as ruderal plants.

Economic Importance: The species of this genus, particularly the
ones not widely distributed (A. lappa, A. minus, A. tomentosum), are
ancient medicinal plants. Their roots are known in the pharmacopea
under the name Radix Bardanae and contain inulin, sugar, and essential
oil. Their various preparations are used for treatment of many diseases.
The oil, which is 25-30% in fruits of burdock (Oleum Bardanae), also
finds medicinal application. The young roots and branches are used as
food. Melliferous plants.

1. All involucral bracts terminating in hook, or inner-most either
gently curved and then apically not broadened, mostly green, less
often reddish; corolla limbs without glandular hairs on outer side.
Common inflorescence corymbose or racemose (in latter case
peduncles’ short’or capitula’ sessile) iis. 2) cctisea.. eerste eco ncoae cues De

+ Inner involucral bracts more or less broadened upward, acute or
obtuse, with short straight cusp at tip, usually purple in upper
part; corolla limbs on outer side with small multicellular glandular
hairs. Common inflorescence corymbose or racemose with more
or, less..distincts peduncles: ew .nteeee ed RS (ee A oe 6.

'Treatment by S.V. Juzepczuk and E.V. Sergievskaya.
*From the Greek name of burdock, arcion or arceion, first mentioned by Dioscorides.

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97

. Common inflorescence corymbose; peduncles more than 1 cm (up

toy 2y[sic]pem)alony wie Steere rie roy earth AREY, Te ratir ens. des 8:
Common inflorescence racemose; peduncles rather short or capitula
SEIS KS ett lane anteater Apc sp ee ad Racial et Sy

. Capitula very large, involucre 20-30 (with cusp 40-SO mm) wide;

middle and outer involucral bracts up to 1.5 mm wide at base of
curved part; achenes about 7 mm long, pitted-rugose...................
Ec ccbbnehts- eee 1. A. platylepis (Boiss. and Bal.) Sosn. ex Grossh.
Capitula 13-20 (with cusp 30-35) mm wide; middle and outer
involucral bracts 1-1.5 mm wide at base of curved part; achenes
5—6(6.5) mm long, pitted-rugose only at apex or near base, or
ALMOSE/ SiO bly Re faery. cane ie” ate: Speers tas eset ae OTE Bante Ca 4.

. Common inflorescence corymbose; peduncles up to 10 cm long.

Inner involucral bracts as long as or longer than florets. Achenes
with distinct longitudinal ribs near apex and base, pitted-rugose,
finekyy PUNCaleKe.>..<ge sere: eee Re. TN. 2 2. A. lappa L.
Common inflorescence corymbose-racemose; peduncles 1-3 cm
long. Inner bracts much shorter than florets. Achenes with indistinct
longitudinal ribs, elsewhere smooth, with large dots..........0.0.0.0..
EREEE «Loves SR REL ES EEE. SONI 3. A. leiospermum Juz. and C. Serg.

. Branches of common inflorescence virgate, often arcuately curved.

Capitula 1.5—2.5 cm wide, with cusp 34 cm; inner involucral
bracts as long as or longer than florets. Achenes 6—8 mm long..
eecaeet .Jie..:t ebouneel Sucwerrameners Meus ee |, ce) 4. A. nemorosum Lej.
Branches of common inflorescence erect, upward spreading.
Capitula globose-ovoid, terminal, 1—1.5 cm wide, with cusp 2—2.5
cm. Inner involucral bracts shorter than florets. Achenes 4-5 mm
LOND? . atalseecrs. . Abbe ...fvvetry...-<- Bone... 5. A. minus Bernh.

. Involucre glabrous or only slightly arachnoid-hairy. Innermost

involucraly-brdctspacuminatelis...A.-2..05. cee. OAR... B.A RE. Th
Involucre more or less distinctly arachnoid-hairy, occasionally
glabrous. Outermost involucral bracts truncate or even emarginate
at apex, with small cusp .....0. 7. A. tomentosum Mill.

. Capitula glabrous or indistinctly arachnoid-hairy. Inner involucral

bracts spatulate at apex, crenate-toothed and crimped along margin,
acuminate. Stem in upper part and branches of common
infloresence almost smooth; leaves weakly toothed, with nonprickly
LOSE) \. atamnpcccueteacs 1... eymeuhames | toa 6. A. palladinii (Marc.) Grossh.
Capitula glabrous; involucral bracts with finely ciliate-toothed
margins, their broader part oblong, not spatulate. Stem in upper
part and branches of inflorescence strongly scabrous, tuberculate;
leaves with large-toothed margin, teeth terminating in distinct
SPINWIE Shave. 4)... BLAST. LEO? »f 8. A. radula Juz. and C. Serg.

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98

Section 1. Eglandulosa Aréne in Bull. Jard. Bruxelles, XX (1950).—
Inner involucral bracts linear-lanceolate, not broadened upward, often
nonpigmented or a little purple. Corolla limbs without glandular hairs.

1. A. platylepis (Boiss. and Bal.) Sosn. ex Grossh. Fl. Kavk. IV
(1934) 164; Klok. Fl. Abkh., IV, 264—Lappa platylepis Boiss. and
Bal. in Boiss. FI. or. II] (1875) 458.—4A. lappa subsp. platylepis Aréne
in Bull. Jard. Bruxelles, XX (1950) 74, 77.—Ic.: Aréne, Ibid. 79, pl.
IIIA.

Biennial. Plants more than 1 m high. Stem erect, strong, deeply
longitudinally sulcate, often reddish, branched, with strong and erect
upward spreading branches almost as long as main stem. Leaves petiolate,
broad-ovate; bases of lower leaves cordate; uppermost leaves truncate,
margin crenate-toothed, with teeth terminating in small thin spine, green
above, glabrous or sparsely pubescent, grayish arachnoid-hairy tomentose
beneath; basal leaves with swollen petioles; cauline leaves greatly
reduced, but uppermost relatively large, often closer to capitula. Common
inflorescence corymbose or corymbose-paniculate, dense capitula
somewhat condensed, although solitary on 1-5 cm, less often up to 7
cm long peduncles, larger than in A. lappa. Involucre 2-3 cm wide
(excluding cusp), 4-5 cm with cusp, glabrous or slightly arachnoid-
hairy. Involucral bracts 1.5—-2.25 mm wide, wider and flatter as compared
to A. lappa; cusps (divergent part) lanceolate from base, up to 1.5 mm
wide at base, with subulate tip (except innermost bracts); all bracts with
terminal spinule; outer bracts often strongly hooked; innermost bracts
more linear, up to 2 mm wide, as long as outer bracts, longer than
florets, pubescent; often all involcural bracts dark violet. Corolla usually
purple, slightly larger than in A. lappa, with campanulate 5.5-6 mm
long limbs and 2.5 mm long lobes, abruptly narrowed into 8-9 mm
long tube (in A. lappa up to 5—6 mm long), much broader at base;
apical appendages of anthers narrow and pointed, 0.3 mm long, basal
appendages simple or two- or three-parted, 1.25 mm long. Achenes 7
mm long and 2.5—3 mm wide, oblong-turbinate, with truncate apex,
longitudinally sulcate, transversely crenulate throughout, rugose, brown
to dark brown with almost black pattern; pappus 3.5 mm long. Flowering
VII-VIII. (Plate VII, Fig. 2).

Forest covered mountain slopes, subalpine tall grasslands.—
Caucasus: Ciscaucasia (State Reserve), western Transcaucasia (Abkhazia),
southern Transcaucasia (Bakuriani), eastern Transcaucasia (northern
Azerbaidzhan). General distribution: Eastern Anatolia. Described from
Habakher in Lazistan. Type in Geneva.

Note. Undoubtedly an endemic species of the Caucasus with a
discontinuous range, known from several places. The material of this

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99

species is heterogeneous, which indicates its tendency to form local
races. For instance, the dark purple color of the involucral bracts recorded
in the description was observed by us only in specimens from the western
Caucasus. The hook of the outer involucral bracts, which are also longer,
characterizes mainly the Azerbaidzhan and Bakurian specimens.
Characteristics not typical of the Caucasian form of this species (slightly
narrower arachnoid involucral bracts, etc.) are mentioned in the
descriptions and figures of typical A. platylepis. The lack of authentic
specimens and the scant Caucasian material do not permit the segregation
of any taxonomic entities.

Aréne’s monograph of the genus recognizes A. platylepis (considered
as a “subspecies” of A. lappa) as “an alpine Pontian race.” We, on the
contrary, are inclined to accept A. platylepis as a Tertiary relict plant,
probably the ancestor of the widely distributed but, apparently always
ruderal and synanthropic A. Jappa. Aréne further reported the existence
of intermediate forms between A. platylepis and A. lappa in Lazistan
and Cappadocia. Incidentally, the Turkestanian form separated by him
as similar to the “intermediate,” in our opinion, belongs to a separate
species, not having any direct relationship with A. platylepis.

2. A. lappa L. Sp. pl. (1753) 816 excl. var. B.—A. lappa subsp.
majus Aréne in Bull. Jard. Bruxelles XX (1950) 75; Fedtsch. and Fler.
Fl. Evrop. Ross. III, 1001; Grossh. Fl. Kavk. IV, 164.—A. majus Bernh.
Syst. Verz. Pfl. Erf. (1800) 154; Syreistsch. Ill. Fl. Mosk. Gub. III,
289.—Lappa vulgaris Hill. Veg. syst. IV (1762) 28.—L. major Gaertn.
De fruct. II (1791) 379; Ldb. Fl. Ross. II, 748; Schmalh. Fl. II, 94.—
Ic.: Rehb. Ic. Fl. Germ. XV, 812; Syreistsch. op. cit. 290; Hegi, Ill. FI.
VI, Il, 828.—Exs.: Woloszcz. Fl. polon, exs. No. 650.

Biennial. Plants 60-150 cm high, reaching up to 3 m. Root branched,
fleshy, thick, fusiform, up to 60 cm long. Stem erect, strong,
longitudinally sulcate, often reddish; branches numerous, erect, spreading,
covered with papilliform hairs mixed with glandular hairs, but mostly
arachnoid-hairy. Leaves petiolate, broadly cordate-ovate, sparsely crenate-
toothed or entire, green and sparsely pubescent above, finely gray-
tomentose beneath with scattered yellowish glandular hairs; basal leaves
up to 50 cm long, on ridged, tumid, more than 30 cm long petioles;
cauline leaves abruptly reduced, uppermost somewhat cordate at base.
Capitula borne sparsely in corymbose-racemose inflorescence, sometimes
on peduncles up to 10 cm or longer, and almost at same height at tips
of branches, globose, large, 13-20 mm wide excluding cusp, with cusps
20-35 mm long. Involucral bracts glabrous or weakly arachnoid-hairy;
lower bracts lanceolate, carinate, 1-1.5 mm wide, weakly ciliate-toothed
along margin, narrowed into linear-lanceolate, erect spiny tip; middle

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100

bracts appressed-pubescent; inner lanceolate, green, sometimes purple,
pubescent, gradually narrowed into weak apical hook as long as or longer
than floret. Corolla purple-red, abruptly narrowed into 5—6 mm long
tube, limb 4—5 mm with 1.5—2 mm long lobes; apical appendage of
anthers 0.2 mm long, narrow and pointed, basal appendages 0.75 mm
long, simple or bifid. Achenes 6—6.5 mm long, 2.5—2.75 mm wide,
narrow-obovate, with narrow areole, pappus attachment surrounded by
four to six tubercles; achenes longitudinally ribbed, transversely rugose
near upper and, to lesser extent, at lower end, grayish to brown, with
zig-zag dark brown to almost black pattern; pappus 3—3.5 mm long.
Flowering VIII-IX. (Plate XII, Figs. 10, 11).

At garbage dumps, near inhabited areas, along roads and fences,
banks of rivers and rivulets, rarely in cultivated fields——European Part:
Balkhash Region, Karcha-Lapland, Ladoga-IImen, Upper Dnieper, Upper
Volga, Volga-Kama, Middle Dnieper, Volga-Don, Trans-Volga Region,
Lower Volga, Lower Don, Black Sea, Upper Dnieper, Bessarabia (?)
Crimea; Caucasus: All regions; Western Siberia: Upper-Tobol; Far East:
Ussurt, Zeya-Bureya, Sakhalin. General Distribution: Scandinavia,
Atlantic and Northern Europe, Mediterranean Region, Balkans-Asia
Minor, Iran, Indo-Himalayas, Tibet, Mongolia, China, Japan, introduced
into North and South America. Described from Western Europe. Type
in London.

Note. In the Far East and Japan, besides the typical A. lappa,
specimens with larger capitula and wider involucral bracts, strongly
resembling the Caucasian A. platylepis in appearance, are often found.
Similar specimens, sometimes collected in the European part of the USSR
(Mogilev), are also known in cultivation (in the Botanical Gardens they
are called “Lappa grandiflora’). | assume that these forms are only a
modification of A. /Jappa and do not have a direct relationship with A.
platylepis. It should also be noted that a part of the Crimean plants and
those from the southern regions of the European part of the USSR are
distinguished by a smaller capitulum, their size being closer to that of
A. minus.

3. A. leiospermum Ji.z. «ad C. Serg. in Bot. Mat. Gerb. Bot. Inst.,
Akad. Nauk SSSR, XVIII (19 °7) 299.—4A. lappa auct. omnium FI. Asiae
Mediae, non L.

Biennial. Tall plants \'ith strong, erect, deeply sulcate stems;
branches numerous, usually turning red, erect, spreading or sometimes
strongly, covered with fine papilliform hairs and dense, yellowish
glandular hairs, without arachnoid-hairy pubescences. Leaves petiolate,
broadly ovate, with cordate base, crenulate-toothed, green, and almost
glabrous above, slightly grayish-tomentose beneath with dense covering

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101

of fine glandular hairs; basal leaves about 30 cm long and 25 cm wide,
with long prickly, tomentose, tumid, often turing red, more than 20
cm long petioles; cauline leaves rather abruptly reduced, upper leaves
with truncate base. Common inflorescence corymbose-racemose. Capitula
in isolated groups, clustered on relatively short, 1-3 cm long peduncles,
globose, medium to large, 1—1.5 cm wide excluding cusps, 2.5-3.5 cm
long with cusp. Involucral bracts similar to those of A. /appa, but inner
bracts much shorter than florets, appressed-pubescent. Corolla purple,
abruptly narrowed into 6—7 mm long tube, limb campanulate, 5-6 mm
long with about 2.5 mm long lobes. Anthers about 5 mm long, apical
appendages up to 0.5 mm long, basal appendages about 0.75 mm. long,
undivided or bifid. Achenes 5—6 mm long, 2—2.75 mm wide; narrow-
obovate, with narrow areola of pappus attachment surrounded by small
tubercles, brown to gray with almost black spots and wide zig-zag stripes,
almost smooth, with indistinct longitudinal ribs; pappus up to 3 mm
long. Flowering VII-VIII, fruiting XI.

Valleys and river banks, along streams and irrigation channels, near
springs, found as a crop weed, on deposits and in orchards.—Soviet
Central Asia: Aralo-Caspian Region, Balkash Region, Tien Shan, Syr-
Darya, Pamiro-Alai, mountainous Turkmenia; Western Siberia: Irtysh,
Altai; Eastern Sibera: Angara-Sayans. General Distribution: Dzhungaria-
Kashgaria. Described from the village of Klyuchevka of the former
Aulieata District of Syr-Darya Region. Type in Leningrad.

4. A. nemorosum Lej. in Lej. and Court. Mag. d’Hort. I (1833)
289; Fedtsch. and Fler. FI. III, 1001; Grossh. Fl. Kavk. IV, 164 p. min.
p.—4A. intermedium Lge. Handl., Dansk. Fl. (1851) 463.—Arctium minus
subsp. nemorosum Boswell. Engl. Bot. V (1886) 25; Aréne in Bull.
Jard. Bruxelles, XX, 96.—A. minus var. nemorosum Th. Dur. in De
Wild. and Th. Dur. Prodr. Fl. Belg. II (1903) 479.—A. macrospermum
Hayek, Sched. fl. Styr. exs. XIII and XIV (1908) No. 29.—Lappa
nemorosa Koern. Schr. Phys.-dkon. Ges. Kénigsb. V (1864) 63 and in
Arch. Ver. Naturg. Mecklenb. XXIII (1870) 189; Schmalh. Flora II,
94.—L. macrosperma Wallr. in Linnaea, XIV (1840) 639.—L. intermedia
Rchb. Ic. Fl. Germ. XV (1853) 54.—L. minor var. nemorosa Crép. Man.
Fl. Belg. ed. 5 (1884) 259.—Ie.: Rcehb. op. cit. tab. 81, fig. 1; Hegi, III.
Fl. VI, II, 826, 830.—Exs.: Fl. Hung. exs. No. 590; FI. Ital. exs. No.
383.

Biennial. Plants up to 2 m high. Stem erect, strong, longitudinally
sulcate, green or reddish; branches numerous, long, lower branches
usually arcuate below, covered with short, papilliform hairs mixed with
glandular hairs and more or less densely arachnoid-hairy. Leaves
petiolate, cordate-ovate, sparsely crenate-toothed, usually thin, green,

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102

and sparsely puberulent or glabrous above, weakly tomentose and grayish-
green beneath, with scattered, fine, yellowish glandular hairs to almost
glabrous; basal leaves up to 50 cm long, with hollow, up to 60 cm long
petioles; cauline leaves abruptly reduced, uppermost leaves weakly
cordate at base, narrowed into petiole, with apical spines. Common
inflorescence racemose, upper capitula solitary or clustered in heads of
two to six, lower isolated, upper crowded, sometimes corymbose; all
capitula short-pedunclulate, or lower ones almost sessile, slightly
compressed-spherical, slightly narrowed at apex, (1.5)2—2.5 cm long.
Involucre mostly glabrous, less often slightly arachnoid-hairy; involucral
bracts lanceolate, carinate; middle bracts densely puberulent on outer
side, ciliate-small toothed, along margin, 1.5—1.75 cm wide, narrowed
into narrow-lanceolate, almost horizontal cusp, up to 1 mm wide at
base, in lower part trihedral, tlat above, terminating in stiff spinule;
inner involucral bracts lanceolate, flat, up to 2 mm wide, green or purple,
gradually and finely acuminate, terminating in spinule or slightly incurved
at apex, about as long as florets or slightly longer. Corolla purple-red,
abruptly narrowed into 5—6 (up to 8) mm long tube, limbs 5-7 mm
with 1.5—2 mm long lobes; apical appendage of anthers narrow, 0.3—0.5
mm long, basal appendages simple or bifid, up to 1 mm long. Achenes
7-8 mm long, narrow, obovoid, with narrow areole and truncate apex,
areole surrounded by fewer small tubercles, transversely rugose
throughout, brownish with black pattern; pappus 2.5-3 mm long.
Flowering VI-—VIII. (Plate VII, Figs. 2, 3).

Forests, mainly broad-leaved forests, their edges, forest glades,
clearings, gorges.—European Part: Ladoga-Ilmen? (Valdai Hills), Upper
Volga (?) (Melenki), Upper Dnieper, Volga-Kama, Trans-Volga, Volga-
Don, Upper Dniester, Black Sea Region, Lower Don, Crimea; Caucasus:
Ciscaucasia (Zheleznovodsk). General distribution: Scandinavia, Central
Europe, Atlantic Europe. Described from France. Type in Paris.

5. A. minus Bernh. Syst. Verz. Erf. (1800) 134, s. str.; Syreistsch.
Ill. Fl. Mosk. Gub. III, 290; Fedtsch. and Fler. Fl. Evrop. Ross. 1001.—
A. minus subsp. eu-minus Syme, Engl. Bot. V (1866) 26.—Lappa minor
Hill. Veg. Syst. IV (1762) 28; Ldb. Fl. Ross. H, 748; Schmalh. Fl. II,
94.—Ic.: Engl. Bot. tab. 702; Reichb. Ic. Fl. Germ. XV, tab. 811;
Syreistsch. op. cit. 291; Hegi, III. Fl. VI, I, tab. 269——Exs.: Pl. Finl.
exs. No. 985; Wolosz. Fl. polon. exs. 836.

Biennial. Plants 0.5—1.20(1.5) m high. Root thick, fusiform, fleshy.
Stem erect, more or less strong, longitudinally sulcate, green or reddish,
branches numerous, erect, spreading, like stems covered with short
papilliform hairs divergent in different directions, mixed with glandular
hairs and Icss often (particularly in lower part of stem) with arachnoid-

103

ATR Sek
AGEN
SMT in

\
Sy 4)
we G

4) Ite
yt 2

vee
ow She
= S
aay “ ti
tay |

101 Plate VII.

1—Arctium platylepis (Boiss. and Bal.) Sosn., upper part of plant; 2, 3—A. nemorosum Lej.,

inflorescence and achenes; 4, 5, 6—A. palladinii (Marc.) Grossh., upper part of plant, achene,

inner involucral bract; 7—A. tomentosum Mill., inner involucral bract; 8, 9—A. minus Bernh..
achene, involucral bract; 10, 11—A. lappa L.. involucral bract, floret.

102

104

hairy pubescence. Leaves petiolate, broad ovate; lower leaves cordate at
base, entire or sinuate-toothed, with short acicular cusp at apex, green
and glabrous or sparsely pubescent above, grayish-green beneath due to
sparse fine tomentum with shining, fine, sessile glandular hairs, less
often white-tomentose; basal leaves smaller than in A. nemorosum,
with hollow petioles; upper cauline leaves short-petiolate, ovate, with
almost truncate or angular base, irregularly crenate-toothed. Common
inflorescence racemose, terminal capitula clustered in heads of three to
six, sessile or short-pedunculate, and sometimes corymbose at tip of
main axis, slightly pressed-globose, narrowed at apex, without cusp 1—
1.5 cm, with cusp (1.5)2—2.2(2.5) cm wide, 1.3—1.8 cm long. Involucre
without or with arachnoid-hairy pubescence; middle involucral bracts
densely puberulent on outerside, lanceolate, carinate, ciliate-toothed, 0.5—
1 mm wide at base, narrowed into linear-lanceolate, horizontal stiff spine;
all bracts, including inner, green or slightly reddish; inner bracts narrow-
linear, gradually narrowed into slightly bent cuspidate tip. Corolla longer
than inner involucral bracts, pinkish-red to purple, abruptly narrowed
into initially violet, later reddish, 4.5 mm long tube; limbs 4-5 mm
with 1.5 mm long lobes; apical appendage of anthers obtuse, not narrow,
0.2 mm long, basal appendage simple or bifid to many-parted 0.5 mm
long. Achenes 4—5 mm long, 1.75—2 mm wide, with 1.5—2 mm long
pappus, turbinate, ridged, often three or four-angled, truncate, transversely
rugose at both ends, with small tubercles near point of pappus attachment,
brownish with dark brown zig-zag pattern. Flowering VII-IX.

Among debris, near inhabited areas, along roads, dumps, and
canals.—European Part: Baltic Region, Ladoga-IImen, Karelia-Lapland,
Upper Dnieper, Upper Volga, Middle Dnieper, Volga-Don, Volga-Kama,
Trans-Volga Region, Lower-Don, Bessarabia?, Black Sea Region,
Crimea. Described from Germany.

Note. This species is reported in the literature as widely distributed
in the Caucasus; however, none of the specimens from there seen by us
with the identification “A. minus” belong to this species; most of them
belong to A. palladinii.

Nevertheless, judging from the description, the plants named as
Lappa glabra y. caucasia C. Koch (Linnaea, XVII, 43) and reported
from Georgia and Armenia should be included only under 4. minus.
However, I did not see the herbarium specimens; therefore, it is included
tentatively among the synonyms of A. minus.

A. minus Bernh. forms the following hybrids.

1) x A. nothum (Riihm.) Weiss in Hallier. Kochs Synops. I (1902)
1512.—Lappa notha Rihm. Jahrb. Bot. gart. Berl. I (1881) 238.—L.
intermedia Schur. in Verh. Mittl. Sibenb. Ver. Nat. Hermansst. IV, 6
(1853) 35, non Lge. nec Rchb.—L. subracemosa Simenk. Enum. fh.

103

105

Transs. (1886) 340.—A. subracemosum Nym. Consp. Fl. Eur. suppl. I,
1 (1889) 179.—Lappa janczewskii Dyb. in Wszechswiat (1904) No. 46.—
A. lappa subsp. majus x A. minus subsp. eu-minus Aréne in Bull. Jard.
Bruxelles, XX (1950) 124.—Exs.: Callier, Fl. Siles, exs. No. 1311.—
European Part: Middle Dnieper (Sumy Region, Grebennikovo), Trans-
Volga Region (Buguruslan’ Region).

2) A. minus Bernh. x A. nemorosum Le}.

Found among shrubs in forest clearings.—European Part: Trans-
Volga Region (Buguruslan Region, Sosnovsko-Arkhangelskaya estate).

Note. So far only one specimen of this hybrid is known from the
territory of USSR. We note that there are no fully reliable published
reports concerning the occurrence of the second parent (A. nemorosum)
from the Trans-Volga Region.

Section 2. Glandulosa Aréne in Bull. Jard. Bruxelles, XX (1950)
112.—Inner involucral bracts more or less broadened upward, root purple;
corolla limbs in lower (undivided) part covered with small, yellowish,
multicellular glandular hairs on outer side.

6. A. palladinii (Marc.) Grossh. Fl. Kavk. IV (1934) 164; Kolak.
Fl. Abkhazii, [V, 264. Papava in FI. Gruzii, III], 438—Lappa palladini
Marc. in Allg. Bot. Zeitschr. VI, 11 (1900) 200; Tr. Bot. Sada Yurev.
Univ. 1, 147.—A. bardana MB. FI. taur.-cauc. I] (1808) 263 p. p.—A.
tomentosum (resp. Lappa tomentosa) auct. cauc. saltem p. max p. non
Mill. (resp. Lam.)—A. tomentosum var. glabrum Aréne in Bull. Jard.
Bruxelles, XX (1950) 118 p. p. non Kém.—4A. transcaucasicum Sosn.
ex Grossh. Opred. Rast. Kavk. (1949) 474; Papava in Fl. Gruzi, VIII,
441.—Ie.: Markovich, op. cit. Fig. 1, 2c, 3c, 4c_—Exs.: GRF No. 927.

Biennial. Plants more than 2 m high. Stem erect, strong, branched,
longitudinally sulcate, green or reddish; branches usually long, numerous,
erect, spreading (often in same plane), rough due to stiff irregular
papilliform hairs, arachnoid-hairy pubescent, mixed with fine glandular
hairs. Leaves petiolate, with cordate base, ovate, entire or crenate-toothed,
cuspidate, green and sparsely hispid above, grayish-green beneath due
to somewhat dense arachnoid-tomentose pubescence, mixed with yellow
glandular hairs. Common inflorescence corymbose with lateral branches
longer than main axis. Capitula globose, broadly open at apex, excluding
cusp 0.9-1.2 cm, with cusp 1.5-—2.2 mm wide and 1.5—2.1 cm long,
readily disintegrating on maturity, glabrous or occasionally slightly
arachnoid-hairy. Involucral bracts ciliate-toothed; lowermost bracts short,
recurved, narrow-lanceolate, spiny-tipped; lower involucral bracts
lanceolate, carinate, narrowed into linear-lanceolate not sharply curved
cusp, terminating in less strongly curved hook than in 4. tomentosum;
inner involucral bracts purple in upper-half, linear-lanceolate; innermost

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106

covered with fine appressed hairs, carinate in middle, often crimped
along upper margin, crenate-toothed, spatulate. Corolla much long than
involucral bracts, reddish-purple, sparsely yellowish glandular-hairy; tube
up to 6-7 mm long; limb about 7 mm long with 3.5-4 mm long lobes.
Anthers up to 5-6 mm long, apical appendages sharply narrowed into
0.2 mm long cusp, basal appendages 0.75 mm long, mostly undivided
or bifid. Achenes 3.5-5 mm long, 2—2.5 mm wide, obpyramidal, with
narrow areole, ribbed, with indistinct tubercles, brownish gray, finely
punctate, mature achenes over entire surface crenate-pitted-rugose; pappus
up to 3 mm long. Flowering VII-VIII, fruiting up to X. (Plate VII, Fig.
6).

Forests, banks of rivers and streams, gardens, weedy meadows, debris
dumps.—Caucasus: Ciscaucasia, Dagestan, eastern, western, and southern
Transcaucasia. Endemic? General distribution: Iran? Described from
Alagir in Northern Ossetia. Type or isotype in Leningrad.

7. A. tomentosum Mill. Gard. Dict. ed. 8 (1768) No. 3; Syreistsch.
Ill. Fl. Mosk. Gub. III, 291; Fedtsch. and Fler. Fl. Evrop. Ross. 1000;
Grossh. Fl. Kavk. IV, 164; Kryl. Fl. Zap. Sib. XI, 2875.—A. lappa var.
B. L. Sp. pl. (1753) 860.—Lappa tomentosa Lam. FI. Fr., Il (1778) 37;
Ldb. Fl. Ross. I], 749; Schmalh. Fl. II, 94.—Ie.: Rchb. Ic. Fl. Germ.
XV (1852) tab. 811; Syreistsch. op. cit. 291; Hegi. Ill. Fl, VI, Il, tab.
270, 833.—Exs.: Wolosz Fl. polon. exs. No. 745; Fl. Ital. exs. No.
1170; Pl. Finl. exs. No. 1398.

Biennial. Plants 60-150 cm high. Root fleshy, thick, fusiform, brown.
Stem erect, strong, longitudinally sulcate, green or reddish; branches
numerous, erect, spreading, less often (in shady places) slightly arcuate,

covered, like stems with papilliform hairs mixed with glandular and
' arachnoid-hairy pubescence. Leaves petiolate, ovate, with cordate to

broadly cordate base, entire or sparsely toothed, obtuse, distinctly
mucronate, green and glabrous or slightly appressed-pubescent above,
densely grayish or whitish arachnoid-hairy tomentose beneath with sessile
golden glandular hairs; upper leaves short-petiolate, ovate or ovate-
oblong, petioles of basal leaves solid, tomentose, often reddish to brown.
Common inflorescence corymbose. Capitula almost corymbose,
numerous, globose, slightly narrowed at apex, excluding cusp 11-18
mm and with cusp 18-30 mm wide, 13-20 cm long, readily disintegrating
on maturity, densely arachnoid-hairy pubescent or occasionally glabrous.
Middle involucral bracts densely puberulent on outer side, sometimes
mixed with glandular hairs, ciliate-toothed; outermost involucral bracts
short, with fine sharply recurved, hooked cusp; outer involucral bracts
lanceolate, carinate, broader at base, 1.25—1.5 mm wide, narrowed into
linear-lanceolate, horizontally divergent hooked cusp; innermost

10

Nn

107

involucral bracts purple, broadened at base up to 2.75 mm, linear-
lanceolate, somewhat obtuse or sinuate, with small prominent but not
hooked cusp; all involucral bracts often purple. Corolla much longer
than inner involucral bracts, reddish-purple, sparsely yellowish glandular-
hairy, campanulate, broadened upward; tube 4.5-6 mm long, broader
at base; limb 4—4.5 mm with 2—2.5 mm long lobes. Apical appendages
of anthers more or less sharply narrowed, 0.2 mm long; basal
appendages 0.6 mm long, simple or bifid. Achenes 5—6 mm long,
turbinate, with obtusely truncate upper end and narrow areole of pappus
attachment, ribbed, transversely rugose, grayish to brown, brownish or
with dark zig-zag spots against lighter background; pappus 2.5—3 mm
long. Flowering VII-IX. (Plate VII, Fig. 7).

Among debris, kitchen gardens, along road, near walls and fences.
A widely distributed weed—European Part: Baltic Region, Ladoga-
Ilmen, Karelia-Lapland, Dvina-Pechora, Volga-Kama, Upper Volga,
Upper Dnieper, Middle Dnieper, Volga-Don, Trans-Volga Region, Lower:
Don, Upper Dniester, Black Sea Region, Bessarabia, Crimea; Caucasus:
Possibly, in Ciscaucasia; Western Siberia: Upper Tobol, Irtysh, Ob, Altai;
Eastern Siberia: Yenisei, Angara-Sayans, Yakutia (introduced); Far East:
Ussuri (introduced); Soviet Central Asia: Aralo-Caspian Region, Balkhash
Region, Tien Shan, Syr-Darya, Pamiro-Alai (north), Dzhungaria-
Tarbagatai. General distribution: Scandinavia, Atlantic and Central
Europe (England), Balkans-Asia Minor, Dzhungaria-Kashgaria. Described
from England. Type in London.

Note. Most of the herbarium collections from the Caucasus attributed
to this species actually belong to A. palladinii, and only plants from a
few localities in the northern Caucasus can be included under 4.
tomentosum.

Economic Importance: To a great extent, ornamental plants; being
extremely common, however, they are not used for this purpose in the
Soviet Union.

A. tomentosum Mill. forms the following hybrids.

1) x A. ambiguum Nym. Consp. Fl. Eur. suppl. II (1889) 179.—
A. lappa x A. tomentosum Beck. Fl. Nied. Oest. (1893) 1228.—L. major
x L. tomentosa Hausskn. in Oest. bot. Zeit. schr. (1864) 206.—L.
ambigua Celak. Prodr. fl. Boh. (1871-1872) 249.—L. tomentosa var.
denudata Lge. Handb. Dansk. fl. ed. 3 (1864) 288.—Ic.: Lge. Fl. dan.
tab. 2915.—Exs.: Wolosz. Fl. polon. exs. No. 631; Callier Fl. Siles.
exs. No. 1310.

Found among debris, together with the original species ——European

Part: Upper Volga, Volga-Don, Trans-Volga Region (Buguruslan
Region).

106

108

Note. A fairly widespread hybrid, readily forming wherever 4.
lappa and A. tomentosum grow together.

2) x A. leiobardana Juz. and C. Serg. hybr. nov.—A. leiospermum
Juz. and C. Serg. x A. tomentosum Mill.

In general appearance and characters of inflorescence, it is more
similar to A. leiospermum, from which it differs by broad and apically
colored inner involucral bracts and sometimes also by a weakly
arachnoid-hairy involucre.

Found in wastelands, along river banks.—Soviet Central Asia:
Dzhungaria-Tarbagatai, Tien Shan.

Note. A hybrid described for the first time, which occasionally is
found in Soviet Central Asia within the range of A. leiospermum.

3) A. nemorosum Lej. x A. tomentosum Mill.—A. minus
3) macrocephalum Rupr. Fl. ingr. (1860) 541.

Wastelands, clearings——European Part: Ladoga-IImen (Kronstadt),
Volga-Don (Gorodishche Region).

— Note. Borshchow’s old collections of this plant in Kronstadt deserve
special attention. Ruprecht (op. cit.) considered these specimens as a
variety of A. minus, but later (on the herbarium label) considered them
as the hybrid A. minus x A. tomentosum. The large achenes compel me
to include the Borshchow’s plant under the above-mentioned hybrid;
apparently, we are dealing here with an introduced plant.

4) x A. mixtum Nym., Consp. Fl. Eur. suppl. Il (1889) 179.—Lappa
mixta Simonk. Term. Fiiz. 2 (1878) 145.—L. ritschliana Aschers. in
Berl. Deutsche bot. Ges. IX (1891) 99.—L. minor x tomentosa Porc. FI.
distr. Roman Nasend. in Transilv. (1885) 113.—A. minus subsp. eu-
minus X A. tomentosum Aréne in Bull. Jard. Bruxelles, XX (1950) 126.—
Exs.: Callier, Fl. Siles, exs. No. 1309.—European Part: Baltic Region,
Ladoga-IlImen (Lake Dubets), Upper Volga (Melenki), Trans-Volga
Region (vicinity of Pugachev, Buguruslan District), Volga-Kama (Kirov,
Ufa). Crimea?

8. A. radula Juz. and C. Serg. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVIII (1957) 300. A. platylepis Sosn. in Sched. non Boiss.
and Bal.

Biennial. A tall plant; stem erect, strong, longitudinally deeply
sulcate, strongly rugose at least in upper part due to strong conical
tubercles, sometimes partly finely tomentose. Leaves petiolate, somewhat
hard, ovate, with cordate base, deeply crenate-toothed, with variable
teeth, often quite large, terminating in rather long and strong spine;
green and sparsely short-setose above; rough, densely grayish- or white-
tomentose beneath. Common inflorescence corymbose with strong
divaricate branches; peduncles up to 1.4 cm long. Capitula crowded,

107

109

globose, excluding cusp 1.5—2 cm, with cusps 2.5—-3 cm wide, 1.5—2.5
cm long. Involucre entirely glabrous; involucral bracts ciliate-toothed,
their cuspids narrow-lanceolate, almost three-ridged, with apical hook,
recurved in outer involucral bracts, rather short, erect and squarrose in
inner bracts; inner involucral bracts linear, gradually broadened upward
and later gradually narrowed into spiny cusp (in innermost bracts without
hook), very scarcely fimbriate-toothed along upper margin, purple in
upper part. Corolla not or slightly longer than involucral bracts, purple
on outer side, with scattered glandular hairs; tube 6.5 mm long, limb
about 8 mm long, with 4 mm long lobes. Anthers 6—7.5 mm long, their
apical appendages abruptly narrowed at apex, basal appendages simple.
Achenes about 6 mm long, 2.5 mm wide, flattened-obovoid-pyramidal,
ribbed, with rather distinct, somewhat sharp tubercles on ribs, brown
with dark brown spots and stripes, somewhat rugose mainly in upper
part and at base. Flowering VII.

Among shrubs, near habitations.—Caucasus: Eastern Transcaucasia.
Described from Chingyly on Lake Sevan. Type in Leningrad.

GENUS 1577. Anura (Juz.) Tschern.!:?

In Bot. Mat. Gerb. Inst. Bot. Akad. Nauk UzbSSR XVII (1962)
107.—Cousinia sect. Anura Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR,
8 (1940) 512, 546.

Capitula homogamous, with fewer florets (three to five), all florets
bisexual, tubular; involucral bracts imbricate, appressed, terminating in
very short, incurved cusp; receptacles setose, receptacular bristles smooth;
filaments free; anthers without basal appendages; achenes with straight
hilum, usually fused with receptacle, as a rule, only one achene in a
capitulum; pappus setose, pappus bristles scabrous, readily shedding
individually.

Perennial herb, densely covered at base with numerous reticulate
remains of old leaves, basal leaves in well-developed rosette, cauline
leaves sessile.

Type species: A. pallidivirens (Kult.) Tschern.

1. A. pallidivirens (Kult.) Tschern. in Bot. Mag. Gerb. Inst. Bot.
Akad. Nauk UzbSSR, XVII (1962) 107.—Cousinia pallidivirens Kult.
in Tr. Sredneaz. Gos. Univ., Ser. VIII C, VI (1929) 3; Juz. in Tr. Tadzh.
Bazy Akad. Nauk SSSR 8 (1940) 546.—Ie.: Kult. op. cit., Fig. 2.

'Treatment by O.V. Tscherneva.
*From the Greek word literally meaning “‘tailless.”

108

110

Perennial. Stem densely covered at base with numerous reticulate
remains of previous year’s leaves, erect, 25-50 cm high, striate-sulcate,
arachnoid-hairy pubescent, sparsely. and paniculately branched from base,
in turn, branches repeatedly branched, each twig with one capitulum;
inflorescence dense, corymbose-paniculate. Leaves coriaceous, glabrous
above with reticulate prominent venation, grayish tomentose beneath;
basal leaves petiolate with basally broadened petioles, ovate, almost entire
or sometimes very finely prickly-toothed, slightly crimped along margin,
apex prickly-acuminate, cauline leaves very abruptly reduced upward,
lower leaves sessile, semiamplexicaul, base large prickly-toothed on both
sides, ovate, very finely prickly-toothed or with almost entire, apex with
short prickly cusp; middle leaves sessile, semiamplexicaul, coarsely
prickly-toothed at the base, ovate-oblong, finely prickly-toothed; upper
leaves oblong, sessile, sparsely prickly-toothed. Capitula numerous, on
fragile short peduncles, 15 mm long and 5 mm wide, somewhat oblong-
cylindrical, glabrous, with fewer florets (three to five). Involucral bracts
appressed, slightly divergent at apex, with very short incurved cusp,
slightly carinate, greenish in the middle, with white border; inner bracts
almost membranous, obtuse, very short-acuminate; corolla whitish or
pale yellow. Anthers without tail-like basal appendage. Receptacular
bristles smooth. Achenes 8 mm long and 3 mm wide, white, with
longitudinal dark striations, oblongate-oval, usually only one achene in
capitulum. Flowering VI—VII; fruiting VIJ—VIII. (Plate OIG Brgy)

Middle mountain belt——Soviet Central Asia: Pamiro-Alai (Nurata
Mountains). Endemic. Described from the Aktau Mountains near Tamana.
Type in Tashkent.

GENUS 1578. Cousinia Cass.!: 2

Cass. in Dict. sc. Nat. XLVII (1827) 503; Bge. in Mém. Acad.
Pétersb. 7 sér. 9, 2 (1865) 6; Boiss. Fl. or. HII (1875) 458; Winkl. in Tr.
Peterb. Bot. Sada, XII (1892) 181; Juz. in Tr. Tadzh. Bazy Akad. Nauk
SSSR, 8 (1940) 507.

Capitula homogamous, all florets bisexual, tubular; involucral bracts
usually imbricate, with strong cusp. Receptacle setose; receptacular
bristles smooth or scabrous. Filaments free; anthers connate into tube;
apical appendages of anthers variously shaped, basal appendages plumose.

'Treatment by O.V. Tscherneva.
"Named after the 19th-Century French historian, Victor Cousin, or after Louis Cousin,
member of the French Academy of 17th Century.

109

111

Achenes with straight hilum, obovoid or obpyramidal; pappus setose,
bristles scabrous, usually yellowish, readily shedding, less often pappus
absent.

Annual, biennial, often monocarpic perennial or perennial herbs, or
semishrubs. Leaves alternate, undivided, pinnatisect, or lyrate; basal
leaves often rosulate. Stems leafy, less often leafless, simple or branched
with capitula aggregated in racemose or paniculate, less often in
semicorymbose inflorescence.

The genus comprises more than 500 species, mainly distributed in
Northwest and Soviet Central Asia.

Type species: C. orientalis (Adams) C. Koch.

Economic Importance: Some species of Cousinia are rubber
producers. For example, according to the results of analysis of the
Laboratory of Rezinotrest, C. resinosa Juz. contains 7.88% rubber and
67.92% resin. The presence of rubber has also been confirmed in other
species close to C. resinosa, as also in C. vicaria, C. dolichophylla and
C. lanata. Some species of the genus are good fodder plants, e.g., C.
triflora, C. resinosa, etc.

Note. The proposed classification of the genus cannot be considered
final, because a phylogenetic classification can be proposed only after a
monographic study of the genus as a whole. The classification proposed
here is mainly a further development of the system proposed by S.V.
Juzepezuk in Tr. Tadzh. Bazy Akad. Nauk SSSR (8, 1940), since, in
fact, the available classifications of the genus by A.A. Biinge (1865)
and K. Winkler (1897), especially by the latter, err toward extreme
artificiality.

Juzepcezuk, in his above-cited work, reported that he borrowed this
classification from a still unpublished review of the Soviet Central Asia
species of Cousinia, which, unfortunately, remains unpublished and is
not traceable even in his archives. Juzepczuk worked on Cousinia for
many years and was a great authority on this genus; unfortunately, he
could not complete the work. Thus, I share the responsibility of
describing, among other species, also those mentioned in the herbarium
of Juzepczuk, whom I consider as my teacher.

1. Annuals, without well-developed rosette of basal leaves, root not
Ehicker ‘than Stetie ee er eee er eter wnca cutee ens ae 2.
+ Biennials or perennials with well-developed rosette of basal leaves
(often withering before flowering or fruiting); root thicker than

stem (or thicker than individual stems, if stems many).......... 16.
2. Receptacular bristles distinctly or weakly scabrous ................... 3:
+ RECEPtacWlar” OLISLLES SMMGOUM tre. eecc ee ener Deere eee eee if

3. Capitula almost globose, glabrous; involucral bracts upright, with

110

1)

fine, smooth, straight cusp; corolla pink .............ccccccsccesseeeseeeeeees
segpraptas pad. ..marsiahencle. aes ieeebe. 244. C. tenella Fisch. and Mey.
Capitula broadly cylindrical or ovoid-oblong; involucral bracts
usually with decurved or horizontal cusp; corolla whitish or grayish
yellow, if pale pink, capitula always arachnoid-hairy ............... 4.

. Capitula clustered terminal, ovoid oblong, 4 mm wide (excluding

CUSPS)}2J.ccett ERE RATR: AS AAG ROM I, 169. C. angusticeps Juz.
Capitula solitary terminal, 6—11 mm wide (excluding cusps)... 5

. Receptacular bristles very long and strongly scabrous; corolla

grayish-yellow; cusps of involucral bracts scabrous ..................
adnssiagitde'en tuaucttg edema sien eeeence wet eae: Seer ORIN 30. C. pygmaea Winkl.
Receptacular bristles very long and weakly scabrous; corolla
whitish, pale pink or creamish, cusp of involucral bracts smooth

. Involucral bracts with sharply decurved weak cusp..............:0008

RS OE A cot eR cee, SN OED. A 167. C. tedshenica Tschermn.
Involucral bracts with horizontal, narrow-lanceolate, long cusp...
facie ogcactareatteas: ceneeet es, Rm aia. wen chee 168. C. murgabica Tschern.
Cauline leaves narrowed toward both ends, not amplexicaul; stem
strongly branched dichotomously from base; involucral bracts with
deltoid-lanceolate or narrow-lanceolate, somewhat uncinate cusp

Cauline leaves (at least upper) amplexicaul; stem somewhat sparsely
branched dichotomously; involucral bracts with subulate-linear or
narrow-lanceolate, but always straight, cusp .............:ccseeseeees 5.

. Stem glabrous, lustrous; involucral bracts thick, carinate, deltoid-

lanceolate, with straight cusp; outer bracts horizontal; inner bracts
WELLS Merete stretcrtte sc eeeneanetee uc enat tn ere nee een enee 232. C. minuta Boiss.
Stem partly floccose-arachnoid-hairy; involucral bracts narrow-
lanceolate, all erect; outer bracts narrowed into weakly uncinate
CUSP;, MANET bracts» PULplew...% 12. estes .n eee 170. C. eugenii Kult.

. Leaves green on both sides; multicellular hairs mainly along veins,

and sessile glandular hairs scattered beneath throughout; arachnoid-

Haihye PUDESCEMES” SPAlSC noc<c.tec.nc-s-dcustans arse -eeseeeeeeree eeeaeer eres teee eee 10.
Leaves grayish or nearly white-tomentose beneath, arachnoid-hairy
pubescences predominant... S282... ere ete tee ee ee. eee 14.
. Involucral bracts straight, erect or horizontal .......0..... eee lid
Involucral bracts more or less decurved or nearly falcate ...... 13.

. Leaves glandular above; capitula 9-10 mm wide (excluding cusps)

MA SIRNA A onisy 4a ae a A aA Aa 08 161. C. polytimetica Tschern.
Leaves with multicellular hairs above, isolated glandular hairs
scattered throughout; capitula 5-8 mm wide (excluding cusps)
iM a Mate ats tin gli ah aime nems Lane ent eA a ee CIEL 1122

111

16.

113

. Capitula 5-6 mm wide (excluding cusps); involucral bracts 5-8

mm long, upward spreading, with fine cusps; achenes glabrous,
with undivided, longitudinal wings as thick ribs...................00
SIS, SPR OER. EER MRE. AL 159. C. dichotoma Bge.
Capitula 7-8 mm wide (excluding cusps); involucral bracts 9-15
mm long, horizontal or erect, but always with strong deltoid cusp;
achenes pitted-rugose, with emarginate longitudinal wings as thick
PANS PRNISILD « Se ete TR TA.. MIMREENS, | sce 160. C. erectispina Tschern.

. Capitula 5—6 mm wide (excluding cusps); involucral bracts with

falcately decurvedicusp Pe: 2008. Ae... 163. C. sylvicola Bge.
Capitula 7-8 mm wide (excluding cusps); involucral bracts with
somewhat decurved cusp .............. 162. C. patentispina Tschern.

. Stem glandular, with arachnoid-hairy pubescence leaves sparsely

tomentose beneath; achenes with undivided longitudinal wings as
thickens Jeacerroe 1G. einer, Oooh, 166. C. orthacantha Tschern.
Stem eglandular; leaves sparsely grayish-tomentose beneath;
achenes with longitudinal emarginate wings, as thick ribs..... LS:

. Leaves glabrous above; capitula 8-9 mm wide (excluding cusps);

involucral bracts eglandular, with horizontal cusp; inner bracts
PUGS Be IK. PERS. LU Ce 165. C. pusilla Winkl.
Leaves glandular above; capitula 1 1—14 mm wide (excluding cusps);
involucral bracts profusely glandular, with erect broad cusp; inner
bracts non-pigmented ............ 164. C. bungeana Rgl. and Schm.
Involucral bracts, at least outer and middle or only middle bracts,
cristate-prickly-toothed, i.e., with two to seven pairs of thin

horizontal spinules, without appendages ..............::::cceeeeeeeees We
Involucral bracts glabrous along margin, or scabrous, never cristate-
spinose-toothed, but some bracts spinescent ..............c0cccceee 25

. Capitula 12-20 mm wide (excluding cusps), solitary on stem or

few; receptacular bristles unequal, longest ones slightly scabrous
or with one or two crenations; corolla pale yellow, whitish, or
lightnpinie ...2.c:ele..20ah...A Vine, MIR DION RU. YLBeRe, 18.
Capitula 4-10 mm wide (excluding cusps), always aggregated at
branch tips in corymbose heads or lax panicles; receptacular bristles
smoothzrcorollas yellowelwsGh We Ueuels. viseiee mova: REAR. 20.

. Capitula globose-cylindrical, 15-20 mm wide (excluding cusps);

leaves basal, long-petiolate, entire with remote, thin, short and
basally frequent spines continuing on petiole; corolla pale yellow;
receptacular bristles very long with one or two crenations ..........
abet SER REELONE, oF. (POR 0. .occitesingutncnnbinccsauebenee 118. C. rigida Kult.
Capitula globose-ovoid or oblong-cylindrical, 12-15 mm wide
(excluding cusps); basal leaves short-petiolate, with broad petioles,
crenate-prickly toothed; corolla whitish or light pink; receptacular
bristles very long, somewhat scabrous ............0.::cccecceeeeeeeeeeees 1:

114

112 rtp

Die

22s

235

24.

25%.

26.

. Leaves on both sides finely gray-tomentose, more densely beneath;

Stem: »white;.arachnoidshairyitiu:...22c2eeies Ae eee... EL...
Schbehelbens sees ls: cae Gel® ob. cies 119. C. mindshelkensis B. Fedtsch.
Leaves light green on both sides, glossy, densely glandular,
sometimes finely white-tomentose along margin or entirely; stem
glandular, with papillae scattered throughout ................ccsceeseeee:
Sea aaeraaies. pee het ieee Shoe ee pete. 120. C. gomolitzkii Juz.

. Capitula with fewer florets, usually with three florets; achenes

usually jsolitanys:..dade.c..nenatioxes.o 125. C. triflora Schrenk.
Capitula with many florets; achenes usually fewer or numerous
aineat.Lawoetleniues embry. attain) abiding ROR EN meee - Zt.
Basal Leaves 18—20 cm long and 3.5 cm wide, oblong-lanceolate;
capitula on very short peduncles, readily shedding; outer and middle
involucral bracts with scarious margin from base, not so in upper
one-third, but with two or three pairs of horizontal spines on both
sides up to apex; latter and apical spines, uncinately incurved;
anther, tube} smocthnrw. atenese es. 46. C. anomala Franch.
Basal leaves 25—50 cm long and 10-25 cm wide, ovate-lanceolate
or elliptical; capitula not shedding; horizontal spines straight along
margin of outer and middle involucral bracts; tip not uncinate; if
apical spine uncinate, then usually up to 3 mm long; anther tube

short-pilose or sparsely sericeous above .............:ccecceeeeteeeeeeeees a
Inflorescence compact, broadly corymbose; capitula 8-10 mm wide
(excludingy cusps) mttshie:, cunt. cst: Lees te. BOR, 2k eee. 28.
Inflorescence lax paniculate; cantina 6-7 mm wide (excluding
CUSPS) cecnrectenetstenacennen AAPM ROENe, SEAS | AA ee Seana: 24.

Capitula 22 mm long and 10 mm wide (excluding cusps); leaves
sparsely arachnoid-hairy above, with glandular hairs scattered
throughout; outer involucral bracts more or less inclined, erect ..
ie Ae, eee. duvnaaeke sl. Ae sees aEy | geal ts 123. C. horrescens Jus.
Capitula 17-19 mm long, 8-9 mm wide (excluding cusps); leaves
sparsely arachnoid-hairy above, later glabrous, eglandular; all
involucral bracts more or less apressed ............::c:ccssceeseeeeeeeneeeeeees
eet WedAA DEOMI. caeet ST aeY MET 121. C. alberti Rgl. and Schmalh.
Basal leaves sparsely glandular; involucral bracts numerous (about
60);;withi-long ainciate;ctisp:? %:.. leaevhal is ceertels eee Te!
PU ENeN os vo ARREE c gad aeaaeaN, BLEED 122. C. pentacantha Rgl. and Schmalh.
Basal leaves densely glandular; involucral bracts about 40-50,
acuminate into long, slightly bent spine ............0.ccceeeeeeeeeeeeeeeees
opto shee RN cepecseena ct osahas sdederan cet 124. C. pentacanthoides Juz.
Involucral bracts with uncinately incurved apical cusp........... 26.
Involucral bracts with straight apical cusp .......0.... cee eeeeeeeeeeee 30.
Involucral bracts (outer and middle) with one or two large glandular
hairs on both sides along margin ...............:ccceeee ou. pabte neh af.

114

30.

33.

34.

Involucral bracts eglandular along margin..............0.e eee 28.

. Capitula densely arachnoid-hairy tomentose ...............:ceeseeeeeeees

weedes “ESE GE: Coeeeee omg ore eeL ike. haa A” - 43. C. tomentella Winkl.
Capitula‘olabrous .cieu ead 6.8 dene 42. C. umbrosa Bge.

. Capitula globose, oblong-cylindrical, or almost ovoid, 10-12 mm

wide, with’ long subulate. cuspstius:.. eee A .. oo)
Capitula ovoid readily shedding, 5-6 mm wide, with short fine
CUISPIS, 522. peePese ese ieen. 0 FREY. acuscennnanens 45. C. lappacea Schrenk

. Basal and lower cauline leaves 20—60 cm long (including petiole),

ovate, more or less cordate, apically roundish and not spinescent,
crenate-lobate, glabrous and green above, finely gray-tomentose
Beneath. Acces. cereus: peeeie. ceeetiena. of 44. C. pseudarctium Bornm.
Basal leaves much smaller, pinnatipartite or pinnatilobate, acuminate-
spinescent, with finely prickly-toothed lobes; green and sparsely
glandular above; green, densely glandular, with arachnoid-hairy
pubescence of long, flexuous, flattened hairs beneath ....................
Pe. er, MRR te ncn ER Yih ae 41. C. arctioides Schrenk
Stem more or less uniformly winged throughout or almost so
(sometimes wingless only in inflorescence branches and on lower

INFEENOMES) nest. Lites ie Wee ee tents SOS a tae 31.
Stem entirely wingless or discontinuously winged because of
Sheathing bases oh canline leaves)... 01.0... ccsbece-ccsdeeesaseeedstieroses- 64.
. Leaves soft, basal and lower cauline leaves entire, crenate-toothed
or with two pairs of teeth only at base ............eeeeeeeeeeeseeeeees 32.
Leaves thin coriaceous, basal and lower cauline leaves pinnatipartite
or pinnatisect, lyrate-lobate or pinnatilobate ...........0.. ee 44.
. Involucral bracts numerous (more than 100), narrow, with fine
SUOUIAGS™ CUSD crete cert acece sete tperthemertbe tenceccacue Rees eee ne 33.
Involucral bracts fewer (20—50), wider, with more or less broadly
lanceolate,.cuspyis! ‘avs: 0) sade cated Tele Ae ee a, 35).

Capitula glabrous, 10-12 ITM Wades er er. COR Been”
d Aiseras chad tzak Sete. aR dntben Ebro! 02 219. C. leiocephala (Rgl.) Juz.
Capitula’ arachnoid, (8—lSwmmnawide aint.ce. i ea, 34.
Leaves acuminate above; capitula globose-oblong, 12—15 mm wide
(excluding cusps); inner involucral bracts with narrow-ovate, about
1 mm wide, gradually narrowed appendages ...................cccsseeceee
R38 95 See Ri. ca aM aa Aine alin as 218. C. severtzovii Rel.
Leaves arachnoid-hairy above; capitula ovoid, 8-10 mm wide
(excluding cusps); inner involucral bracts with broadly dvate, about
4 mm wide, abruptly narrowed apical appendages. ..................0
seteusn apa, « Sanasdeas. eas. seb eeeiaves. emma, oes 220. C. pulchella Bge.

. Capitula in groups of few, terminal on branches, on short peduncles

or sessile, readily shedding, 4—5(6) mm wide (excluding cusps)
se Be Nie. ik «3 RR Male joy OAM at eas te tC eat 36.

115

116

3h:

38.

39:

40.

41.

42.

43.

Capitula solitary terminal on branches, persistent, (6)8—25 mm wide
andusenesnaeceasnescunessthetOORIEUE, MAMIE AeLCHL EEE Se, veaeetee, EAS ERCE | Bulle

. Plants about 60 cm high, with 2-3 mm wide wings on stem;

involucral bracts 25, about 2.5 mM Wide ........ccccccececcecccccccccccecceeees
LAN. DOR o. DEU, 21 SOY 3 47. C. pseudomollis Winkl.
Plants 15—40 cm high, with narrower, 1-2 mm wide wings on
stem; involucral bracts more numerous, about 40, approximately
LES prianineyide, Me oe carseat 48. C. leptacantha (Bornm.) Juz.
Biennials with one stem; stem wings usually crenate-toothed, 3-8
mm wide; inner involucral bracts membranous, often extended in
appendagessisall. | 0c. Dake As. 2eOIISI OO MGD SIRT? 38.
Perennials, usually with few stems; stem wings entire, 1-2 mm wide,
inner involucral bracts gradually narrowed, without membranous
appendages s 2600), WRAL VAI, IGM BATH Ae oierge 42.
Plants almost entirely nonprickly in vegetative parts, leaves and
stem wings undivided; corolla greenish yellow ................cc:0000
Soe RANGER NED sc coztacaeecccunemaeannate shorts 222. C. regelii Wink.
Plants more or less prickly, leaves and stem wings spinose-toothed;
corollagpimkon purplescne a Sh MUN Seo Ry’ Cane 2. 32
Receptacular bristles scabrous; inner involucral bracts membranous,
brown with light icolored! margin 10.222. 2908.. ese Pie Ts
Spc deci sic hsv ont tee 223. C. arachnoidea Fisch. and Mey.
Receptacular bristles smooth; inner involucral bracts with mem-
branous, light colored, 3-5 mm wide appendages.................... 40.
Basal and lower cauline leaves obovate-orbicular or ovate, crenate-
spinose-toothed or weakly lobate.... 172. C. macroptera C.A.M.
Basal and lower cauline leaves oblong, remotely lobate or finely
SPINOSE-1OOUN EM ..cocteccrworeennarccwcheracewsceenenneceercaen AAO 4 ean eee 41.
Plants entirely gray-tomentose; capitula 10-12 mm wide (without
cusps); outer involucral bracts about 2 cm long; inner bracts with
obtuse, about 3 mm wide appendages... 227. C. ortholepis Juz.
Plants less pubescent; capitula 7-9 mm wide; outer involucral bracts
shorter, about 1 cm long; inner bracts with broader, about 5 mm
wide, appendages isin o.n sete. 226. C. platylepis Schrenk
Capitula on short peduncles, aggregated at branch tips usually in
twos; involucral bracts with fine Short ierect "cusp ei...) sc.)
SERIA IS. oS o2s seta. nasecwnsvaiag aemeaaernnentgesaesnases 140. C. alaica Juz.
Capitula usually rsolitaey at tips of stem and branches; involucral
bracts withistine traleatewousp 201. aan eek Re eO 43.
Inner involucral bracts broadened above, membranous, roundish-
ovate, with short and fine cusp; corolla deep yellow; receptacular
bristles; ‘snideth Eiken, 0. Ie) wo 141. C. fallax Winkl.
Inner involucral bracts linear, thicker, acute; corolla whitish;
receptacular bristles scabrous ..................00 139. C. grisea Kult.

116

44.

45.

46.

47.

48.

49.

50.

51

SZ.

Receptacular-DriStleS: SCADTOUS foe. et-ce se seeeteccste tect eiderecereretes 45.
Receptacular bristles smooth ..................... i deen i tt 32.
Wings of stem up to (4)5—7 mm wide, spinose-toothed or almost

pinnatifid; capitula 10-25 mm wide; inner involucral bracts with
wide membranous appendages, if appendages inconspicuous, then
capitulaalways large,"22 mm ‘wide We. 2 8. Se .. 46.
Wings of stem 2—4 mm wide, sometimes with solitary teeth;
capitula 6—10 mm wide; inner involucral bracts without appendages

Stem weakly branched with three or seven capitula; leaves soft,
almost-glabrous;- allleaves“pinnatisect iieiiuui sce cetrecdeactceseasseeseee
Pceipiocccasneteerty: Sean braMire in eee dae AID 147. C. ulotoma Bornm.
Stem strongly branched, with much larger number of capitula;
leaves thin-coriaceous, weakly spinose; middle and upper leaves
distinct in shape from basal and lower cauline leaves and not
PInTatiSece alot. SRO RNG OF Oe BEAN e a ae dil 47.
Involucral bracts squarrose with straight cusp; corolla pink or (light)
STrayish-Velows ess aun es eer 48.
Involucral bracts with arcuately bent cusp; corolla deep yellow..
So concer cs cceneer Mba ar niin beth BS madi aa ii Ati teil 49.
Stem branched only in upper half, with about 20 capitula; involucral
bracts about 45; florets 2-3 mm longer than involucral bracts,
with purple corona .............c.cceeeee 143. C. kuekenthalii Bornm.
Stem branched from base, with 25-65 capitula; involucral bracts
about 75; florets about 5—7 mm longer than involucral bracts, with
grayish-yellow corona ............::cccscceseeereeeees 144. C. neglecta Juz.
Entire plant moderately pubescent; capitula 6 mm wide (excluding
cusps); leaves lanceolate, almost undivided, with one or two pairs

OF tect at" base ee Ce Lee ees 146. C. decurrentifolia Juz.
Entire plant densely pubescent; capitula about 10 mm wide; leaves
Pinnatisecmed, HOG) mE ON OY Mee ee 145. C. dissectifolia Kult.
Corolla“yellow= eee Ee eens 157. C. botschantzevii Juz.
Corollat pink eet ROS eG el TOT OR SCNT Se ee oie

Basal leaves pinnatisect, almost without spines, grayish on both
sides, arachnoid-hairy tomentose; capitula densely arachnoid-hairy;
involucral bracts narrowed into long carinately sulcate spine ......
Le EE Ce A BROOD er) ES 221. C. pauciramosa Kult.
Basal leaves lyrate-pinnatilobate, green above, loosely arachnoid-
hairy, capitula weakly arachnoid-hairy; involucral bracts narrowed
LALOMS EONS ASOT CUISPSirorrvsectttere: nto Ue cOnMT DES MNT OT. 52:
Plants green, scantily arachnoid-hairy pubescent; capitula 15-25
mm wide (excluding cusps); inner involucral bracts broadened
above into almost roundish membranous appendage, usually pink
(O/0) 01.0) Slane are ren ene enemy Mt 224. C. microcarpa Boiss.

117

118

56"

54.

35

56.

Sh

58.

a).

60.

Plants grayish-green, with more densely pubescent stem and leaves;
capitula 10-15 mm wide; inner involucral bracts broadened into

ovate whitish or light brown appendages ...... 225. C. ninae Juz.
Capitula 4-10 mm wide (excluding cusps).............c::ceseeeseeeees 58.
Capitula 11-15-25 mm wide (excluding cusps).............::.00000+ 54.

Capitula globose-pyramidal, 25 mm wide (excluding cusps);
involucral bracts erect or somewhat horizontal, stiff, carinate;
achenes without pappus .................-. 3 oa 171. C. hystrix C.A.M.
Capitula broadly ovoid to broadly cylindrical; 11-15 mm wide;
involucral bracts with subulate, decurved thin cusp; all achenes
With PapPPUS....o...-.::-:.. ducemeressioe.eoage ter the. eee eae a2.
Stem weakly branched; branches tone usually wingless below
capitula, single headed; basal leaves deeply pinnatipartite, with
linear crenate-spinose-toothed lateral lobes ................::cecceeee 36:
Stem cymosely branched above, branches somewhat reduced,
narrow-winged right up to capitula; basal leaves pinnatisect, with
unequal, narrow-lanceolate lateral lobes, in turn pinnatipartite and
SPINES CONE ye. nak sn las ccass sevens eda eas cn scurucsunnse sunt ue eee EE eS Syl
Capitula ovoid; 12-14 mm wide (excluding cusps); involucral bracts
narrowed into strongly divergent, 44.5 mm long, falcate cusp ..
pio oetonagtatisqoauewes Gh tearaate. eibena, Wo eee oe 23. C. dolichoclada Juz.
Capitula globose-cylindrical, 15 mm wide; involucral bracts
narrowed into weakly divergent 2-3 mm long falcate cusp .........
nota lasaan uate. EO OMe c Petnas ceases 22. C. sogdiana Bornm.
Stem and leaves white arachnoid-hairy; capitula globose ovoid,
12-15 mm wide (excluding cusps); involucral bracts falcate with
subulate, up to 3 mm long cusp... 20. C. dissecta Kar. and Kir.
Stem and leaves white-tomentose; capitula oblong-ovoid, 11-12
mm wide; involucral bracts with abruptly recurved, 6-7 mm long
CUS Digs. cocci T40- trees oc teateensos 21. C. perovskiensis (Bornm.) Juz.
Corolla whitish or pink; if pale yellow, then plant very densely
white-tomentose; capitula small, with three to six florets, aggregated

at branch tips; achenes glabrous, smooth, roundish above...... ao.
Corolla yellow; anther tubes purple; plants never densely white-
tomentose; achenes three or four-angled, toothed above......... 60.

Plants biennial; capitula, like entire plant, very densely white-
tomentose with three to six florets; corolla pale yellow, later turning
Pith, .: tescond .. xia cometen: ete nlseesates ee 50. C. mollis Schrenk
Plants perennial; capitula, like entire plant, more pubescent, greenish,
with 8-10 florets; corolla pink.................. 49. C. fascicularis Juz.
All involucral bracts, except inner, falcately decurved ........... 61.
Involucral bracts erect, sometimes outer bracts attenuate into
GIVEFS ENE CUSPs cre. .denienmescecteoreoppes dees tar upese < Becta e- eee ee: 63.

118

61.

G2.

63.

68.

69.

119

Wings of stem 1—2 mm wide; capitula truncate at base; involucral
bractsysparsely. glandularcdorsally’..: 2ccee0n.i2d...alivend. meetin...
Be. Ee TS Eel sare hak 28. C. turkmenorum Bornm.
Wings of stem 4-5 mm wide; capitula roundish at base; involucral
braets, eglandulatycce tsccin.< Actecssteceus tameptereate iets Pree. 62.
Capitula 4-5 mm wide (excluding cusps); involucral bracts
cuspidate with short, 1—-1.5 mm long, slightly bent cusp ..........4:.
Pe OI EE UE 5 SINE OEE Re oe 25. C. oxiana Tschern.
See 9-10 mm wide (excluding cusps); involucral bracts
narrowed in long, about 5 mm long, abruptly falcately decurved
(SEES eR ela aah ie te EEN 24. C. alata Schrenk
Wings 2—3(5) mm wide, all involucral bracts erect, sometimes with
MEG SSIMORE, CUSD) cosa. coccioattacasscetscccnsserses: 26. C. psammophila Kult.
Wings usually up to 3-15 mm wide; outer involucral bracts
attenuate in deflected erect cusp; middle bracts with erect cusp
esa eee nied it en eG rR be nda as Diente -Schistoptera haz. -

64(60) semishrubs Or small shrubs (2.02 Sire Nee 2 65.

ekos: Perennial “Or -Orelnial ...crreesktecetcce teste eee eee neeeccoteeese ees 74.

. Capitula 4—6 mm wide, aggregated at stem and branch tips..... 66.

Capitula solitary, on long slender peduncles at stem and branch
UST RS TE SE OR CRO RUE OS eS a 6S.

t Corolla pink“on whitish 2a AINA Pere ee CAS 67.

Corolla yellow, with purple anther tubes... eee eeeeeeee eens
stevesiussreseecsdvecausesaccazsesactessseassaies 6. C. hypopolia Bornm. and Sint.

. Leaves narrow oblong-lanceolate; basal leaves 8-9 cm long, 15-

17 mm wide (including lobes); involucral bracts with thin, some-
what falcate cusp; corolla pink or purple... eee eters
Al AOPTEP SEY IME EER ON, « 4. C. leptocephala Fisch. and Mey.
Leaves broad lanceolate, 10-15 cm long and 40-45 mm wide;
involucral bracts with thin, short, deflected erect cusp; corolla
whitish vine. balsas 5. C. albiflora (Bornm. and Sint.) Bornm.
Capitula 5 mm wide (excluding cusps); involucral bracts closely
appressed, .Wwaeh vetyaShOrtserect VCUSp fo7...copapte Sees tee

Capitula 7-12 mm wide; involucral bracts narrowed in thin,
arcuately or somewhat falcately decurved, less often with almost
hor Zonta USP sce cccnccese coe eeeeehy saa caeat, oleae LM vee meee one eee 69.
Leaves glabrous and green above, densely white-tomentose beneath;
involucral bracts with short, almost arcuate cusp; corolla whitish,
wath, purple anther, tubes.....--f.cb--acrerets 18. C. antonowii Winkl.
All leaves grayish-green and arachnoid-hairy above, densely white-
tomentose beneath; involucral bracts with strongly falcately
decurved or horizontal cusp; corolla yellow or flesh-red........ 70.

119

120

TS

. Tall, strongly flexuously branched semishrub; leaves of vegetative

shoots sessile, in groups of three to six leaves on flowering shoot
alternate, one or two, oblong-lanceolate, 3-8 cm long and 3-5
mim) ‘widel:isi suspen: eae ol emiben spree 1. C. badghysi Kult.
Short, many stemmed semishrubs; leaves sessile, usually amplexicaul,
basal leaves petiolate, 7-10 cm long and 20-35 mm wide......... lt

. Corolla flesh-red; involucral bracts with more or less horizontal

CUS OME ROA Pe GE Me RR Ror etesena cesses nntaes 9. C. oreoxerophila Kult.
Corolla yellow; involucral bracts with arcuately decurvec cusp ..
Boa ph et tht nh da ea a Amel Dory AAMC a Oa at T2

. Stem sparsely leafy; Upper leaves few, strongly reduced, usually

NOE ENClOSIND: ‘Capitlla <.-2:..0..-ccesceecrers 10. C. chaetocepkala Kult.
Stem densely leafy; upper leaves numerous, usually enclosing
Poe) | PE abe inns Aetna nel neem yi obs he EM NE AMD cal oi Ss Bs
All leaves crenate-spinose-toothed .............cscceseceesceeeeceeeeeseeeeneeeeaeees
RR nee were een 8. C. stahliana Bornm. and Gauba
All leaves almost entire, only with one or two short spinescent
TCCBie. Peccciaect Bases Sich, taps Reet it rater NE ae a 7. C. mucida Kult.

. Cauline leaves few, almost undivided, somewhat undeveloped

(much smaller than basal leaves), hence stem appearing almost
leafless (especially above); corolla golden-yellow; stem always with
remains of petioles of old leaves near base; if remains present,
then mnombranmOusecrgel ..0)...2).--scscoescoesenesdeect tiedeecs Ree ee TS:
Cauline leaves (at least lower and middle) more or less well
developed, stem more or less leafy throughout; corolla purple, white
or pale yellow, or golden yellow, but in latter case leaves
amplexicaul or pinnatisect; stem basally with, sometimes without,

remainswoftoldypetiolesee.t i24h....2ee ek tee. See 82.
Basal leaves undivided or weakly pinnatilobate.........0....000...... 76.
Basal leaves pinnatipartite, sometimes undivided only in lower half
Or pilinatisece throughoute, Ree ee, Se To:

. Involucral bracts with more or less strongly deflexed cusp...........

et TRIE: ETE » RO Nae AUS 128. C. refracta Juz.
Involucral bracts with erect or horizontal cusp ..................08 ies

. Capitula 10 mm wide (excluding cusps); involucral bracts with

erech thing 2=4> mimiMonge Cusp ae ee Eee sce neers 78.
Capitula 10-12 mm wide; involucral bracts with horizontal, 5—7
mitlong cusp eee eee eae 127. C. medians Wink.

. Basal leaves usually undivided, sometimes with one or two pairs

of tateralMlobes: at oases reer ees eee eee 126. C. aurea Winkl.
Basal leaves pinnatilobate throughout or only in upper half ........
IESE ee eRe te RE BRON a 129. C. schmalhausenii Winkl.

120

Te

80.

81.

82.

83.

84.

87.

88.

89.

Basal. leaves, pinnatisectythroushoutli.. 208.2 528%.. 900... 80.
Basalgleaves undivided in lower Nall. csccc-.c.ucceccttesuresugccessoevs-cxase.
Li. LATOR A UCM. AER EET, 130. C. karatavica Rgl. and Schmal.
Involucral uracts with erect or erect-spreading, very short,
sometimes almost dart-shaped CUSDp ............::cccccscceseseseeseeeteeeeeseeenees
CDE seS REM ECHS MRA AY. 2 133. C. korolkovii Rgl. and Schmal.
Involucral bracts with divergent, falcate or strongly deflexed cusp

Cusps of involucral bracts divergent, falcate ..........000..ccceeeeee eee
oe, POR, Mead wath re TUN EN a Beak oe feted, 131. C. haesitabunda Juz.
Cusps of involucral bracts strongly deflexed, straight ...................
ic ona a xe aa Ra. a as RNa 132. C. chlorantha Kult.
Leaves green above, hard coriaceous, with very prominent, white,
glossy, veins, usually with scattered white spines throughout......
saiasacaissanesenacesannnde aa: HEREEAROD - EE. Ae ROE, 40. C. raddeana Winkl.
Veins of leaves never bearing spines, often with or without apical

SPINE site: uarethe, sera: tie Taek ae coy aol meri 83.
Biennial or perennial monocarpic plants with one-leafed, usually
MOIMMOTOGSATOOL). cee et eee (AIR es OAT RAO LAL 84.

True perennials with many-branched strongly fibrous root... 183.
Outer and middle involucral bracts abruptly narrowed medially,
later broadening in rather wide spatulate appendages ............. 85.
Outer and middle involucral bracts gradually narrowed in cusp or
abruptly narrowed at transition in narrow, trihedral, subulate cusp

tech crcce sngs-e ssa Atami). b NORE ARENAS RRC REMAN NIS 96.
. Receptacular bristles smooth or crenulate ................ccccceeeees 86.
Receptacwlansbnistles SCADTOUS \........cacsesswensoonsaeveavcenaseesionesoavec ss 90.
. Monocarpic perennials, with thickened root collar, covered with
remains of old leaves, and Woody root.........c.ccccccceccsseeseeeees 87.
Biennials, usually with unthickened root collar, roots not woody
OF eR, PR cancnecmantinannsnacesnenomcnns COMRMER.. Bee. 89.

Anther tubes velutinous due to fine, flexuous, arachnoid-hairs ....
POPPE Pere eer Se ccmoanicmamekent 148. C. lasiosiphon Juz.
Amther tubespelabraus tess 1. AL. I a. 88.
Branches and stem densely leafy; leaves long, broadly decurrent;
appendages of involucral bracts carinate, with two opposite teeth
at jbase, SpINeSCeMmt,:.0., ceere. reek 155. C. beckeri Trautv.
Leaves usually crowded in lower part of stem, stem and branches
almost leafless above; leaves not decurrent; involucral appendages
coriaceous, green, more or less sulcate, deltoid-triangular, shortly
spinescent, smooth along margin, 7-12 mm wide at base and 12-
SAU a5 CTH Ra (a age a NR 154. C. freynii Bornm. and Sint.
Stem arachnoid-hairy floccose; leaves almost entire; capitula

12]

122

90.

91.

92:

96.

97.

globose, 25-35 mm wide (excluding cuspS)...........0..ceeceeeeeeeeeee
a Rem RE ead 230. C. simulatrix Winkl.
Stem less arachnoid-hairy; leaves finely spinose-toothed; capitula
ovoid-oblong, smaller, 15-20 mm wide... eee eeeeeeseeeeneeeeeees
b ascashivatin) ahcgisnnsmaeisaatcce dae ee a Ee 231. C. vvedenskyi Tschern.
Monocarpic perennials; appendages of involucral bracts rather wide,
with two to four pairs of spinescent teeth along margin; achenes
ribbed. toothed SADOME\.:..scsycccsaeas oe scssneisassea custosescane tech ecemrne re 91.
Biennials; appendages of involucral bracts usually smooth or
scabrous along margin, but always without spinescent teeth; achenes
smooth, apically obtuse or indistinctly longitudinally mbbed, but
them:carolla Ayellow 2 eV ee TEs eS od 94.
Corollaryellow; anther tubespink.!net. eile i eee
Tesi el a eta 150. C. cynaroides (MB.) C.A.M.
Corolla pink; anther tubes pink or yellow... eee eeeeeeeees 92.
Basal leaves lyrate, with large, roundish-cordate terminal lobe;
cauline leaves broadly and shortly excurrent along stem...............
ils, dbelwacleoad stats AMUIL oA LIES 152. C. lyrata Bge.
Basal leaves broad-lanceolate, pinnatilobate, or finely and crenate-
spinose-toothed; cauline leaves sessile, broadly and long decurrent
were hapertis.. whbowrel .ahoea dd Lee OM On | SUITE I ATA O38.

. Plants 80-100 cm high, capitula 40 mm wide (excluding cusps)

cues leads aeanes seine: ulated Layass 149. C. onopordioides Ldb.
Plants 15-30 cm high; capitula 15—17(25) mm wide (excluding
(AUIS 01S) Pa et eee re, enn aaa SO AO DE 153. C. purpurea C.A.M.

. Corolla yellow; achenes indistinctly ribbed ................ceeeeeeeeeeees

ea ia Mi a ae 151. C. macrocephala C.A.M.
Corolla pink; achenes smooth, globose above.............. ee oS:

. Basal leaves lyrate, with large ovate terminal lobe; inner involucral

bracts membranous, two times as long as adjacent involucral bracts,

fOMMMINE, CCOTONA ae ieee cra eee 228. C. ferganensis Bornm.
Basal leaves pinnatisect into narrow-deltoid lobes; inner involucral
bracts not forming corona ......... 229. C. subappendiculata Kult.

Monocarpic perennials with strongly thickened root collar; root
collar resinous-sticky in living plant, sometimes covered with

numerous¢nemams ofvolddleaves tt. e5 Lek. A eae ee. DA
Biennials with relatively slender root, if monocarpic, then always
without resinous-sticky root collar... eeceeseeeceeeceeeeeeeees 126.

Stem glabrous, whitish, glossy, sometimes finely arachnoid-hairy
pubescent; corolla pink; leaves usually not decurrent, often all
leaves, except upper, petiolate or sessile, and very indistinctly
GeCUITentyht. Ley eHt cc. PA. | cccccosmsenmnevesucckpenenee ante dae ne ai 98.
Stem always pubescent, never glossy; corolla yellow or whitish;
cauline leaves often very broadly long-decurrent .................... gD).

98

122

99

+

100.

101.

104.

123

Stem repeatedly divaricately branched, terminating in umbelliform
branching, comprising two to eight short branchlets bearing
capitula; capitulum solitary borne at base of fork; umbelliform
branching appears to have common involucre of modified leaves;
capitula 6-7 mm wide (excluding cusps); involucral bracts 12-14,
middle bracts 35-40 mm long, with sulcate-deltoid divergent cusp;
receptacular bristles weakly scabrous abOVE..............:cccccseeeeeerees
TORN, SOI RD, Gari, PO COE, 51. C. eryngioides Boiss.
Stem repeatedly dichotomously branched; capitula at base of
dichasia sessile, others on peduncles, 4-5 mm wide; involucral
bracts more numerous, about 25, middle bracts 15 mm long,
terminating in short, almost horizontal, deltoid cusp; receptacular

BHISEES (SINOOUN A -. .-ccssegert)vdecassosoddecoossdesiceoes 52. C. margiana Juz.
Ricrets three toimive mucapitulunri Sapa a ee es 100.
Florets more numerous in capitulum, in any case not less than 10
ts A ne 9 aa nd ONE NN eee Seta ead ce AEF Ae cL 101.

Capitula almost sessile, cylindrical, with globose or somewhat
narrowed base; lower and middle involucral bracts with somewhat
deflected cusp; corolla whitisn ............. 88. C. dichromata Kult.
Capitula on short peduncles, ovoid, with truncate base; lower and
middle involucral bracts terminating in short and erect cusp; corolla
Velo a BB AICP ies. ccceoceetaloatebeneieiiehs 89. C. egens Juz.
Capitula numerous, all or at least those terminal on branches of
common inflorescence more or less approximate or aggregated in

Tiere Ceara eres okie minnie Ammons meter rn rena riscr 102.
Capitula few, solitary, in lax, more or less paniculate inflorescence
TR TS Sh MCA re Mie COMM belts DO ee LEE Mere ie a Sa a 118.
Capitula aggregated at apices of branches in groups of 15—40,
forming heads; capitula narrow-cylindrical .......0...0..c cee 103.
Capitula aggregated in groups of 3-12 in relatively dense
inflorescence, but not forming heads !2/0. ea 104.

. Heads dense; apical leaves almost as long as head, as if forming

common involucre; stem white-tomentose; lateral lobes of basal
leaves tenminatins sin Strong -CUSpI: QUIN Eee Reese...
wcesceu tse vecearuas sae adbeaslemameae Vcenameeribes Meee 91. C. agelocephala Tschern.
Heads more lax; apical leaves shorter than head and usually not
resembling common involucre; stem floccose-arachnoid-hairy;
lateral lobes of basal leaves terminating in short weak cusp .......
wines LERLIE SRO MOLL. SaeSE RE ECT Petar e noes ee tok 90. C. congesta Bge.
Capitula narrow-ovoid or cylindrical, 3-6 mm wide (excluding
EUSPS) cssozesacarersusaraenniass. RTT ee Le ene eet eats 105.
Capitula ovoid or broadly ovoid, 7-14 mm wide (excluding cusps)
EHDA. TIE «cbc REMIT. « IIL. SCD, MERIT OT PRIMO Re thas steno 109.

123

107.

108.

109.

110.

. Involucral bracts few (10-15), 3—4 mm wide, coriaceous, carinate

toward) apexsncuter: (tars. es... 87. C. platystegia Tschern.
Involucral bracts somewhat narrower, herbaceous, somewhat flat,
sradually, attenuate; into; CUSPiyeu:s. .v.cel..2). ees eae. Gore 106.

. Capitula narrow-cylindrical; involucral bracts closely appressed with

short, 2-4 mm long, erect cusp, without constriction at base ......
sone sacsnnis need in oienn sans Rete: Seem emaregee. Sa) ame 86. C. maracandica Juz.
Capitula narrow-ovoid; involucral bracts with distinct constriction
at the transition into longer (about 10 mm long) erect cusp.........
Pele «<SGSoeR AR ARBRE - Poneto = cee nae anes. SUGAR, ahaa nD OE TEN 107.
Cauline leaves rather long-decurrent; capitula about 20 mm long
and 6 mm wide (excluding cusps); receptacular bristles weakly
COOPMEM ADO VE fF oc cocessucsestocsansenyau erates 85. C. horridula Juz.
Cauline leaves short-decurrent; capitula usually somewhat longer
than 10-17 mm and about 4-6 mm wide; receptacular bristles
SF TT CLO he ee Ht OE Me RIN RS AIRE A 108.
Capitula aggregated in relatively dense inflorescences, 10 mm long
and 4 mm wide (excluding cusps); achenes 3 mm long...............
» BIRR EE OEE «nlc OM ae cosines neannnct REE: «M 71. C. syrdariensis Kult.
Capitula usually solitary on peduncles, in relatively lax inflorescences,
17 mm long and 6 mm wide; achenes 44.5 mm long .................06.
Soc Goose AMG GS «EAMES A ats et Se 72. C. turkestanica (Rgl.) Juz.
Plants tall, (40)60—100(150) cm high, with lax inflorescences;
petioles of basal leaves usually spinose along sides up to base...
zi ct aeRO RON RATA DARING VENT EOI, TSO RITE aE A 110.
Plants shorter, 15-60 cm high, with more compact, almost
semicorymbose inflorescence; petioles of basal leaves without
SPINES JIN lOWER Paktlsecere:..dg.. eee... Le. AISNE 113.
Very tall plants, 100-150 cm high, with almost 2 cm thick stem,
involucral bracts flat, green, sparsely glandular dorsally, closely
appressed to each other, with only slightly divergent short cusp.
Saad. othe cpipan Meisinst am. REC. Bas. RE. MOM. 92. C. androsowii Juz.
Plants somewhat shorter, (40)60—100 cm high, with slender stem;
involucral bracts without glands, somewhat carinate, with almost
trihedralt.diviersent, “StrOME CUSP -2..c..nds2osse--c05-cntsesnecasocactesennnns Ths

. Capitula 10 mm wide (excluding cusps) broadly ovoid; involucral

bracts 20, upward spreading............. 79. C. sporadocephala Juz.
Capitula 6—9 mm wide (excluding cusps), ovoid or narrow-ovoid;
involucral bracts about 25, with somewhat recurved cusp.... 112.

. Middle involucral bracts usually not exceeding (in capitulum)

florets; innermost bracts ‘almost linear ..;,..3..0:.:;200+4--20-24-eeegeeees-- oes
dan ane DERE oth b «ERE « AIEEE. Pe Gowen o ASRS EINE 77. C. polycephala Rupr.
Middle involucral bracts distinctly (by almost 5 mm) exceeding
florets; innermost bracts almost spatulate.... 78. C. finitima Juz.

124

122.

125

. Capitula broadly ovoid, 12-14 mm wide (excluding cusps) ... 114.

Capitula smaller, 7-10 mm wide (excluding cusps) .............. MS.

. Capitula 13-14 mm wide (excluding cusps); middle involucral

bracts 35 mm long, lanceolate, gradually acuminate in strong,
eaammtely sulcate, 12-15 mm long, almost falcate-attenuate cusp
nepcunt era loee apie tlede Mist. eben. Rina. . bees 76. C. scleracantha Kult.
aera 12 mm wide (excluding cusps); middle involucral bracts
20-22 mm long, gradually acuminate in divergent, 6-10 mm long
CUSPpestericees. sunhoaie. wre ICT LE Lea atoa a 82. C. ambigens Juz.

. All plants densely pubescent; capitula densely arachnoid-woolly

frees Ll rien ci" open tins le ene opiate PD) Dass el Oe 116.
Plants sparsely pubescent; capitula finely arachnoid-hairy.... 117.

. Leaves short-decurrent; capitula 8-10 mm wide (excluding cusps);

receptacular bristles scabrous................. 80. C. spiridonovii Juz.
Leaves broadly long-decurrent; capitula 7-9 mm wide; receptacular
bristles smooth...........000..0..0000s000..s000008 81. C. transoxana Tschern.

. Stem floccose-tomentose; leaves green and sparsely arachnoid-hairy

above; capitula 8-10 mm wide (excluding cusps); achenes 5.5—6
PANNA, MTN ico cava en detest no EE 84. C. kazachorum Juz.
Stem densely white-tomentose; leaves grayish-green due to fine,
appressed, white tomentum; capitula 7-8 mm wide; achenes 4.5

TIN CNA Be ass ebb naarncnsinn 83. C. resinosa Juz.
Receptacular bristles unequal, longest bristles scabrous above.....
Baik at eccerenee! tential eee ya aubeone emma $i f 1. Tehawelele, tl pene Veh
All receptacular bristles, smooth xs: isnccies322. ..ceeetactunt. alan 122.
. Capitula about 20 mm wide (excluding cusps); all involucral bracts
densely scabrous dorsally ................ 73. C. juzepezukii Tschern.
Capitula 12-17 mm wide; involucral bracts dorsally smooth.......
Bence cabin seen seca saee on SSE, RO eae AS, emer: ey See 120.
. Capitula basally somewhat roundish.............. 75. C. pungens Juz.
Capitula ‘basally noticeably foveate 3.0... Aveta. aeeneee: 2s

. Leaves green and finely arachnoid-hairy above, white-tomentose

beneath; upper cauline leaves ovate; involucral bracts 25—27; middle
bracts narrowed in middle into carinate, erect cusp ............::cce
b Sa jecnadrsseady sAaeds. ads SEPESREELLED SMa CARR a8 en 820 74. C. umbilicata Juz.
Leaves on both sides arachnoid hairy-tomentose; more densely
beneath; upper cauline leaves narrow-ovate or almost lanceolate;
involucral bracts about 40; middle bracts abruptly narrowed from
ovate base in long, almost horizontal cusp ...........0.eceeeeeeeeeeeeees
Se LEE ere cerh cee tse eset ee Ot ETE 65. C. dolicholepis Schrenk
Involucral bracts somewhat short, narrowed into short, erect, cusp
HAR, len Hea ch ., PRT ERE SOUPE UN obs uk MisGoad ckbe. te 66. C. vicaria Kult.
Involucral bracts narrowed into long, more or less divergent cusp
MM se ot es ene ca ei canlee ek tiene ace -a Balase bona Su cen Mae pew eee nate le 123:

125

125.

126(96).

130.

13k

132%

. Cauline leaves divided into narrow lobes almost up to midrib;

involucral bracts weakly scabrous at base ............ccceeeee 124.
Cauline leaves less deeply divided; or upper leaves often undivided;
involucral bracts scabrous entirely ..........ccccccecscceseeeeseeeeeeees 125:

. Basal leaves bipinnatisect, with almost linear, prickly-pointed

pinnules; capitula ovoid, 12-15 mm wide (excluding cusps);
involucral bracts about 40.0.0... eee 70. C. tenuisecta Juz.
Basal leaves pinnatisect; with pinnatilobate spinescent lateral lobes;
capitula broadly cylindrical, (12)17—20 mm wide; involucral bracts
aboutr6Ocxe..viseaeh. olanans. . 20908 69. C. minkwitziae Bornm.
Capitula ovoid, 10—15 mm wide (excluding cusps); involucral bracts
strongly scabrous, weakly arachnoid-hairy; with somewhat squarrose
ofrecurved) widevcuspl 21:8. 210062. ORAS 68. C. scabrida Juz.
Capitula almost globose, 18-20 mm wide; involucral bracts with
dense arachnoid-hairy tomentum, almost covering their more or
less rough surface, and with strongly recurved, narrow cusp.......
NOU MONG d OTT NS, SEO ES See A 67. C. angreni Juz.

Receptaculagabristles) smooth. 22101. 00 Yh BES aN On! 127.
Receptacular bristles scabrous -c2:4i:2.se2se<evacoostessvassanwacem eras 132)
iwAcheneswelabrousy ismoothins..c28 00 RU... es 128.
Achenes ribbed or flattened with rather wide wings ............. 129:
E. Conollaypuiwnk LN. : soecserceesavencvreraoe Tansee. 188. C. glabriseta Kult.
Corollanyellaw: 2s e287... CURIE, Be 189. C. triceps Kult.

. Capitula globose, 13-15 mm wide (excluding cusps); involucral

bracts numerous, terminating in fine, falcate cusp ..............cce
ASV AROCIN A L ARORA BAO G ty 29. C. olgae Rgl. and Schm.
Capitula ovoid-cylindrical, if globose, then large, 25—30 mm wide;
involucral bracts fewer, with broader, strong, usually spreading

erect on/decurved: butjnot faleate.,cusp wtecnonicre.uet eee 130.
Capitula ovoid-cylindrical, 7-10 mm wide (excluding cusp);
achenesinibbed ..cdscie..vgu ti G ORAS, BRR, WARES » BAO. Pane

Capitula globose, 25—35 mm wide (excluding cusp); achenes
flattened with wide wings along margin..............c:cccsccceceeteeeeeneees
Sealed a ics ROR. ERR CAS EEDA 158. C. spryginii Kult.
Involucral bracts about 50, dorsally glandular and arachnoid-hairy;
middle involucral bracts narrowed from a broad, appressed base
into 22-24 mm long, divergent cusp; corolla yellow.....................
SOL MOE. phone 217. C. centauroides Fisch. and Mey.
Involucral bracts about 80, arachnoid-hairy; middle involucral bracts
with 15-22 mm long, decurved or almost horizontal cusp; corolla
pale tpurple2.0)3): 2aw GRE aN, a 216. C. divaricata Winkl.
Involucre glabrous or weakly arachnoid-hairy, of numerous narrow
bracts, terminating in subulate cusp, almost as wide as thick (at

126

133.

Involucre usually arachnoid-hairy, less often glabrous; bracts not
sO numerous, wider, terminating in narrow, broadly linear, or
lanceolate cusps, more or less distinctly wider than thick (with
maximum width 1—6 mm at base or above) ..................::0008 149.
Innermost involucral bracts membranous, white or purple, very
prominent forming well-developed “crown” or “collar”, surrounding

MOTELS SRN TINE ONY RIED. ON UT AY RUE 134.
Innermost involucral bracts not prominent and not forming well-
developed “crown” or “collar”, surrounding florets............... 139)

. Involucral bracts with erect or erect-spreading cusp.................00

Sewn Pea AIRGAS A Ita eoa AEN, ss Ta Ree STR oat 173. C. radians Bge.
Involucral bracts with more or less strongly spreading, often arcuate
GUUS Preece cece vsbea tests nts Cepeedetavaedt ee dickee lean Reman, 9 7 MR Sie, 135:

. Capitula 18-22 mm wide (excluding cusps)............ccceccsseseesseeees

Btseea dunes Likhena chs) wire ah a Masse enen nia, es 2 AE Ne 174. C. coronata Franch.
Capitula not more than 15:mm wide (excluding cusps) ........ 136.

. Stems with numerous, up to 50, capitula; involucral bracts with

short, very fine falcate cusp (latter much shorter than half width
OU TC AP IRVIN): erect ere eeeseesatese.<cscs 178. C. leptocampyla Bornm.
Stems with fewer capitula (usually not exceeding 20); involucral
bracts usually with longer, stiff, horizontal, not curved or ‘slightly
curved cusp (latter, at least in middle bracts, as long as or longer
PHanalt Width Of Capit sco acscneccess ts <--cacesecccesesesanencosece 13g:

. At least lower cauline leaves deeply pinnatipartite or lobate .......

Bee eects e ae Reteee nas te sunaie nese nace ncancmeantee 176. C. alpestris Bornm.
@auline leaves. MNGUVIGE . (ocels se ee aun nae emean ec 138.

. Middle involucral bracts with long cusp, as long as width of

CAP UCU os serccseecconcscccovecyoett teh suse oe 175. C. pulechra Winkl.
Middle involucral bracts with shorter cusp, as long as half width
of capitulum or only slightly longer ...............ccccccscssesscseeeseseessees
Be Uno sewah saciaia choison suacelasie i heeies eee 177. C. mulgediifolia Bornm.

. Involucral bracts with erect or spreading straight cusp ......... 140.

Involucral bracts with more or less horizontal, divergent or
decurved) often arcuate cuspree ts ate Le, Be 143.

. Capitula narrow-ovoid, 9-10 mm wide (excluding cusps); involucral

bracts with very short cusp or obtuse and without spine; corolla
whitish, or plac, VeMOw. .stsccajenrcsecbacneasds 243. C. jassyensis Winkl.
Capitula ovoid or broadly ovoid, 8—17 mm wide; but corolla always
pink or purple; involucral bracts with well-developed cusp, or cusp
Shoney Ame MOB EIS CHE et ones ati scab dmeasascbyhablaasaaas 141.

127 141.

148.

128. alole

Involucral bracts with short, more or less obtuse, roundish or short-
pointed cuspee: seek eee. 234. C. submutica Franch.
Involucral bracts with weak, well-developed, 3-6 mm long,
Spreadinise CUSpiaerAtom.. RAEI. TARO, AUT ee ee 142.

. Capitula ovate, 8-12 mm wide (excluding cusps); involucral bracts

with 3—4 mm long cusp .............. 239. C. hastifolia Winkl.
Capitula broad-ovoid, larger, 17 mm wide; involucral bracts with
longer, 5—6 mm long, cusp .................. 240. C. transiliensis Juz.

. Capitula 6—7 mm wide (excluding cusps); involucral bracts with

short, recurved cusp............. 233. C.. krauseana Rgl. and Schm.
Capitula 10-18 mm wide (excluding cusps); involucral bracts with
well-developed, horizontal, divergent or arcuate cusp............ 144.

. Basal leaves lyrate, with a large, entire terminal lobe...................

sectors « sasbieba meet ated dd tect bh Tac ee ences ae ll 237. C. dubia M. Pop.

Basal leaves: pintiatisectyenicten.. Sie ieee ee eee 145.
. Involucral bractssancuatese ioe ee eee Ee ee ee 146.
Involucral bracts horizontal or squarrose ............::::cssceceeeeeeees 148.
. Corolla whitish or pale yellow............. 242. C. margaritae Kult.
Corollaypiniks Messed ee. echoes aida bariecsekceateeticReRie ee 147.

. Cauline leaves finely arachnoid-hairy above, white-tomentose

beneath, similar to basal leaves, pinnatisect, with narrow-linear
lateral SG@MERtS .cccccececssecsiane 238. C. kokanica Rgl. and Schm.
Cauline leaves glabrous above, green, grayish-tomentose beneath,
narrow-lanceolate or lanceolate, almost entire, with two to four
pairs of teeth only near base ...... 241. C. waldheimiana Bornm.
Capitulum globose, 11-12 mm wide (excluding cusps) strongly
truncate at base; involucral bracts with nearly horizontal cusp;
eorollayyelloware. ase... 328h.. Reee 236. C. omphalodes Tschern.
Capitula broadly ovoid, 15-17 mm wide; involucral bracts
squarrose; corolla pink or whitish ............. 235. C. tamarae Juz.

. Achenes glabrous, smooth, roundish above ................::c:ce+ 150.

Achenes four-angled, narrow-winged along edges, toothed above
Sau hee xian. ss sae, sae haeak me, Sed NN eae 156. C. smironowii Trautv.

. Leaves soft, entire, almost without spine, decurrent......................

WSIS NT en sy, ANA, «ha cI RT ok 215. C. integrifolia Franch.
Leaves not decurrent, less often weakly decurrent or with sheathing
base? andiitheny more sorilesstspimosert -2--c.ses. -ceeereete esac es: iSite
Inner involucral bracts very prominent, usually reaching or
exceeding florets (less often, as in C. stellaris, shorter than florets)
and forming well-developed “collar” or “corona” around them...
Diba MOM a Mce Minas tonr shasncpisespanes soneerniesedastecs eee MUete ot eee REE RET he2-
Inner involucral bracts not prominent shorter than florets, not
forming ‘collar or Scorona;waround them), 20-2 ie neck 168.

129

160.

161.

129

mnCorollaliyellow -stiesne..2telewiel.aiew..ad: 213. C. dimoana Kult.

Gorollagpurple.. pinks.on whites: ce, .2sc2<c4...c.ceesycaceek.-.aveeversetcntten: ji35

. Involucral bracts numerous (more than 100), relatively weak, narrow

lanceolate, 2 mm wide at base, usually falcate ...............0.... 154.
Involucral bracts less numerous (40-—60-100), stronger, 3-5 mm
wide at base, erect, if divergent, as in C. strobilocephala, then
cauline leaves short-decurrent ..................ccc:ccccceceeeccseeeseseeeeeeees |e

. Basal leaves more or less lyrate, spinulose-toothed along margin

wagner, Levetneel AIS. Nee NAN id ERR » SMB Se 8.5 209. C. rosea Kult.
Basal leaves pinnatisect, spinose-toothed along margin................
itt SAU LI AMEE haces tec. ges sarc) . rb Lg 210. C. rhodantha Kult.

. Basal and lower cauline leaves pinnatisect, pinnatipartite’ or

pinnatilobate, terminal lobe not larger than laterals .............. 156:
Basal and lower cauline leaves lyrate or almost lyrate, with large
temmumall tobe. sseenhe. .. cs isdetis Maven Seriile atseveche ly. Peet 161.

. Capitula ovoid or ovoid-oblong, 18—25 mm wide (excluding cusps),

SMARSelyMAACHMOIG-MAIGY: ....:.0...c2-c06-scancccceescpsenaeratedoeeseeceevsenees Sige
Capitula flattened-globose or globose, (20)25—35 mm wide, densely
arachnoid-hairy or almost tomentose ..............c::cccescccesseeceeteeees 158.

. Leaves light green and finely arachnoid-hairy above, arachnoid-

hairy beneath; lower cauline leaves pinnatipartite or pinnatisect
pcere lint tree. porumneetie sete t tbody. maciel 204. C. tianschanica Kult.
Leaves grayish green and finely arachnoid-hairy above, grayish-
tomentose beneath; lower cauline leaves pinnatilobate .................
gets. Ct. obras tae: merent clare. $n 198. C. gulezensis Kult.

. Middle involucral bracts squarrose, strong ............c.ccccceseees 59:

Middle involucral bracts arcuately decurved, comparatively weak
we. eboenark . Leereulon pent. ode tebe aac leery. RAMS. Melek Bam ie 160.

. Basal and eres cauline leaves pinnatilobate; involucral bracts

scabrous, carinate-sulcate, cuspidate, 3—3.5 mm long ...................
nes, weaned. Leva sdousses, eget 207. C. praestans Tschern. and Vved.
Basal and lower cauline leaves pinnatisect or pinnatipartite;
involucral bracts less scabrous, with 2—2.5 mm long cusp............
ph oP meet aha aele Ee beet 206. C. butkovii Tschern. and Vved.
Leaves densely or sparsely grayish-arachnoid-hairy on both surfaces;
involucral bracts numerous, about 100 ...........ceeceeeceesccceeeeeeeeeeteeees
Se sn ee 208. C. strobilocephala Tschern. and Vved.
Leaves grayish green and finely arachnoid-hairy on both sides,
more densely beneath, involucral bracts about 50—60...................
a ets cath cn hes Aiud doco mtaé eines «: ARENA EMMERE 203. C. laniceps Juz.
Inner involucral bracts with broadly lanceolate or almost roundish,
about 5 mm wide appendage, forming short, about 5-8 mm long,
ebllar, notxenclosingytiorets}..pex!.34..ascew eine Bibione a6 162.

130

130

164.

165.

166.

167.

168.

Inner involucral bracts with lanceolate, apically gradually acuminate
appendage, forming long-radiate collar, equaling or exceeding
florets. .L AO YASH2), LE RU) IOI RO rs SOB IS, LE eM ee 163.

. Middle involucral bracts with strong, arcuate cusps; appendages of

inner bracts dark purple.................00 186. C. grigoriewii Juz.
Middle involucral bracts with horizontal cusps; appendages of inner
involucral bracts whitish...........00.0.00- 182. C. stellaris Bornm.

. Basal leaves with large terminal lobe and one to four pairs of

lateral lobes; involucral bracts with strong tri-hedral spine, 2.5—3
mm widel'at base; SQUAITOSE!()ii.ii.s00. Lec ARE si beeh 164.
Basal leaves with smaller terminal lobe and four to six pairs of
lateral lobes; involucral bracts with thinner and shorter spine, 1.5—
2 mm wide at base, almost all erect, only outermost sometimes
decurved 3s). 20.004.20.. SRN) Fede SUR Ao) dee Teeee 167.
Leaves glabrous, above with roundish, almost cordate terminal lobe,
and one or two (three) pairs of small lateral lobes.............0.........
sare coun snthaliscisiagncet ts Ueaatenues aesiheteabe casas 211. C. rotundifolia Winkl.
Leaves arachnoid above, with greater number (three or four pairs)
of. larger: lateral..lobes.... 22 Q0AW0l. REE. SO. MUR 165.
Entire plant densely pubescent; capitula floccose-tomentose, ovoid-
globose, (20)—25 mm wide (excluding cusps); inner involucral bracts
narrow, 2.5 mm wide at base, exceeding adjacent middle involucral _
bracts by only 8-10 mm..............ce 201. C. dshisakensis Kult.
Plants sparsely pubescent; capitula densely arachnoid-hairy, globose-
cylindrical, 25-40 mm wide; inner involucral bracts 5-7 mm wide
at base, exceeding adjacent middle involucral bracts by 15-35 mm
LEP RIOD, SOOO MALOU, a RT, Lele eny 145 ek Do. 166.
Plants strong; leaves spinulose-dentate; involucral bracts strong,
inner bracts long-radiate, exceeding adjacent involucral bracts by
25-55, MDS, ee IF, 0! 199. C. schtschurowskiana Rgl. and Schm.
Plants less strong; leaves spinose-toothed; involucral bracts weaker,
inner bracts radiate, exceeding adjacent middle involucral bracts by
15—20 mm ......... 200. C. pseudodshisakensis Tschern. and Vved.
Leaves green and very finely arachnoid-hairy above, finely grayish-
tomentose beneath; basal leaves lyrate, with four or five pairs of
oblong lateral lobes; involucral bracts more or less flat...............
wet STE OC IS A. BERT INET oR: SS, 202. C. princeps Franch.
Leaves grayish-green and sparsely pubescent above densely tomentose
beneath; basal leaves lyrate, with five or six pairs of ovate or ovate-
oblong obess, bracts, carim ate oti. qcct ancien iteckverntne tt oceecte eccrine
10, TORE. Nae 205. C. mogoltavica Tschern. and Vved.
Leaves glabrous above; capitula glabrous; involucral bracts about
25; middle bracts more or less flat, broad lanceolate, 10-14 mm
wide at base, acuminate .................. 185. C. knorringiae Bornm.

169.

170.

17S:

174.

eS:

131

Leaves usually arachnoid-hairy above, if glabrous, then capitula
always white- or rusty-tomentose or arachnoid-hairy; involucral
bracts more numerous (40-100); middle bracts narrower, usually
GUSPId Aleta... .asihiad tech acctennes ieanshettisneed. «tester dee «qty edacenn tee cebeht 169.
Plants usually strongly branched; branches of common inflorescence
and peduncles slender, greatly elongate; capitula up to 50, 10-15
mm wide (excluding cusps); involucral bracts with 1.5—2 mm wide
CUSP Staten... Pceteenest wasceetebecs. pee adasedeak wed acsoet. smchatinan ssnaaddautes 170.
Plants usually weakly branched with rather thick branches of
common inflorescnece and peduncles; capitula usually not more
than 10-(15); capitula (13)15—35 mm wide, involucral bracts with
145—Summ mwide | cusps':(at. baSe) ..! aise eae. ak ele 2 ale
Cauline leaves terminating with relatively short and soft cusps;
capitula weakly arachnoid-hairy; involucral bracts about 50, with
somewhat short cusps, not as long as width of capitula...............
PUREE! O00), 10. STO) 20. Be ps Tit), oie 214. C. darwasica Winkl.
Cauline leaves terminating with very long and strong cusp; capitula
densely white-pubescent or rusty-tomentose; involucral bracts about
80, with long cusps, usually as long as or longer than width of
(OEY Oy UY Cia eh ey ae ee Rien ee 179. C. alpina Bge.

. Inner involucral bracts more or less distinctly broadened upward,

GatkeseOlOTci er ee eee a Meee 184. C. hilariae Kult.
Inner involucral bracts gradually narrowed in thin and short cusp;
stally We hits COMOKEG hie eae ee Teor ain zo) vaeaee «eeu a, Ve

. Cauline leaves sessile, short-decurrent; capitula strongly truncate

at base or even concave, densely rusty-tomentose; involucral bracts
numerous, about 100, with falcate Cusp ...........ccceccccsscetseeseeeeeees
£258 GNSSAREEE BEING EGS PoRAET Dna oe only: ees 212. C. ferruginea Kult.
Cauline leaves sessile, sometimes short-decurrent; capitula roundish
or weakly truncate at base; involucral bracts not so numerous, 40—

OL) eesti pele aa il pT so Ei Arts nb eri, Made tlet Mi Pg 173.
All involucral bracts erect; cauline leaves broad-lanceolate,
narrowed towards both ends................... 192. C. speciosa Wink.

Involucral bracts usually variable; outer bracts either decurved or
almost horizontal, middle often squarrose or horizontal, inner erect;
cauline leaves sessile, semiamplexicaul, usually broader at base
seta seaside uate wane ns acto ana Nie ag dau teind sede antetieta se casero arent, veered 174.
Stems sparsely branched, with long, usually almost leafless or
sparsely leafy, one-headed branches; capitula 20-35 mm wide

(OXIME. COSDS aos tie can aan 5 Baan ctees cele seesemaseaoe ie b75:
Stems branched, with short, usually leafy (often very dense) one-
headed branches; capitula 13-25 mm wide................c0000 179.

Leaves green and finely arachnoid-hairy above, grayish-arachnoid-
hairy or grayish-tomentose beneath; achenes 4—5 mm long .... 176.

132

152

176.

177.

178.

1S2)

Leaves green and glabrous above, white-tomentose or glabrous
beneath with flattened flexuous hairs only along midrib, or finely

sarachnGid=hatrys 2.2; oe ik cies vk. eee eee, een, oe 178.

Basal leaves almost bipinnatisect, petiolate, with usually dark
petioles; cauline leaves pinnatipartite, sessile, with small auricles
iobte Senden ed SEAS | SERRE | ROT 195. C. oxytoma Rech. f.
Basal leaves pinnatilobate, petiolate with short green petioles;
cauline leaves prickly-lobate or spinose-toothed, sessile ....... 177.
Capitula flat-globose, 20 mm wide (excluding cusps) weakly rusty-
tomentose; involucral bracts about 80, outer and middle bracts
15=23 immi long 92041. 200. oe. 196. C. ovezinnikowii Tschern.
Capitula broad-pyramidal, 20-25 mm wide, arachnoid-hairy;
involucral bracts almost horizontal, less numerous, about 50, outer
and middle bracts almost horizontal, 30-32 mm long..................
ncczainne Se EREAE OSES: « ots AUEMION, «RA. SANE, RN EON 193. C. batalinii Winkl.
Basal leaves oblong-ovate, undivided or more or less lyrate, with
large, broadly ovate terminal lobe and three or four pairs of wide,
sessile, oblong lateral lobes petiolate; petioles long, almost as long
as lamina, arachnoid-hairy like stem; achenes 7 mm long............
aes Ss FI, SR nn aA a Bae Sad beh al EN gi toe 197. C. peduncularis Juz.
Basal leaves pinnatisect into numerous (12-15 pairs) narrow-
lanceolate lateral segments, petiolate; petioles thickened near base,
densely pubescent with thick, flexuous, flattened hairs throughout,
like" lower halt“of*stemm 2000.2 eee 194. C. calva Juz.

. Outer and inner involucral bracts arcuately decurved or pendant

wit cet ta, enh ORE i IM, (oben Acde A. Cia Rea Re Se aD 190. C. fetissowii Wink].
Fey Oe a td MN re aks LMR NEL een Gem e e ae 180.
ae | MAND soninticd ck tS ae Mami, Se nL AN Rs OREN ND: oat A te aR Ne 181.

Capitula globose-cylindrical, 25—30 mm wide ..............:::cssceeeeeees
SEN Birt Mei Ltt MO: era ere MrCEat Sco me 191. C. schischkinii Juz.

. Basal leaves lyrate, with large, ovate, basally cordate lobe, finely

spinose-toothed along margin; capitula ovoid, 13-15 mm wide
(excluding “cusps) neh. e8. ag 183. C. buphtalmoides Rgl.
Basal leaves pinnatipartite or lyrate, then with oblong-roundish basal
lobe, cuneately excurrent on petiole; capitula ovoid-globose, 15—
OA VES AVES eee ee Se nemee EER Gan Ue NR Sore ee mana Genomes ee rere ee 182.
Cauline leaves sessile, short-decurrent; involucral bracts about 60,
inner bracts whitish; achenes about 4 mm long ............... eee
prvehiny rd sama RRS CAE area! Siok 180. C. carduncelloidea Rgl. and Schm.
Cauline leaves sessile, semiamplexicaul, not decurrent; involucral
bracts about 75, inner bracts often purple above; achenes about 4—
SPUTUM OME. AA MPRE eR anne 181. C. outichaschensis Franch.

183 (83)

133

+

184.

133

Basal leaves pinnatisect, with linear-lanceolate or almost linear
entire segments, usually appearing like whorl around almost entire
iniformalygw INP SUC MEtIOlS aco. .can- can eS caarainn Pemcmnesenon 184.
Basal leaves undivided or pinnatilobate, less often pinnatipartite,
then lobes deltoid, often boldly toothed or lobed, but never
appearinp slikeuwhorlses!...wiedsbiowdauan. kesares.. mea pA
Leaves soft, usually on both sides grayish and appressed-arachnoid-
hairy; lateral segments of leaves usually in one plane..................
ps aceeretin ins chewed circles cua lssboreatay eke 272. C. caespitosa Winkl.
Leaves usually stiff and coriaceous, if pubescent on both sides,
then. lateral segments of leaves never in one plane............... 185.

. Involucral bracts with long, falcate cusp; receptacular bristles

smooth; achenes wingless.............. 245. C. glaphyrocephala Juz.
Involucral bracts with erect cusp; receptacular bristles scabrous, if
smooth;«then,achenes .wingedl mcrs..tecdie.ataen!. ea 186.

. Plants 60-120 cm high; inflorescence spicate or at least with

tendency to form spikes, or almost paniculate, then achens 5—7
mm wide and 12-14 mm long, winged .............:::scceeeeeeeee Si.
Plants shorter; inflorescence never spicate.............. ccs 189.

. Involucral bracts smooth; achenes 5 mm long and 3 mm wide...

ae eee ee Seas vai nS sain euiguap tas coeun 248. C. pseudolanata M. Pop.
Involucral bracts scabrous due to spinulose surface; achenes 10—
14mm, Jong and 5—7 mmywide..... aiaves..:. genio. soe) 188.

. Leaves glabrous above; general inflorescnece spicate or subspicate;

receptacular bristles weakly scabrous or almost smooth ...............
pert. wteasark. peviiineen, ammo nccd . tyoc cue 247. C. lanata Winkl.
Leaves scabrous above from spinulose surface; common inflorescence
more or less paniculate; with erect spreading lateral branches;
receptacular bristles strongly scabrous.......... 246. C. magnifica Juz.

. Leaf segments 3-7 mm wide, somewhat flat; wings of petiole more

onedessiilat), well-developed... sis). Aversa... eterna. . feet 190.
Leaf segments 0.5—2 mm wide, usually recurved; wings of petiole
ASU ET ECO MLYC(EG Fav, SRM 2 a mame ete Ue OU lea ce 194.

. Leaves petiolate, scabrous from dense glandular hairs.......... Ki

Leaves entirely glabrous or with fine sessile glandular hairs
Scattered throuphotit .cemets, . anmeaerctste eyerss. pie ae eee) ie2.

. Stem arachnoid-hairy above; leaves green beneath or white-

tomentose only along margin; capitula finely white arachnoid-hairy;
involucral bracts sparsely glandular; corolla light yellow .............
eyesore, Ooarem ley. drteseltr ar. suerte: . Sepacuny 251. C. xanthina Bornm.
Stem sparsely glandular above; leaves white-tomentose beneath;
capitula rusty arachnoid-hairy; involucral bracts densely glandular;
Gorollappoulatish...225.....0ccc<e.00-. 2. 252. C. trachyphylla Juz.

134

134

193.

194.

. Capitula entirely glabrous; involucral bracts bright green, outer

and middle bracts ciliate along margin.............e cc eeeeeeecceeeeeeeeeeeees
gees Bi Ra se 5 Oe splendidavw ink!:
Capitula finely arachnoid-hairy; outer and middle involucral bracts
alwaysdentire), Mu) NOOR NS, Aes eae 193.
Stem appressed arachnoid-hairy; leaves usually with fine and
appressed pubescence above, less often glabrous, but always without
glandst2458. 2230. BUR) Bonen 254. C. stephanophora Winkl.
Stem glabrous or sparsely glandular above; leaves glabrous, smooth
or less often sparsely glandular, with fine sessile glandular hairs
paed di ahiubenssOnOATe RROD Woes ISI cats Baa eee cc 253. C. chrysantha Kult.
Most involucral bracts (except outer) broadened in upper part
forming wide membranous appendage or with wide membranous

Border W220 200k, SERS ERED dee ee. LLL, Mec, AEE E 195.
Involucral bracts without membranous appendage (excluding inner
Braet) 1.280. eee ee UN te Bees 196.
/2@orollaryellowr Ae. Ane eee 260. C. sororia Juz.
Corollai pink. 2... 28ers. Be ae 4 255. C. semilacera Juz.
. Capitula more or less densely arachnoid-hairy ..................0.. 197.
Capitula without arachnoid-hairy pubescence, glabrous or glandular-
SCA ROS LEON oP BMS. ccose nce venosccannsuonsdeswansossuties ta tacnsaadosean: 209.
. Sorolla: vellows 222.) Ny, BURR. aah LEONE 198.
Corolla. whitish. or. purple: ...007..29%. 3.25 Oee.. Be ae al. 204.
. Capitula 8-10 mm wide (excluding cusps)..............::eeeeee 199.
Capitula 12-18 mm wide (excluding cusps)...............0:seeee8 200.

. Stem profusely branched; branches spreading, usually longer than

main stem, and in turn divided into slender twigs, forming widely
spreading, almost corymbose inflorescence; inner involucral bracts
coriaceous, densely scabrous due to abundant spinules.................
NEE DO SOG. MELE, LOT, AOE, | 262. C. ramulosa Rech. f.
Stem sparingly branched into few, short, one headed branches;
inner involucral bracts scarious abOVE .............:ccesceeeeeeeeeeeeeeeeeeeees
sides Sins dace Mle des sdceed Saag MR Weta Nea 261. C. abbreviata Tschern.

. Involucral bracts few, about 35; capitula with curly, yellowish

glandular hairs ................... 256. C. allolepis Tschern. and Vved.
Involucral bracts more numerous, about 50-80; capitula arachnoid-
hairy gwithout, clands 22/80... .Q000e. VE ee ORE RR ee 201.

. Plants pubescent; leaves sericeous beneath............... cee 202.

Pubescence of plants mot SeriC€OUS..............:cceeceeeseeeeeeeeeeeeeeees 203.

. Leaves appressed-pubescent above, without glands or appressed-

clandulammai-ahuies.. 2ay68). ses. 257. C. newesskiana Winkl.
Leaves covered with papillate glandular hairs above alternating
with long, silky, simple hairs ............ 258. C. trichophora Kult.

135

203.

204.

207.

210:

211.

135

Stem erect, simple, strong; innet\involucral bracts with membranous
appendage, irregularly incised along margin...
1s, NPRORE. aktrarernt: Guests. obec, 6 263. C. acrodroma Tschern.
Stem relatively weak, usually branched; inner involucral bracts
without membranous appendage .......... 259. C. schugnanica Juz.
Outer involucral bracts sharply distinct from others, usually
divergent or almost horizontal, longer than middle bracts, as long
as or longer than width of capitulum.................. pevcairtnts Ney. M09.) 8

Outer involucral bracts shorter than or as long as middle, usually ©
TV cE SOMIN GE y ROTM LAGE i ees estae cine ompclenonejstncnttom aneahin ORE 205.

. Leaves glandular above; capitula 20 mm wide, densely ferruginous

‘FRROULGVIAL YO! af Serre), META) .oby 264. C. rubiginosa Kult.
Leaves without glands above, if with scattered glandular hairs,
then capitula always 8-(12) mm Wide .............cceceeeeeeeteeeeee 206.

. Leaves on both sides covered with very long, silky, entangled,

divergent hairs; all involucral bracts erect spreading ...................
BSNS REE REIE TOL SILO eh SOL os eee AEE, Bs 266. C. eriotricha Juz.
Leaves glabrous above or with appressed pubescence; involucral
bracts, especially outer and middle, divergent or sometimes even
honzZon tales sarcrcwascucaaswwg Re Md ase. 08, PORE, MUO LEY 207.
Stem densely white-tomentose; capitula densely white arachnoid-
hairy; achenes longitudinally narrow-ribbed .......0..... cee eeeeeeeeees
heed cRisselietiet ice cach brceiuslouaboomiienenier aula SINR 265. C. multiloba DC.
Stem finely tomentose, pubescent or arachnoid-hairy; capitula finely
arachnoid-hairy; achenes glabrous, smooth.................. sce 208.

. Capitula (8)12 mm wide; involucral bracts about 40 .........0000000...

M..AWOOe PAT RO IMO. 16 PANO. |e 269. C. bonvalotii Franch.
Capitula 15—22 mm wide; involucral bracts more numerous, up to
GORRY Bb oe OI Te AL oo RA SUR UE 268. C. verticillaris Bge.

. Outer and middle involucral bracts 5-10 mm wide near base,

glabrous, smooth, all except outer scabrous above dorsally .........
sew aul ak/anbe caren pent catras eM LRU, REL AUEEES 270. C. ceratophora Kult.
Outer and middle involucral bracts narrower, all glandular-scabrous
1s ERT, EY, DROID EE 10), SUNOS, NAG SET LL RG 210.
Achenes more or less 3-angled, narrowly winged along ribs; outer
involucral bracts erectspreading, usually nearly half as long as

minddlle ay etOCH. SSG NE ELL wae aL 249. C. adenophora Juz.
Achenes smooth; outer involucral bracts horizontal, usually longer
thantnmiddles).. UNGER eae 271. C. seravschanica Winkl.

Leaves soft, undivided without or with three to five scarcely visible
teeth along margin, less often pinnatilobate, or cauline leaves finely
SPINOSE=tOOtMEM (EG. 0 UR GRIN SIO, fai RG eo 212s

136

213.

214.

219.

Leaves usually more or less coriaceous, usually incised or lobate,
always ‘spinose:..1..gi@et Rater Ramee ales, eiahemeets DAS,

. Leaves tomentose on both sides, more densely beneath, cauline

leaves sessile; capitula scabrous, 12—13 mm wide (excluding cusps),
fernipinouset 0.2. RAS: jae 142. C. baranovii Juz.
Leaves usually finely arachnoid-hairy above, tomentose or densely
arachnoid-hairy beneath; cauline leaves decurrent; capitula ovoid
or cylindrical-ovoid, 7-13 mm wide (excluding cusps) ........ 213.
Stems numerous; cauline leaves short-decurrent; capitula 7-8 mm
wide (excluding cusps); inner involucral bracts usually pinkish
above w..c20 le ee ee ee 214.
Stems usually solitary or few; cauline leaves rather long-decurrent;
capitula 9-13 mm wide (excluding cusps); inner involucral bracts
notipigmentediins. ie. 2 138. C. leucantha Bornm. and Sint.
Involucral bracts 7-22 mm long, with long, subulate-deltoid, almost
horizontal cusps; achenes 4 mm long: sebis..ctevd oe eee tend... ee
Soenoenencte Sete en UnRe Senet ebro: Liana 137. C. stenophylla Kult.
Involucral bracts 5-14 mm long, with deltoid, erect spreading cusps;
achenes 4.5—5 mm long.... 136. C. komarowii (O. Ktze.) Winkl.

. Root usually many-headed, with thick caudices, usually covered

with woody remains of old petioles ................cccccccsseceeseeeessees 216.
Root many-headed, often with thick caudices, but without woody
remains of old petioles, sometimes with scarious remains ... 236.

t ®eceptaeulan bristles’ Smooth Noe ans cons enaseeeanaecneens a 20:
Receptacular: bristles) ‘scabrousa) 2a iret lee. ee 232.
. Plants (40)45—100 cm high; petioles of old leaves woody, usually
falcately decurved or divergent, almost horizontal ................ 218.
Plants 10—40(60) cm high; petioles of old leaves short, usually
CRESS E ease cals. eee Sat ed EN ok ee ee Ce Oe ieee 2D.

. Common inflorescence broad, lax corymbose; leaves nearly

glabrous and green above, white-tomentose beneath; capitula .
numerous, globose-ovoid, 6—8(10) mm wide and 15-18 mm long
(with erect florets); involucral bacts about 35 .............c ccs

Seabee es nemteene hte PE asa Rhrnsed Bene nl AAU end 3. C. linezewskii Juz.
Common inflorescence more or less spicate or paniculate; leaves
on both sides grayish-green, glandular and arachnoid-hairy or gray-
tomentose; capitula flat-globose, globose or cylindrical, 10-30 mm
wide (excluding cusps); involucral bracts numerous, sometimes
more} than! OOM chee Veh tere toon tere 212.
Lower cauline leaves oblong-ovate; capitula flat-globose, 20—30
mm wide (excluding cusps); middle involucral bracts 26-30 mm
long, more or less horizontal ................. 39. C. glochidiata Kult.
Lower cauline leaves oblong-lanceolate or oblong; capitula globose

137

220.

224.

Nagy,

137

or cylindrical, 10-20 mm wide (excluding cusps); middle involucral
bracts "erect spreading ‘or divergent \sie.f/202 <click. nccee-cceee tees PPA:
Inflorescence almost spicate; upper cauline leaves short-decurrent;
capitula cylindrical, 10-40 mm wide (excluding cusps); inner
involucral bracts lanceolate, long acuminate, 20-25 mm long...
As pole Seale op ah Daa ar ld 37. C. oreodoxa Bornm. and Sint.
Inflorescence paniculate; upper cauline leaves sessile, not decurrent;
capitula globose, 18—20 mm wide (excluding cusps); inner involucral
bracts somewhat broadened above, short-acuminate, about 20 mm
JO ek ete eal SSE Oe Sdn asl re hei 38. C. apiculata Tschern.

. Involucral bracts appressed, with erect cusp ..........:.cccceeeeee 222.

Involucral bracts with falcate or divergent, sometimes almost
herzontal wuspsnewer. settee een ee 225.

. Capitula 4-5 mm wide, with fewer florets ............. eee eeeeeeeeeeeees

nent i siteddlen ene: NAME EL OUR GENe ere 64. C. microcephala C.A.M.
Capitula 7-10 mm wide, with many florets ..............e eee 223)

. Capitula aggregated in groups of two to five at tips of branches,

somewhat enclosed by apical leaves; basal leaves oblong, pinnati-
lobate eee eat 0 63. C. orientalis (Adams) C. Koch
Capitula solitary terminal on branches, not enclosed by apical
leavess basal” leaves piinatisecte mies. ete te eee 224.
Capitula oblong-ovoid, 7-8 mm wide (excluding cusps); stem
moderately leafy, corymbosely branched; achenes five-ribbed .....
bb vi etd a SEE LEA Reem ate gee ETERS 53. C. chlorocephala C.A.M.
Capitula ovoid, 10 mm wide (excluding cusps); stems densely leafy,
with spreading branches; achenes usually 10-ribbed ............05......
MS tat in eh pe ee atl at BP ey 54. C. daralaghezica Takht.

. Leaves, at least basal, pinnatipartite or pinnatifid ................. 226.

EGAVes spinniatilo bate pasty: esse tele esctarsaeiroomtarverter hentnsnces, 229).

. Corolla purple; capitula 12-13 mm wide (excluding cusps).........

pha 3AM ee tee er RE ee th 57. C. brachyptera DC.
Corolla yellow, capitula 9-10 mm wide................:ceeeeeeee 227].

. Involucral bracts with straight, divergent cusps; capitula sparsely

AKACHMOM MALTY”. .22..5...c2cctteteere etnias 56. C. fedorovii Takht.
Involucral bracts with falcate or arcuately divergent cusps; capitula
weakly pubescent or sparsely arachnoid-hairy..................006 228:

. Cauline leaves very short-decurrent, or almost not decurrent;

capitula sparsely arachnoid-hairy; involucral bracts with falcate
CRISPS Samana teet eepa de 62. C. hohenackeri Fisch. and Mey.
Cauline leaves narrowly short-decurrent; capitula subglabrous;
involucral bracts with arcuate cusps......... 55. C. armena Takht.
Capitula 15-17 mm wide (excluding cusps); leaves grayish-green,
with network of prominent veins, gray arachnoid hairy-tomentose
on both sides, less often glabrous ..... 59. C. erivanensis Bornm.

138

138

234.

235.

Capitula 7-10 mm wide (excluding cusps); leaves glabrous on both
sides, sometimes finely glandular or finely arachnoid-hairy above,
and) eray-tomentose, beneath )..5 54... yao. ean coset ce-abcnoycenoe araraeee 230.

. Involucral bracts with cusps falcately decurved; leaves finely

plandularpabOVe: o.2...ccsosishos-+ ggace ses obs t eace 61. C. lomakinii Winkl.
Involucral bracts with divergent cusps; leaves always eglandular
abowe pgcsaghes eee Oy ON RS ae AN ea 231"

. Leaves glabrous on both sides; capitula glabrous ....................00

nN Mea ENG, CULT De, SE Nia 58. C. araxena Takht.
Leaves finely arachnoid- hairy above, gray-tomentose beneath;
Capitula, pubesce tts cs: saesdrorssacbhised Wena ercss 60. C. iljinii Takht.

. Common inflorescence, almost spicate or partially spicate, less often

compound spike ti). G6. LAAN EC dope ee) ote [sso
Common inflorescence, at least in upper part, with well-developed
lateral branches, i.e., racemose or paniculate ............... 234.

. Leaves green, very finely glandular or finely arachnoid-hairy above,

white-tomentose beneath; capitula oblong-cylindrical; achenes about
5 mm long; lower and middle involucral bracts with cusps far
exceeding (sometimes two times as long) capitulum ....................
sage Rigab bea 2 RIOR CEES RO 31. C. albertoregelia Winkl.
Leaves green and finely glandular on both sides, finely arachnoid-
hairy beneath; capitula broad ovoid; achenes 3.5 mm long; lower
and middle involucral bracts with cusps not exceeding capitulum
cy. Aik tease ea Bg So MRR QUO OES, 32. ctardisodnads 32. C. bobrovii Juz.
Leaves glabrous above, white or gray-tomentose beneath; capitula
more or less globose, roundish at base .............eeeeeeseeeeeees 235.
Leaves green on both sides, with sessile glandular hairs above;
capitula oblong-cylindrical, narrowed at base ...............eeeeeeeeeee
i UIE space truce sige Ma tae 35. C. stricta Tschern.
Root collar very strongly thickened, almost ovoid; involucral bracts
with cusps not exceeding capitulum............... 33. C. oopoda Juz.
Root collar weakly thickened; involucral bracts with cusps usually
one and one-half times as long as capitulum............ dee eee
Fs ts eer ee eee eee ee 34. C. nojnenclineeait Juz.

. Leaves more or less densely arachnoid-hairy or tomentose on both

sides, concolorous (white, grayish-green or glaucous)........... 237.
Leaves glabrous above or only weakly arachnoid-hairy, finely
arachnoid-hairy or densely white-tomentose beneath............. 254.
4 Receptaculan bristles smooth tiga. 20223: ssn eek eee 238.
Receptacular-bristles sscabrous 33350.) iahgec Sete scer ceert pee tage 251.

. Capitula 10-15 mm wide (excluding cusps); middle involucral

bracts two or three times as long as width of capitulum; less
numerous (about, 40) iene a atin tog: coos erctetece soars: ago:

240.

241.

242.

139

Capitula (12)15—-25 mm wide (excluding cusps); middle involucral
bracts shorter, usually not exceeding or only one and one-half times
as long as width of capitulum, numerous (about 60)............ 241.

. Basal leaves narrow lanceolate or oblong-lanceolate, pinnatipartite,

strongly spinose; lobes numerous, narrow-deltoid, each with long
and stiff spine, closely placed, petioles one-sixth to one-fifth as
long as lamina; capitula oblong-ovoid, 10-12 mm wide (excluding
GEIS) Meese see ess muta decode tesdenencsnecascrencesSaleaveasuerestemeeen eaten notaries 240.
Basal leaves petiolate, oblong-obovate or oblong-elliptical, lobed,
lobes broad; petiole one-fourth to one-third as long as lamina;
capitula larger, 12-15 mm wide (excluding cusp)................:000
sands tebe tN 2 hah Sie ABR a Lo a CR 113. C. gnezdilloi Tschern.
Basal leaves narrow and oblong-oblanceolate (4.5 cm wide
including cusps), pinnatipartite into numerous narrow-deltoid,
strongly spinose lateral lobes................ 114. C. leptoclada Kult.
Basal leaves oblong-lanceolate (8.5 cm wide including spines),
pinnatipartite into deltoid, weakly spinose lobes ...........000..
= iste eons RR nate ay oa 112. C. leptocladoides Tschern.
Middle involucral bracts 10—12(15) mm long, wider, with short,
up to 3 mm long, strong spine; lower cauline leaves lanceolate
eid CHR, dunia gen A Dearne baie oa le a hl ea 242.
Middle involucral bracts 15—25 mm long, comparatively narrow, with
long, 5—12 mm long, fine spine; lower cauline leaves oblanceolate,
oblong-oblanceolate, narrow-elliptical, oblong-elliptical or broad-
Der oi eC be aie te a a Sa amie rae Mea ee I Be A a ieee 243.
Terminal branches (floriferous) long, 4-6 cm long; leaves finely
arachnoid-hairy, grayish-green above, finely gray-tomentose
beneath; basal leaves minutely lobed with remote lobes; capitula
globose; middle involucral bracts up to 10 mm long ...................
Soria bedi totes Cestaite L418. bah eMac Re ate a eitdb rik | 108. C. proxima Juz.
Terminal branches (floriferous) short, 2—2.5 cm long; leaves white-

tomentose on both surfaces; basal leaves with large approximate

lobes; capitula broadly ovoid; middle involucral bracts longer, upto

DSMTITMLON Gee ee ee ee ee ee ee ecco 105. C. litvinovii Kult.
. Capitula ovoid or oblong-cylindrical, 12-18 mm wide (excluding
CEES PN) eres. eee eee Sere aes SE Cee CREE, MEET OI Sn 244.
Capitula globose or ovoid-globose, larger, 15-25 mm wide
CexCladinsMenspy ieee. ee esters erters deco ek cases cestnoieee oe 247.

. Terminal branches (floriferous) short, about 15 mm long; capitula

oblong-cylindrical, 12-15 mm wide (excluding cusp), aggregated
into heads; involucral bracts deltoid, acicular, erect ..........0.0........
tat Roan a a te ML taAL Ebel Ma 104. C. leptacma Tschern.
Terminal branches (floriferous) long, 3-12 cm long; capitula

140

141

140

247.

250.

oblong-ovoid, 12—20 mm wide (excluding cusps); inflorescence lax;
involucral bracts deltoid, subulate, weakly arcuate................ 245.

. Basal leaves short-petiolate or subsessile, oblong-lanceolate,

oblanceolate or oblong-elliptical ........... eee eee eeneeeneeeeeeeneeeees 246.
Basal leaves long-petiolate (petioles half to two-thirds as long as
lamina), broad-elliptical .................. 111. C. podophylla Tschern.

. Basal leaves subsessile, oblong-oblanceolate; capitula 17-20 mm

wide (excluding cusp) :..2082...02088.2 110. C. candicans Juz.
Basal leaves short-petiolate, oblanceolate or oblong-elliptical;
capitula 12-15 mm wide (excluding cusp) .............c:cccscccesseeeeneeees
Tee Sie cea ee saa Retaa tS BiB) 109. C. subcandicans Tschern.
Basal leaves very weakly spinose, boldly large toothed or minutely —
lobed; teeth or lobes with very short, up to 2 mm long, thin spine
SPIES, ERAS RES. GS, 2S oe eae ae 117. C. coerulea Kult.
Basal leaves strongly spinose, boldly toothed or lobed; lobes and
teeth with strong, 7-15 mm long spine ...............cc:ccesceeeeeeees 248.

. Capitula ovoid-globose or broadly ovoid; involucral bracts squarrose

on decuived inucrohiate’yisi.2.. 20 ee. Paty wel. acer 249.
Capitula globose; involucral bracts horizontal, abruptly acuminate
mani MOrUh sense setae sand te damhesee anh VaceaeZty aut ac baad ds eee Me een ABE 250.

. Basal leaves oblanceolate; capitula 18-25 mm wide (excluding

cusp); involucral bracts stiff; erect-spreading, spinescent with short
5, mimi longi spine e..4 ees er rem 106. C. amoena Winkl.
Basal leaves oblong-elliptical; capitula 15—17(20) mm wide
(excluding cusp); involucral bracts weak, falcate, spinescent with
7 mm_ Jong (spine =:...2..2 tee. 107. C. campyloraphis Tschern.
Lower and middle cauline leaves broad-elliptical, spinose-lobed,
10-12 cm wide (including spine) ..... 116. C. corymbosa Winkl.
Lower and middle cauline leaves oblanceolate, 5-6 cm wide
(including spine), pinnatipartite into long spinescent lobes ..........
Se ee Ne nie Seatac Oa, CS TAU 115. C. hoplophylla Tschern.

. Cauline leaves long-decurrent; stem simple or sparingly branched;

capitula one to fewer ...............06 99. C. semidecurrens Winkl.
Cauline leaves short-decurrent or almost nondecurrent; stem simple,
alwaysione=headed ack. A266 wit Monin eee, Descente is 252:

. Plants 25.60 cm high; capitula very densely ferruginous or white-

tomentose, almost globose; outer and middle involucral bracts
horizontal or slightly decurved, (15)20—25 mm long ....................
Pe eRe cei idee A 2 ult: UT Nee Reena, Ree 96. C. pannosa Winkl.
Plants 15—40 cm high; capitula sparsely ferruginous or white
arachnoid-hairy, cylindrical or ovoid; involucral bracts usually erect,
or (outer ones) horizontal, or slightly decurved, 10-15 mm long
Se ak ae ae MES, cs chs aha ae alia aL 1 an i Lm DD3:

142

258%

254.

255:

256.

259%

14]

Stem sparsely leafy; capitula ovoid, white arachnoid-hairy, 10-15
mm wide (excluding cusp); involucral bracts reduced, outer and
middle bracts 10-12 mm long; achenes glabrous, smooth, about 7
TATION SERIE «seek todas -<toovecsaveoncans 98. C. pannosiformis Tschern.
Stem densely leafy; capitula cylindrical, sparsely ferruginous-
arachnoid hairy, 15—18 mm wide (excluding cusp); involucral bracts
about 15 mm long; achenes indistinctly ribbed longitudinally, about

GEIL AR ce ieec sorte es aceacecteereseresveees 97. C. franchetii Winkl.
Achenes obpyramidal, three or four-ribbed, along ribs thickly
narrow-winged or narrow-winged, sharply toothed above..... JUDY

Achenes obovate, glabrous, smooth or with scarcely visible
longitudinal filiform ribs, roundish above or with two or three
IeLIStING tate ether: MOR IR ROU eset Se, 263.
Leaves green and glandular above, grayish-green beneath; capitula
strongly foveate at base, involucral bracts sparsely glandular
dorsally etter. shed Na eR: 19. C. glandulosa Kult.
Leaves green above, glabrous, glaucous or finely arachnoid-hairy,
eglandular; capitula roundish or somewhat foveate at base;
involucral bracts eglandular, arachnoid-hairy or glabrous, smooth
ORASCADLOUSOHN. «sasisvan nities nevivc Ree Ma tee NS. OREN: ROTA 256.
All involucral bracts scabrous dorsally, arachnoid-hairy, inner
involucral bracts membranous, dark colored, unevenly incised;
achenes 6—6.5 mm long............-eeeeeeeeee 36. C. sclerophylla Juz.
Involucral bracts usually smooth, glabrous or arachnoid-hairy,
membranous, scabrous along margin, usually light colored; inner
involucral bracts membranous, sometimes dorsally scabrous,

BCheENnES (3: Siam lone Ade HA hole ee RE 257:

. Capitula cylindrical or narrow-cylindrical, 4—6.5 mm wide

(excluding cusp), slightly extended at base... eee 258.
Capitula ovoid or globose, larger, 12—20 mm wide (excluding cusp),
somewhat foveate or roundish at base ..................c00seeeeeeeeeeeees 259.

. Inflorescence lax panicle; capitula 4 mm wide (excluding cusp) and

15 mm long (excluding corolla); corolla yellow; receptacular bristles
almost smooth or weakly scabrous; basal leaves pinnatipartite;
cauline leaves broadly short-decurrent ...... 17. C. cryptadena Juz.
Inflorescence usually raceme; capitula 5—6.5 mm wide and 18 mm
long; corolla whitish; receptacular bristles scabrous; basal leaves
crenate-spinose-toothed; cauline leaves sessile ..............c:ccceceeeees
dards asactianuchc-nletence:. 0, Ae eee! | 16. C. xanthiocephala Tschern.
Receptacular bristles smooth or with one or two crenations;
involucral bracts with falcate CUSPS ..........:ceccesceeceeeeeneeereeeeees 260.
Receptacular bristles scabrous or weakly scabrous; involucral bracts
WAGHOSGUM ATE OSE? CUSPS.......0.<cnncsecveenncvecesuteevewaccearecaesered Ssdue MAIS 262.

142

143

. Stem strongly divaricately branched in upper half; basal leaves

oblong-ovate, in well-developed many-leaved rosette; receptacular
bristles with one or two crenations; achenes 4—5 mm long .........
Tel eats sestsestteessteeseeeeeeee, 13 C, talassica (Kult.) Juz.
Stem weakly branched; basal leaves lanceolate, fewer in rosette;
receptacular bristles always smooth; achenes 3 mm long..... 261.

. Plants tall, 20-60 cm high; capitula 18-20 mm wide (excluding

CUSP)MSIOBOSEN) cure e ee a 12. C. affinis Schrenk
Plants shorter, 10-30 cm high; capitula smaller, 10-12 mm wide,
ovoidiets. janine. ois 11. C. astracanica (Spreng.) Tamamsch.

. Basal leaves oblong-lanceolate, crenate-lobed; capitula ovoid, 12

mm wide (excluding cusp); involucral bracts with divergent cusps,
middle bracts 12-15 mm long; achenes 5 mm long; receptacular
bristles; scabrousyieives: awn ee 14. C. pseudoaffinis Kult.
Basal leaves oblong-ovate or cuneately narrowed toward base,
oblong-oblanceolate, large spinose-toothed; capitula oblong-ovoid
or flat-globose, 10-14 mm wide; involucral bracts with squarrose
cusps, middle bracts 10-12 mm long; achenes 4—4.5 mm long;

receptacular bristles weakly scabrous......... 15. C. hamadae Juz.
Receptacular bristles scabrous...................0. 187. C. rava Winkl.
Receptacular bristles smooth ................c:cccscessssssseeeseeeseceeeesenes 264.
. Basal leaves ovate, undivided, crenate-lobed or with two or three
lobes near base, or more or less lyrate ..............c.cccccsceeseeeeeees 265.
Basal leaves pinnatisect or pinnatipartite .................:.ccceeeeee 266.

. Basal leaves more or less lyrate, very large, about 40 cm long and

25 cm. wide, green above, glabrous and lustrous; capitula 12—13
mm wide and 18-20 mm long (excluding projecting florets);
involucral bracts with membranous border, inner bracts without
membranous appendage; all bracts yellow ..............cccccssceeseeereeeeees
‘Sage RON WONG a POU TNE ToS ES NUNES ees 135. C. grandifolia Kult.
Basal leaves entire, crenate-lobed or with two or three lobes only
near base, 15—20 cm long and 5-6 cm wide, very finely arachnoid-
hairy; capitula 8 mm wide and 20-22 mm long; involucral bracts
with narrow membranous border; inner bracts with membranous,
unevenly incised, usually somewhat purple appendages; corolla
Ww hitishts ve ane hee eae ee 134. C. vavilovii Kult.

. Involucral bracts without appendages ..............:cccesesseseeeeeeeeeee 267.

All or at least inner involucral bracts with membranous or coriaceous
(and then case ciliate-toothed) appendages ................::scccceesee 268.

. Basal and lower cauline leaves 30 cm long and 12 cm wide (with

spines), long-petiolate; capitula aggregated at tips of branches and
stems in dense racemose inflorescence; outer involucral bracts
Hale ater sie nr eNel Cia nes nee er anne 93. C. abolini Kult.

144

143

+ Basal and lower cauline leaves 15-20 cm long and 2-3 cm wide,
short-petiolate; capitula solitary terminal on stem and branches;
outer involucral bracts divergent ........ 103. -C. macilenta Winkl.

268. Capitula in lax, usually paniculate, inflorescence, 10-17 mm wide
(excluding cusp), ovoid or narrow-ovoid; basal and lower cauline
leaves oblong-lanceolate, 25-30 cm long and 5-11 cm wide......
Uy, Denar eee aie bee edie eae dbxesse didi renin eo election, fH 209.

+ Capitula solitary terminal on branches and stem, sometimes in more
or less corymbose inflorescence, 18-28 mm wide (excluding cusp),
globose; broadly ovoid or somewhat cylindrical; basal and lower
cauline leaves 14-60 cm long and 3.5—6 cm wide ............... 270.

269. Capitula glabrous, lustrous, 13-17 mm wide (excluding cusp); inner
involucral bracts almost roundish above, entire; leaves finely

_arachnoid-hairy above............... ARIES). Pe 94. C. egregia Juz.

+ Capitula arachnoid-hairy, matty, 10-12 mm wide; inner involucral
bracts spatulate, small-toothed, usually incised in middle into two
ALVES ee fen eee eee 95. C. dolichophylla Kult.

270. Plants 100-120 cm high; inflorescence corymbose; leaves glabrous
above, green; basal and lower cauline leaves about 40-60 cm long
CMAMOMEM” WAUE 5. Stes erecceesetreurer toes: 101. C. korshinskyi Winkl.

+ Plants 35-60 cm high; stem one-headed or seldom with two or
three one-headed branches; leaves finely arachnoid-hairy above;
basal and lower cauline leaves about 14-25 cm long and 3.5-5
CIM WUC scare ccs coceces JsSielhcresecet sper tat cna hana Mie el, duapechornn.» Abana DA

271. Lateral lobes of basal leaves with one or two pairs of spinescent
lobes at base; cauline leaves sessile, semiamplexicaul; all involucral
bracts (except inner) with coriaceous appendages; appendages
CilALCCOOHMEG, neo cectsnccGatarceus ues 100. C. fedtschenkoana Bornm.

+ Lateral lobes of basal leaves deltoid, along margin spiny-lobed;
cauline leaves sessile, short and broadly decurrent; all involucral
bracts (except outermost) with membranous, almost transparent
appendages; appendages irregularly toothed along margin............
Sire tues een | Septias dan kad bates oaata 102. C. pterolepida Kult.

Section 1. Badghysia Tschern. in Bot. Mat. Gerb., Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 77.—Capitula solitary terminal on one-
year-old stems; involucral bracts gradually narrowed into thin, usually
strongly falcate cusp; corolla yellow; receptacular bristles more or less
smooth; achenes three or four-angled, toothed above. Semishrubs with
thick, strongly flexuously branched woody stems; one-year old stems
whitish, with short branches.

Type of section: C. badghysi Kult.

145

144

1. C. badghysi Kult. in Pochv. i Bot.-Geogr. Iss]. Bass. rr. Amu- i
Syr-Dari, 2 (1916) 76; Kult. in Fl. Turkm. VII, 189.—Ic.: Kult. op. cit.

- (1916), Plates 23a, 23b.

Cultivated. Stem thick, short, strongly flexuously branched, covered
below with gray, shredded dead bark; one-year old floriferous shoots
numerous, short-branched above, covered with smooth whitish bark,
densely white-tomentose when young, glabrescent. Leaves of woody
branches sessile, in whorls of three to six; leaves of floriferous shoots
alternate, one or two, oblong-lanceolate, 3-8 cm long and 3—5 mm wide,
with small, narrow-lanceolate, short acuminate, remote lobes, all leaves
coriaceous, recurved, grayish-green, sparsely pubescent, glabrescent
above, densely white-tomentose beneath, gradually reduced upward on
stem and branches. Capitula narrow-ovoid, 7-9 mm wide (excluding
cusp), 15-16 mm long (excluding florets), finely arachnoid-hairy.
Involucral bracts 75, deltoidly broadened in lower part, closely appressed,
scabrous along margin, gradually attenuate toward apex into thin,
subulate, strongly falcate cusp; inner bracts coriaceous, linear-lanceolate,
acuminate into thin cusp, scabrous along margin; receptacular bristles
unequal, longest slightly broadened above, very conspicuously serrate,
almost smooth. Corolla yellow; anther tubes yellow. Achenes 4 mm
long and 2 mm wide, three or four-angled, three or four-toothed above,
rugose, light gray, lustrous, with black spots. Flowering VI.

Loamy-gravelly mountain slopes.—Soviet Central Asia: mountainous
Turkmenia (Badkhyz). Endemic. Described from the vicinity of Lake
Er-Oilan. Type in Tashkent, isotype in Leningrad.

Section 2. Leucocaulon Tschern. in Bot. Mat. Gerb., Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 77.—Orthacanthae Winkl. in Tr. Peterb.
Bot. Sada, XII (1892) 214 p. p.——Capitula numerous, in lax corymbose
inflorescence or solitary, terminal on slender branches. Involucral bracts
few, flat, short or very short and sharp-acuminate. Corolla pale yellowish
or whitish. Receptacular bristles smooth. Achenes tetraquetrous,
obpyramidal, with small teeth above. Perennials with thick root collar,
covered with remains of old petioles and coriaceous, sessile, cauline
leaves or semishrubs with woody, basally warped and glabrous, whitish
one-year old stems.

Type of section: C. turcomanica Winkl.

2. C. turcomanica Winkl. in Tr. Peterb. Bot. Sada, XI (1889) 133;
Winkl. Ibid. XII, 217; Kult. in Fl. Turkm. VII, 182.—Serratula
microcephala Trautv. in Tr. Peterb. Bot. Sada, IX (1886) 458.—<Arctium
turcomanicum QO. Ktze. Rev. Gen. 1 (1891) 308.—TIc.: Winkl. op. cit.
(1889), Plate 2, Fig. 1.

146

145

Cultivated. Semishrub with woody, basally warped stems; one-year
old stems numerous, finely arachnoid-hairy, glabrescent, white, glossy,
leafy, weakly branched in upper half, with long, slender, usually one-
headed branches. Leaves hard, coriaceous, with network of prominent
veins, green, punctate with fine glandular hairs above, finely arachnoid-
hairy beneath, entire or minutely prickly-toothed, narrow-lanceolate or
more or less lanceolate, narrowed toward both ends into short petiole
below, and short cusp above; lower leaves 4—5 cm long, petiolate
(petioles 8-12 mm long), other leaves gradually reduced upward, with
shorter petioles; uppermost leaves sessile, strongly reduced. Capitula 5
mm wide and 20 mm long with projecting florets, almost glabrous.
Involucral bracts closely appressed, with short erect cusp, weakly
scabrous in upper part or dorsally due to fine glandular hairs, very finely
arachnoid-hairy, ovate-lanceolate or somewhat lanceolate; inner involucral
bracts membranous, brownish, linear-lanceolate, scabrous along margin,
mucronate. Receptacular bristles smooth. Corolla whitish; anther tubes
purple. Achenes narrow-pyramidal, 3.5 mm long and 1—1.5 mm wide,
narrowly ribbed, with three small teeth above. Flowering VI; fruiting
VU-VIil.

Clayey and stony debris foothill slopes.—Soviet Central Asia:
mountainous Turkmenia. General distribution: Iran. Described ‘from
Kopetdag, near Germab and Geoktepe. Type in Leningrad.

Note. Two localities were reported in the description of this species;
a plant with the following label is proposed as the lectotype: “Montes
Kopetdagh prope Germab et Geok-Tepe. Antonow.”

3. C. linezewskii Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3
(1936) 304.

Perennials. Roots woody, black-fibrous. Stem erect, 60-90 cm high,
with numerous remains of old petioles near base, whitish, glossy,
cylindrical, sparsely branched in upper half; branches and twigs long,
slender, forming wide lax corymbose inflorescence. Leaves coriaceous,
green above, almost glabrous, densely white-tomentose beneath; basal
leaves petiolate, with 3 cm long whitish petioles deltoidly broadened at
base, oblong or elliptical, slightly narrowed towards both ends; lamina 16—
25 cm long and 5.6—9 cm wide, crispate-lobed along margin; lobes seven
or eight on each side, broadly lanceolate or almost semicircular, prickly-
toothed and prickly-acuminate; cauline leaves abruptly reduced, lower
cauline leaves similar to basal leaves, other sessile, prickly-toothed,
uppermost cauline leaves strongly reduced. Capitula numerous, globose-
ovoid, 6—8(10) mm wide, 15—18 mm long (with projecting florets), sparsely
glandular and very weakly arachnoid-hairy. Involucral bracts about 35,
flat, ovate to lanceolate-ovate, closely appressed, with short, thin, weakly

147

146

protruding cusp; inner bracts roundish-obtuse, broadened above, cuspidate,
weakly scabrous along margin. Receptacular bristles smooth, whitish.
Corolla pale yellow, tube glandular. Achenes obpyramidal, with narrow
longitudinal ribs, with small teeth above, dark brown with blackish spots,
44.5 cm long. Flowering VI; fruiting VII—VIII.

Stony slopes.—Soviet Central Asia: mountainous Turkmenia
(Badkhyz, southeastern Kopetdag). Endemic. Described from Badkhyz
in the Pul-i-Khatum District. Type in Leningrad.

Section 3. Leptocephalae Tschern. in Bot. Mat. Gerb., Inst. Bot.
Akad. Nauk UzbSSR, XVII (1962) 77.—Stenocephalae Bge. in Mém.
Acad. Pétersb. 7 sér., 9, 2 (1865) 24 p. p.—Squarrosae Boiss. FI. or. III
(1875) 460 p. p.—Brachyacanthae Winkl. in Tr. Peterb. Bot. Sada. XII
(1892) 204 p. p.—Capitula solitary terminal on branches or aggregated,
ovoid or narrow cylindrical. Involucral bracts gradually acuminate in
long or short, falcate or erect thin cusp. Receptacular bristles smooth,
less often scabrous. Corolla pink, whitish, yellow or flesh-red; anther
tubes glabrous. Achenes obovoid or obpyramidal, almost triquetrous,
narrowly winged along margin.

Semishrubs, usually with sessile or narrow or broadly decurrent
cauline leaves.

Type of section: C. leptocephala Fisch. and Mey.

Series 1. Leptocephalae Tschern.—Capitula small, aggregated;
receptacular bristles unequal, longest bristles scabrous. Cauline leaves
narrowly long-decurrent.

4. C. leptocephala Fisch. and Mey. in Bull. Soc. Nat. Mosc. XII
(1839) 159 (nomen) and ex Bge. in Mém. Acad. Pétersb. 7 sér., 9, 2,
25; Boiss. Fl. or. III, 487; Winkl. in Tr. Peterb. Bot. Sada, XI, 135; XII,
205; XIV, 220; Kult. in Fl. Turkm. VII, 203—Exs.: H.F.A.M. No. 46.

Cultivated. Short, many-stemmed, densely leafy semishrub,
caespitose. Stem white-tomentose, 10-15 cm high, cymosely branched
above, with short branches. Leaves hard coriaceous, grayish-green above
due to finely arachnoid-hairy pubescence, finely gray-tomentose beneath;
lower cauline leaves petiolate, narrow oblong-lanceolate, 8-9 cm long,
8 mm (at notches), 15-17 mm (with lobes) wide, narrowed toward both
ends, crenate-lobed, with narrow deltoid and spinescent lobes; other
cauline leaves similar, but gradually somewhat reduced; middle leaves
short-petiolate; upper sessile, narrowly long-decurrent, 2—2.5 cm long.
Capitula aggregated, on short peduncles, narrow-cylindrical, 4-5 mm
wide (excluding cusps) and 10-12 mm long (excluding florets),
arachnoid-hairy. Involucral bracts about 50, imbricate, closely appressed,

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apically narrowed in thin, falcate cusp; inner bracts membranous, narrow-
lanceolate, short-pointed into thin, erect cusp, scabrous along margin
dorsally; receptacular bristles unequal, longest slightly broadened above,
scabrous, shorter, smooth. Corolla pink or purple; anther tube purple.
Achenes obovoid, 3—3.5 mm long, denticulate above. Flowering . VII—
VII; fruiting VIII-IX.

Mountain slopes.—Soviet Central Asia: mountainous Turkmenia
(Bolshie Balkhany). Endemic. Described from Bolshie Balkhany. Type
in Leningrad.

Series 2. Hypopoliae Tschern.—Capitula aggregated or solitary
terminal on branches. Receptacular bristles smooth. Cauline leaves
broadly long-decurrent.

- 5. C. albiflora (Bornm. and Sint.) Bornm. in Beih. bot. Centralbl.
XXXIV, 2 (1916) 285; Kult. in Fl. Turkm. VII, 201—C. hypopolia B.
albiflora Bornm. and Sint. in Bull. Herb. Boiss. 2 sér. 7 (1907) 215.—
C. lepida auct. non Bge.: Winkl. in Tr. Peterb. Bot. Sada, XI (1890)
136; Fedtsch. Perech. Rast. Turk. IV, 249.—Ic.: Bornm. and Sint. op.
cit. tab. 7, Fig. 2.

Cultivated. Short many-stemmed semishrub. Stems erect, almost
tetraquetrous, white arachnoid-hairy, leafy, usually branched from middle.
Capitula aggregated at stem and branch tips. Leaves coriaceous, greenish
above, usually finely arachnoid-hairy, with fine network of somewhat
prominent veins, white-tomentose beneath; lower cauline leaves usually
appearing rosulate, petiolate, 10-15 cm long, 44.5 cm wide, crenate-
toothed, with prickly-acuminate teeth, broad lanceolate, narrowed toward
both ends, spinescent; other leaves gradually reduced, sessile, decurrent,
otherwise similar to lower leaves. Capitula on short peduncles, 8-9 mm
long (excluding florets) and 3.5-4 mm wide (excluding cusp), very finely
arachnoid-hairy, ovoid-oval. Involucral bracts 60, all except inner closely
appressed, ovate to oblong-ovate, apically narrowed in very short, slightly
divergent cusp; inner involucral bracts membranous, narrow lanceolate,
shortly spinescent, scabrous along margin; receptacular bristles smooth,
whitish. Corolla whitish; anther tube pink. Achenes obovoid, 3-4 mm
long, ribbed, with small teeth above. Flowering VI-VII; fruiting VII-—
VUI. (Plate VIII, Fig. 1).

Gravelly slopes.—Soviet Central Asia: mountainous Turkmenia
(Kopetdag). Endemic. Described from Kara-Kal, Sundzodag Mountain.
Type in Weimar.

Note. Plants from Syunt are transitional to C. hypopolia Bornm.,
although in their characters they are closer to C. albiflora and need to
be examined in nature.

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148

6. C. hypopolia Bornm. and. Sint. in Bull. Herb. Boiss. II sér. 7
(1907) 214; Bornm. in Beih. bot. Centralbl. XXXIV, 2, 285; Kult. in
Fl. Turkm. VII, 202.—Ic.: Bornm. and Sint. op. cit. tab. 7, fig. 4; Kult.
op. cit. Plate 28, Fig. 2.

Cultivated. Short many-stemmed semishrubs. Stems erect, white-
arachnoid, densely leafy, cymosely branched above. Leaves coriaceous,
green, glabrous, glossy or finely arachnoid-hairy above, gray-tomentose
beneath with prominent veins; lower leaves petiolate, usually smaller
than middle, 8-10 cm long, 2-3 cm wide, broad-lanceolate, somewhat
narrowed toward both ends, with shallow and wide, acute crenation along
margin; middle leaves ovate-lanceolate, 12-14 cm long, 5 cm wide,
sessile, broadly relatively short-decurrent (over 2—2.5 cm), with roundish
auricles; upper leaves like middle, but shorter, 4-5 cm long; uppermost
leaves reduced. Capitula one to three, terminal on each branch and stem,
on peduncles, 5—6 mm wide (excluding cusp), 12 mm long (excluding
florets), finely arachnoid-hairy. Involucral bracts about 60, all except
inner imbricate, ovate to oblong-ovate, 2-7 mm long, closely appressed,
acuminate into somewhat short, thin, almost horizontal or weakly falcate
cusp; inner involucral bracts narrow-lanceolate, erect, short spinescent,
scabrous dorsally and along margin, 12—-12.5 mm long. Receptacular
bristles unequal, smooth. Corolla yellow; anther tubes purple or pink.
Achenes obpyramidal, 5 mm long and 2 mm wide, tetraquetrous, narrow-
winged along angles, toothed above. Flowering VII—VIII; fruiting VIII.

Debris, slopes in middle mountain zone.—Soviet Central Asia:
mountainous Turkmenia (Kopetdag). General distribution: Northern Iran.
Described from Kopetdag. Sulyuklyu (Saratovka). Isotype in Leningrad.

7. C. mucida Kult. in Bot. Mat. Gerb. Inst. Bot. Akad. Nauk
UzbSSR XVII (1962) 97.

Cultivated. Short many-headed semishrub. Stems erect, almost
tetraquetrous, white arachnoid-hairy, leafy, usually simple, one-headed.
Leaves coriaceous, finely gray-tomentose on both sides, more dense
beneath, with fine network of weakly prominent veins, all leaves ovate
or oblong-ovate, narrowed toward both ends, almost entire or very
inconspicuously and finely spinose-toothed, with short apical spine; basal
and lower cauline leaves petiolate, others sessile, decurrent on stem.
Capitula 7-8 mm wide (excluding cusps), finely arachnoid-hairy, ovoid.
Involucral bracts ovate to oblong-ovate, all closely appressed, apically
narrowed in thin, falcate cusp; inner involucral bracts membranous,
narrow lanceolate, finely prickly-acuminate, scabrous along margin.
Receptacular bristles smooth. Corolla yellow. Achenes obovoid, ribbed,
toothed above, 4 mm long. Flowering VII.

Debris slopes.—Soviet Central Asia: mountainous Turkmenia

149

Plate VIII.
1—Cousinia albiflora (Bornm. and Sint.) Bornm.; 2—C. affinis Schrenk.

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150

(Kopetdag). Endemic. Described. from the Misinev Range. Type in
Leningrad. —

8. C. stahliana Bornm. and Gauba in Mitt. Thir. Bot. Ver. 47
(1941) 117.—C. chejrabadensis Kult. in Fl. Turkm. VII (1960) 380.—
C. hypopolia auct. non Bornm. and Sint.: Kult. in Herb. Fl. As. Med.
No. 45.—Exs.: H.F.A.M. No. 45.—Ic.: Bornm. and Gauba, op. cit. tab.
13.

Cultivated, short many-stemmed semishrub, 20-30 cm high. Stems
erect, finely white arachnoid-hairy, densely leafy, sparingly short-
branched above; branches short one-headed, less often two or three-
headed, usually entirely enclosed by apical leaves. Leaves strongly
coriaceous, gray arachnoid-hairy on both sides, more densely beneath,
with prominent veins; basal leaves petiolate, broad oblong, crenate-
prickly-toothed acuminate in strong divergent cusp at apex; cauline leaves
very insignificantly reduced toward stem apex; upper leaves sessile,
decurrent. Capitula ovoid, 8-9 mm wide (excluding cusps), arachnoid-
hairy. Involucral bracts, except inner, narrow-lanceolate, closely appressed
at base, free in middle part, somewhat arcuately decurved, short
acuminate; inner bracts narrow-lanceolate, erect. Receptacular bristles
smooth. Corolla yellow; anther tube pale yellow or light pink. Achenes
about 4 mm long, obpyramidal, ribbed, toothed above. Flowering VI—
VII; fruiting VII—VIII.

Dry slopes in upper mountain zone.— Soviet Central Asia: mountainous
Turkmenia. General distribution: Northern Iran. Described from region
between Shirvan and Ashkhabad. Type in Weimar.

9. C. oreoxerophila Kult. Fl. Turkm. VII (1960) 380.

Cultivated, many-stemmed semishrub, 20-30 cm high, with numerous
caudices at base. Stems numerous, leafy, with few short branches above,
2-8 cm long lateral branches appressed white arachnoid-hairy, pubescence
sparse above and dense below; bark pinkish. Leaves light grayish-green,
coriaceous, dense, with thin remote spines along margin and very
prominent veins on both sides; basal leaves about 10 cm long, 1.5—2.5
cm wide, long-petiolate, with 5 cm long petiole, ovate-lanceolate,
cuspidate, sparsely and finely spinose along margin, with 3-4 mm long
spines; middle cauline leaves like basal leaves, but short-petiolate or
sessile, with larger lamina; upper cauline leaves ovate-oblong, sessile
with broad base, weakly decurrent, broadly crenate-prickly-toothed along
margin; uppermost leaves 2.5 cm long and 1.5 cm wide, reaching almost
up to capitulum base, like upper. Capitula ovoid, 12 mm wide (excluding
cusps) and 20 mm (including cusps), solitary terminal on short or long
branches, arachnoid-hairy, pale green. Involucral bracts numerous (more

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than 60), closely appressed at base,‘with almost horizontal cusp; outer
bracts 4 x 2, in upper part about 5 mm long; middle bracts 8 mm long,
3 mm wide, in upper part 6 mm long; inner bracts 15 mm long, 2 mm
wide, linear, coriaceous, scabrous along margin, gradually narrowed
toward apex. Receptacular bristles 10 mm long, smooth. Corolla 13 mm
long, flesh-red (?); anther tube pink. Achenes 4 mm long, 1.5 mm wide,
dark brown, obpyramidal, with unequal teeth above. Flowering VI-VIII.

Upper mountain zone.—Soviet Central Asia: mountainous Turkmenia
(Kopetdag). Endemic. Described from the Dogri-Dere Ravine. Type in
Ashkhabad.

Note. A very doubtful species. It is described here for the first
time, as no material for this species could be found. The type, as
indicated by the author, should be in Ashkhadad.

Series 3. Chaetocephalae Tschern.—Capitula solitary terminal on
branches; receptacular bristles smooth. Cauline leaves sessile, short-.
decurrent.

10. C. chaetocephala Kult. in Tr. Sredneaz. Gos. Univ., Ser. VIIIc,
7(1929) 3.—Ie.: Kult. Ibid., Plate 3.
Cultivated, short many-stemmed semishrub. Stems 35—40 high, erect,

‘arachnoid-hairy, moderately leafy, branched above, branches short, one-

headed. Leaves coriaceous, grayish-green on both sides, finely arachnoid-
hairy (more densely beneath), with prominent veins, oblong-lanceolate,
crenate-prickly-toothed; lower leaves petiolate, with petioles almost as
long as lamina, narrowed towards both ends, spinescent, 7.5—8.5 cm
long including petiole, 2 cm wide including spines; middle leaves sessile,
semiamplexicaul, short-decurrent somewhat shorter than lower; upper
leaves reduced, narrow-lanceolate, sessile, short-decurrent. Capitula ovoid
or ovoid-oblong, finely arachnoid-hairy, 8-10 mm wide (excluding
cusps), 18 mm long (excluding florets). Involucral bracts about 60, all
except inner closely appressed at base, with thin, -subulate, deltoid,
falcately recurved cusp; ovate to oblong-lanceolate; inner bracts flat,
membranous, linear, 15—17 mm long, short and thin acuminate, scabrous
above and along margin. Receptacular bristles somewhat broadened
above, smooth, creamish. Corolla yellow; anther tube purple. Achenes
obpyramidal, triquetrous, with weakly thickened ribs, toothed above, 5
mm long and 2 mm wide. Flowering VII—VIII; fruiting VIII.

Stony slopes.—Soviet Central Asia: mountainous Turkmenia
(Kopetdag). General distribution: Northern Iran. Described from
Khorassan. Type in Tashkent.

Section 4. Leiocaules Bge. in Mém. Acad. Pétersb. 7 sér., 9, 2 (1865)
42 p. max. p—Sphaerocephalae Beg. Ibid. (1865) 35 p. p.—Squarrosae

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Boiss. FI. or. III (1875) 460 p..p——Hamatae Boiss. Ibid. (1875) 461.—
Drepanophorae Winkl. in Tr. Peterb. Bot. Sada, XII (1892) 206 p. p.;
Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8 (1940) 550 p. p.—
Nudicaules Winkl. op. cit. (1892) 191 p. p—Heteracanthae Winkl. op.
cit. 225 p. p—Capitula solitary terminal on branches, numerous or few;
involucral bracts ovate, with strong, arcuate or almost erect, long or short
cusp; corolla yellow or whitish; receptacular bristles smooth or scabrous;
achenes obovoid or obpyramidal, three or four-angled, toothed above.
Perennial herbs with woody many-headed root, thick collar, and sessile
cauline leaves, or semishrubs.
Lectotype of section: C. boissieri Buhse.

11. C. astracanica (Spreng.) Tamamsch. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XVI (1954) 468.—Carduus astracanicus Spreng.
Fl. Hal. Mant. Prim. (1807) 49.—Carduus wolgensis MB. in Willd.
Enum. 2 (1809) 839; Spreng. Syst.-veg. 3 (1826) 385.—Cousinia
wolgensis C.A.M. in DC. Prodr. VI (1837) 555; Ldb. Fl. Ross. II, 678;
Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2 36; Fl. Yugo-Vost. VI, 384;
Pavlov, Fl. Tsentr. Kazakhst. III, 297.—C. transcaspica Juz. ex Kult. in
Fl. Turkm. VII (1960) 210 (descr. ross.).—Arctium wolgense O. Ktze.
in Tr. Peterb. Bot. Sada, X (1887) 198.—ic.: MB. Cent. Pl. Rar. 2
(1810) tab. 44; Fl. Yugo-Vost. 697.

Perennial. Roots woody, blackish-fibrous, with thick, densely white-
tomentose collar, covered with remains of old leaves. Stem erect,
arachnoid hairy-tomentose, with somewhat floccose tomentum, more or
less cymosely branched, 10-30 cm high. Leaves green above, glabrous,
white-tomentose beneath; basal and lower cauline leaves oblong,
petiolate, crenate-toothed, teeth finely prickly-acuminate; other cauline
leaves gradually reduced upward, sessile, semiamplexicaul, oblong-ovate,
or ovate, or uppermost leaves more or less orbicular, shallow crenate-
toothed. Capitula fewer, solitary terminal on short branches, ovoid, 10—
12 mm wide (excluding cusps), arachnoid-hairy. Involucral bracts
numerous, ovate to oblong, with strong, subulate, relatively long,
arcuately recurved cusp; inner involucral bracts thin-coriaceous, with
short and thin cusp, scabrous along margin, usually whitish. Receptacular
bristles smooth, helical below. Corolla whitish or yellowish. Achenes
obpyramidal, more or less tetraquetrous, with narrow wings along ribs
and short toothed above. Flowering VI; fruiting VI-VII.

Clayey-stony slopes——European Part: Lower Volga; Soviet Central
Asia: Aralo-Caspian, Kara-Kum. Endemic. Described from Sarepta. Type
in Berlin?

12. C. affinis Schrenk, Enum. pl. I (1841) 41; Ldb. Fl. Ross. II,
678; Bge. in Mém Acad. Pétersb., 7 sér. 9, 2, 36; Boiss. Fl. or. III, 488;

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153

Winkl. in Tr. Peterb. Bot. Sada, XII, 212; XIV, 221; Pavlov, Fl. Tsentr.
Kazakhst. III, 297; Kryl. Fl. Zap. Sib. XI, 2883; Grubov, Konsp. FI.
Mong. 273.—C. wolgensis var. affinis Rgl. in Tr. Peterb. Bot. Sada, VI
(1879) 317.—Arctium affine O. Ktze. Rev. Gen. 1 (1891) 307.

Perennial. Root fusiform, blackish-fibrous, woody, with usually thick
whitish-fibrous collar. Stem 20-60 cm high, erect, arachnoid-hairy,
glabrescent, light colored, almost smooth, usually solitary, weakly
branched above, branches one-headed. Leaves very finely arachnoid-
hairy, green above, gray-tomentose beneath, large prickly-toothed or
lobed along margin; basal leaves lanceolate, short-petiolate lobed, lobes
remote, spinescent; lower cauline leaves like basal, but sessile; middle
cauline leaves oblong, sessile, with roundish auricles, lobed; upper leaves
ovate, sessile, large prickly-toothed; uppermost leaves reduced. Capitula
globose, 18-20 mm wide (excluding cusps), weakly arachnoid-hairy.
Involucral bracts more numerous, imbricate, closely appressed near base,
apically narrowed in subulate falcate cusp; outer bracts half as long as
middle; middle bracts 1.5—2 cm long; inner bracts membranous,
somewhat flat, lanceolate, gradually prickly-acuminate, 2—2.5 cm long.
Receptacular bristles smooth. Corolla pale yellow or whitish; anther tube
purple. Achenes obovoid, three or four-angled, thickened along angles,
grayish, dark spotted, toothed above, 3 mm long. Flowering VI—VII;
fruiting VII-VIII. (Plate VIII, Fig. 2).

Along clayey and sandy knolls, at the outcrops of bedrocks in deserts
and semideserts.—Soviet Central Asia: Balkhash Region, Aralo-Caspian,
Kyzyl-Kum, Tien Shan (Trans-Ili Alatau, Chu-Ili mountains). General
distribution: Dzhungaria, Mongolia. Described from the Lepsa District.
Type in Leningrad.

Note. The two northernmost species of the genus, C. astracanica
(Spreng.) Tamamsch. and C. affinis Schrenk, with quite wide and
contiguous ranges, are well distinguished only in extreme forms. An
examination of quite large collections of these species shows that most
of the characters used to distinguish them are extremely variable. A
detailed study of the abundant material collected from different sites in
the distribution areas is essential to determine the constancy of one
character or another; the material of these species in herbaria does not
make it possible to determine the stability of the characters.

It is quite possible that we are dealing with a single, widespread
species that, depending on the conditions of growth, varies slightly in
its appearance. It has been found that the plants growing in the northern
deserts on gravelly slopes of knolls are shorter, with smaller leaves,
while the plants on sands in the river valleys are more robust, with
larger, usually densely pubescent leaves. It is also possible that we are
dealing with several, sometimes strictly local species.

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Following tradition, I accept for the present both species until a
detailed study of all the forms can be made.

13. C. talassica (Kult.) Juz. comb. n. in sched.—C. affinis var.
talassica Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIIIc, VI (1929) 13.

Perennial. Root woody, blackish-fibrous, with thick, usually white-
tomentose collar. Stem erect, solitary, finely arachnoid-hairy or
subglabrous, strongly divericately branched in upper half, with one-
headed branches forming corymbose inflorescence. Leaves coriaceous,
green, glabrous or finely arachnoid-hairy above, white-tomentose beneath;
basal leaves in conspicuous rosettes of many leaves, very short-petiolate,
with thickened petioles, crenate-prickly-toothed, oblong-ovate or ovate;
cauline leaves shorter upward, sessile, semiamplexicaul, with roundish
auricles, orbicular or broad-ovate, crenate-prickly-toothed fewer. Capitula
globose, 11-16 mm wide (excluding cusps), densely arachnoid-hairy,
somewhat foveate at base. Involucral bracts numerous, appressed at base,
narrowed in subulate, strong, usually aracuate cusp; inner bracts
membranous, lanceolate, somewhat flat, with short and thin cusp,
scabrous along margin and dorsally. Recetpacular bristles smooth, only
some with one to three crenations. Corolla yellowish. Achenes obovoid,
4—5 mm long, three or four-angled, with sharp, winged thickened angles,
with three or four teeth above, light gray, with dark transverse bands.
Flowering VI—VII; fruiting VIII.

Loamy and conglomerate slopes in the foothills—Soviet Central
Asia: Tien Shan (west). Endemic. Described from the Talass River Basin,
Ur-Maral River. Type in Tashkent.

14. C. pseudoaffinis Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIIIc,
VI (1929) 10.—Ie.: Kult. Ibid., Plate 11.

Perennial. Root woody, with thick collar covered by remains of old
leaves. Stem 25-80 cm high, erect, almost glabrous, whitish, glossy,
almost smooth, profusely branched above. Leaves glabrous, green above,
white-tomentose beneath, boldly crenate-prickly-toothed or crenate-lobed;
basal leaves rosulate, usually falling by anthesis, short-petiolate, oblong-
lanceolate, crenate-lobed, lobes spinescent; lower cauline leaves sessile,
like basal; middle and upper cauline leaves ovate, sessile, auriculate,
prickly-toothed, gradually reduced upward, sparse; uppermost leaves
strongly reduced, Capitula ovoid, 12 mm wide (excluding cusps), finely
arachnoid-hairy. Involucral bracts few, remote, imbricate, somewhat flat,
appressed at base, squarrose; outer bracts one-third to half as long as
middle; middle 12-15 mm long, oblong-ovate, 3 mm wide at base,
abruptly narrowed in upper one-third into thin, deltoid, divergent spine,
adjacent bracts not exceeding middle; inner bracts membranous,

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lanceolate, with 17-20 mm long terminal spine. Receptacular bristles
scabrous. Corolla white; anther tube purple. Achenes obovoid, 5 mm
long, tetraquetrous, with wing-shaped thick ‘angles, three or four-angled,
above. Flowering VI-VII; fruiting VII—VIII.

Outcrops of varicolored rocks in the foothills —Soviet Central Asia:
Syr-Darya. Endemic. Described from Kaplanbek. Type in Tashkent.

Note. The type is not designated in the description of the species; I
propose to consider the plants with the label “Kaplanbek. Popov. 1920
VII 19, Nos. 780, 787” as the lectotype.

15. C. hamadae Juz. in Tr. Tadzh. Bazy. Akad. Nauk SSSR, 8
(1940) 523, 550.—C. pseudaffinis var. bucharica Kult. in Sched. Herb.
Fl. As. Med. 20 (1928) No. 485; Tr. Sredneaz. Gos. Univ. Ser. VIIIc,
VI, 11.—Exs.: H.F.A.M. No. 485.

Perennial. Roots woody, many-headed. Stems fewer, erect, 25-50
cm high, finely arachnoid-hairy, glabrescent and then glossy, light
colored, smooth, weakly branched above or from base, sparsely
arachnoid-hairy, green above, white-tomentose or finely gray-tomentose
beneath, boldly spinose-toothed or lobed with spinescent lobes or teeth,
sessile; basal leaves forming dense rosette, oblong-ovate or cuneately
narrowed toward base, oblong-oblanceolate; lower cauline leaves like
basal; middle and upper leaves fewer, sparse, ovate or oblong-ovate,
auriculate, almost half as long as basal; uppermost leaves reduced.
Capitula oblong-ovoid or flat-globose, 10-14 mm wide (excluding cusps),
arachnoid-hairy, subglabrous. Involucral bracts less, numerous, sparse,
somewhat flat, appressed near base, broad-ovate, abruptly narrowed
upward in three-angled divaricate, strong spine; outer bracts decurved,
almost half as long as middle; middle bracts 10-12 mm long; inner
membranous, lanceolate, spinescent, scabrous along margin and dorsally,
12—20 mm long and 2.5 mm wide. Receptacular bristles weakly scabrous
above. Corolla creamish or whitish; anther tube light purple. Achenes
obovoid, almost tetraquetrous, with sharp, wing-like thickened angles,
with four teeth above, 44.5 mm long. Flowering V—VI; fruiting VII-—
VI.

-Submountainous stony deserts, on the outcrops of varicolored rocks
in foothills and on bare mountains.—Soviet Central Asia: Kyzyl-Kum,
Pamiro-Alai. Endemic. Described from region between Kyzyl-Tepe and
Malik. Type in Tashkent. |

Note. This species has a quite wide range and its characters vary
somewhat. At present, I do not have sufficient material to determine the
range of character variation. I have tried to separate the plants from the
northwestern part of the range (Kuldzhuktau Region, Tamdytau) as a
separate species, naming them C. victoris Tschern.; these plants differ

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from C. hamadae proper by larger, almost glabrous capitula and less
pubescent leaves and stem. However, we consider it essential to verify
carefully the constancy of these characters in populations, before
resolving the question of segregating the species.

The northeastern locality, disjunct from the main range, also requires
a special study: the Turkestan Range, ascending from Aksu to Zambar.
Only very young plants with still undeveloped capitula were collected
from this region.

In establishing C. hamadae, S.V. Juzepczuk qualifies it as a nom.
nov., and includes in the synonyms pseudaffinus var. bucharica Kult.,
which was described by M.V. Kultiasow on the basis of specimens from
the area between Kyzyl-Tepe and Malik, issued in H.F.A.M. (No. 485).
These specimens, and not the specimens from Kugitang, as assumed by
S.V. Juzepczuk, should be considered as the type of the species.

16. C. xanthiocephala Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 107.

Perennial. Root vertical, woody, with thick somewhat pubescent collar,
usually covered with remains of old petioles. Stem erect, whitish, glabrous,
glossy, with short branches in upper half; branches one-headed, sometimes
very short, and capitula almost sessile, in racemose inflorescence. Leaves
coriaceous, pale yellow above, glabrous, with network of prominent veins,
densely white-tomentose beneath; basal leaves oblong-obovoid, petiolate,
crenate-prickly-toothed; cauline leaves gradually reduced, sessile,
semiamplexicaul; lower cauline leaves like basal; middle and upper cauline
leaves orbicular-ovate, or orbicular more or less obtuse, spinose-dentate.
Capitula cylindrical, 5—6.5 mm wide (excluding cusps) and 18 mm long
(excluding corollas), almost glabrous or very finely appressed arachnoid-
hairy; all involucral bracts closely appressed, ovate to oblong-ovate, free
in upper half or one-fourth, narrowed in strong, almost horizontal, 3-4 mm
long, yellowish cusp; inner bracts flat, lanceolate, acuminate, dorsally
scabrous, erect. Receptacular bristles white, somewhat broadened
above, scabrous. Corolla whitish; anther tube creamish. Young achenes
obpyramidal, tetraquetrous, toothed above, 4—4.5 mm long. Flowering
VII.

Outcrops of red clays——Soviet Central Asia: Syr-Darya. Endemic.
Described from the Alymtau Mountains, Sarypaya-Sai Ravine. Type in
Leningrad.

17. C. cryptadena Juz. in Bot. Mat. Gerb. Glavn. Bot. Sada, V
(924) 111; Kult. in Fl. Turkm. VII, 201.—C. leucocitrina Kult. in Bot.
Mat. Gerb. Glavn. Bot. Sada, V (1924) 143.

158

sea]

Perennial. Root woody, many-headed. Stem erect, with remains of
old petioles near base, sparsely arachnoid-hairy below, usually
subglabrous above, paniculately branched, branches divergent, bearing
numerous capitula, aggregated in lax panicles. Leaves coriaceous, green,
almost glabrous above, white-tomentose beneath, with prominent veins;
basal leaves ‘petiolate, pinnatipartite, with broadly deltoid, spinescent,
15 cm long and 5 cm wide lateral lobes; lower cauline leaves like
basal, but with shorter petioles, 12 cm long, 5 cm wide; middle and
upper cauline leaves sessile, broad-lanceolate or almost ovate, broadly
short-decurrent, with roundish auricles, crenate-prickly-toothed, gradually
reduced upward, strongly reduced in inflorescence. Capitula on short
peduncles, numerous, oblong-cylindrical, 15 mm long and 4 mm wide
(excluding cusps), finely arachnoid-hairy, weakly divergent at base, with
fewer florets. Involucral bracts about 30; outer bracts 6 mm long, 2 mm
wide, oblong-ovate, short-acuminate; middle bracts oblong, appressed
at base, gradually acuminate in carinate weakly recurved cusp, 12-15
mm long, 2—3 mm wide near base; inner bracts linear-lanceolate,
membranous, acuminate in thin and short cusp, scabrous along margin.
Corolla yellow; anther tube purple; receptacular bristles white, almost
smooth or weakly scabrous. Achenes oblong-cylindrical, 5 mm long, 2
mm wide, longitudinally nbbed, narrow-winged along ribs, with sharp
teeth above. Flowering V; fruiting VII.

Soviet Central Asia: mountainous Turkmenia (Kopetdag). Endemic.
Described from Chuli District. Type in Leningrad.

18. C. antonowii Winkl. in Tr. Peterb. Bot. Sada, XI (1889) 135;
XI, 227, XIV, 200; Kult. in Fl. Turkm. VII, 212.—Arctium antonowi
O. Ktze. Rev. Gen. 1 (1891) 307.—Ie.: Winkl. op. cit. (1889), Plate 2,
Fig. 2——Exs.: H.F.A.M. No. 41.

Cultivated. Semishrub with thick woody root. Stem erect, 50-70
cm high, slender, glabrous, glossy, sometimes very finely arachnoid-
hairy, usually divaricately branched above, with slender one-headed
branches. Leaves coriaceous, glabrous, green above, densely white-
tomentose beneath; lower leaves usually petiolate, 15-18 cm long
including petiole (petiole 4 cm long), oblong, pinnatilobate, lobes obtuse,
with short apical spine, sparsely spinose-toothed; other cauline leaves
sessile, semiamplexicaul, auriculate, gradually reduced upward, oblong
or ovate-cordate, undivided, with short apical spine indistinctly crenate-
prickly-toothed. Capitula ovoid, 8 mm wide and 17-20 mm long
(including florets), finely arachnoid-hairy. Involucral bracts about 80,
broad-lanceolate, closely appressed, acuminate into short arcuately
recurved cusp; inner bracts thin coriaceous, lanceolate, short acuminate,

160

158

scabrous along margin, 11-12 mm long, erect. Receptacular ,bristles
smooth. Corolla whitish; anther tube reddish. Achenes obovoid, 4.5—5
mm long, with three or four ribs, very narrowly and discontinuously
winged along ribs, blunt-toothed above. Flowering VI—VII; fruiting VII—
VIII. _

Loamy debris slopes in foothills and lower mountain zone.—Soviet
Central Asia: mountainous Turkmenia. Endemic. Described from. Geok-
Tepe. Type in Leningrad.

19. C. glandulosa Kult. in Tr. Sredneaz Gos. Univ. Ser. VIlIIc, VI
(1929) 13; Kult. in Fl. Turkm. VII, 192.—Ic.: Kult. op. cit. (1929),
Plate 12.

Perennial. Root vertical, woody, many-headed, with thick collar.
Stems numerous, somewhat thickened at base, white-tomentose, with
fewer remains of old leaves, white, glossy, smooth, sparsely branched
in upper half. Leaves green, glandular above, grayish-green, finely
arachnoid-hairy beneath; basal and lower cauline leaves petiolate, oblong,
pinnatisect into almost undivided lateral segments, gradually reduced
downward, lanceolate, more or less obtuse, with very short thin cusp,
segments remote on narrow-winged stalk; other cauline leaves less
numerous, gradually reduced, sessile, semiamplexicaul; uppermost leaves
strongly reduced. Capitula 15 mm long (excluding florets), 9-10 mm
wide (excluding cusps); globose-ovoid, strongly foveate at base.
Involucral bracts deltoidly brpadened at base, appressed, attenuate above
into weakly falcate cusp, sparsely glandular dorsally; inner bracts
membranous, short acuminate, scabrous along margin and dorsally,
arachnoid-hairy. Receptacular bristles smooth, somewhat flattened.
Corolla whitish, anther tube purple. Achenes obpyramidal, 5 mm long,
three or four-angled, toothed above. Flowering VI; fruiting VII.

Clayey and stony slopes.—Soviet Central Asia: mountainous
Turkmenia (Kopetdag). Endemic. Described from Kyurendag, Aeishem
Mountains. Type in Tashkent.

Section 5. Chrysoptera Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 78.—Recurvatae Bge. in Mém. Acad.
Pétersb. 7 sér. 9, 2 (1865) 19 p. p.—Squarrosae Boiss. Fl. or. III (1875)
460 p. p—Drepanophorae Winkl. in Tr. Peterb. Bot. Sada, XII (1892)
206 p. p.; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8 (1940) 550 p.
p.—Heteracanthae Winkl. op. cit. (1892) 225 p. p.—Capitula solitary
terminal on branches or numerous, more or less aggregated; involucral
bracts acuminate into long or short, thin, falcate, less often erect cusp;
corolla yellow, often with purple anther tubes; receptacular bristles
smooth; achenes obovoid, three or four-angled, toothed above. Biennial

159

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161

160

herbs with winged stem and sessile cauline leaves having excurrent
wings.
Type of section: C. alata Schrenk.

Series. 1. Dissectae Juz.—Capitula comparatively large, solitary,
terminal on branches; stem narrow-winged, prickly-toothed or almost
smooth.

20. C. dissecta Kar. and Kir. in Bull. Soc. Nat. Mosc. XV (1842)
391; Ldb. Fl. Ross. II, 679; Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2,
23; Boiss. Fl. or. III, 477; Winkl. in Tr. Peterb. Bot. Sada, XII, 209;
Juz. in Bot. Mat. Gerb. Glavn. Bot. Sada, II, 54; Pavlov, Fl. Tsentr.
Kazakhst. III, 296.—C. dissecta var. genuina Trautv. in Tr. Peterb. Bot.
Sada, IX (1886) 456.—Arctium dissectum O. Ktze. Rev. Gen. 1 (1891)
307.

Biennial. Stem erect, 25—35 cm high, white arachnoid-hairy, narrow-
winged throughout, corymbosely branched above, branches spreading
upward, one-headed, narrow-winged; wings of stem 2-4 mm wide,
grayish-green, densely arachnoid-hairy, crenate-prickly-toothed. Leaves
densely arachnoid-hairy above, white-tomentose beneath; basal and lower
cauline leaves short-petiolate, pinnatisect in unequal, narrow-lanceolate
segments, in turn pinnatipartite in spinescent lobes on winged leaf stalk;
cauline leaves like lower, but sessile, decurrent into wings; uppermost
leaves narrow-lanceolate, spinose-toothed, short, reaching almost to
capitula. Capitula globose-ovoid, 12-15 mm wide (excluding cusps),
arachnoid-hairy. Involucral bracts numerous, all, except inner, appressed
at base, falcate in middle, carinate dorsally, subulate-acuminate with
thin, usually yellow, 3 mm long cusp; inner bracts erect, narrow-
lanceolate, scabrous along margin, gradually acuminate upward into short
and thin cusp. Receptacular bristles smooth. Corolla pale-yellow. Achenes
almost tetraquetrous, narrow-winged along ribs, with four teeth above,
5 mm long and 1.5—2 mm wide, dark gray. Flowering V—VI; fruiting
VI-VIII. (Plate IX, Fig. 1).

Along borders of hummocky sands, in sandy steppes, sandy and
pebbled valleys of steppe rivers—Soviet Central Asia: Balkhash Region.
General distribution: Dzhungaria-Kashgaria. Described from the Arganaty
Mountains. Type in Leningrad.

21. C. perovskiensis (Bornm.) Juz. comb. n. in sched.—C. dissecta
var. perovskiensis Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916)
[53:

Biennial. Plants morphologically much resembling C. dissecta Kar.
and Kir., but differing by more dense, loose, white tomentum on stem

162

161

and leaves, 11-12 mm wide, oblong-ovoid capitula (excluding cusps),
and numerous involucral bracts with long, subulate, usually yellowish
6-7 mm long, deflexed cusp, otherwise like C. dissecta Kar. and Kir.
Flowering VI-VII; fruiting VIII. (Plate IX, Fig. 2).

On saline plains, wormwood steppes, sometimes at the foot of sandy
hillocks.—Soviet Central Asia: Balkhash, Aralo-Caspian, Syr-Darya,
Kyzyl-Kum. Endemic. Described from Kyzyl-Dzhar. Type in Leningrad.

22. C. sogdiana Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916)
154; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 551.—C. dissecta
var. macrocephala Rgl. and Schm. in Izv. Obshch. Lyubit Estestv.
Antrop. Etnogr. 34, 2 (1882) 49—C. subinermiceps Juz. in Bot. Mat.
Gerb. Glavn. Bot. Sada, II (1921) 55.

Biennial. Stem 30—70 cm high, erect, floccose-tomentose, narrow-
winged, sparsely branched from middle, branches long, one-headed, wings
2-3 mm wide, crenate-prickly-toothed, arachnoid-hairy. Leaves gray-
tomentose beneath, finely arachnoid-hairy, grayish-green above; basal
leaves petiolate, linear-lanceolate, pinnatipartite, often remote, prickly-
toothed, terminal part long, lanceolate, acuminate, finely spinose-toothed,

_ lower cauline leaves sessile, decurrent to stem wings, like basal leaves;

middle cauline leaves lanceolate, sessile, decurrent; uppermost leaves
reduced to spines. Capitula 15 mm wide (excluding cusps), globose-
cylindrical, densely pubescent. Involucral bracts imbricate, densely
pubescent with simple hairs, appressed at base, attenuate in upper one-
third into falcate, 2-3 mm long, stiff, thin spine; outer bracts half as
long as middle; middle bracts 12 mm long; next row flat, long-lanceolate,
15 mm long, with long and thin terminal spine; inner bracts linear, with
thin, erect spine. Receptacular bristles smooth. Corolla yellowish.
Achenes ovoid, tetraquetrous, angles narrow scariously-winged, forming
fringe of small teeth at apex. Flowering V—VII.

On stabilized and hummocky sands in deserts.—Soviet Central Asia:
Kyzyl-Kum, Pamiro-Alai (Zeravshan River Valley). Endemic. Described
from the Dzhizmanskii Ravine. Type in Leningrad; cotype in Tashkent.

23. C. dolichoclada Juz. in Bot. Mat. Gerb. Glavn. Bot. Sada, II
(1921): 57.

Biennial. Stem 30-80 cm high, erect, grayish arachnoid-hairy-
tomentose, with sparsely spinose-toothed narrow wings, branched in upper
half; branches one-headed, wingless below capitulum. Leaves grayish-
green, finely arachnoid-hairy with white-tomentose veins above, gray-
tomentose beneath; basal leaves petiolate, deeply pinnatipartite, almost
pinnatisect below, with linear, crenate-prickly-toothed lateral segments;
cauline leaves sessile, decurrent on wings, like basal leaves; uppermost

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162

leaves strongly reduced. Capitula 12-14 mm wide, almost ovoid,
tomentose. Involucral bracts numerous, densely woolly dorsally, closely
appressed, narrowed in upper half into thin, falcate, 44.5 mm long
cusp; innermost bracts lanceolate, terminating in thin, slightly curved
cusp. Receptacular bristles smooth. Corolla pale yellow. Achenes
obovoid, tetraquetrous, narrow-winged along angles, with four teeth
above, whitish, about 5 mm long. Flowering V—VI.

On sands.—Soviet Central Asia: Kyzyl-Kum. Endemic. Described
from the Amu-Darya Basin, between Tyuya-Muyun and Mesheklin-Kala
fortress. Type in Leningrad.

Note. The species was described without designating the type. As
the lectotype, I recommend the plants with the label “Near Amu-Darya,
between Tyuya-Muyun.and Mesheklin-Kala fortress. Sands. 19 V 1915.
Krascheninnikov, No. 179a.”

Series 2. Alatae Juz.—Capitula in wide, relatively compact
inflorescences; stem wings narrow or often in many rows more or less
wide, usually deeply incised.

24. C. alata Schrenk. Enum. pl. nov. I (1841) 40; Ldb. FI. Ross. I,
678; Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2, 22; Boiss. Fl. or. IL, 178;
Winkl. in Tr. Peterb. Bot. Sada, XII, 209; XIV, 220; Pavlov, Fl. Tsentr.’
Kazakhst. III, 296; Kult. in Fl. Turkm. VII, 196.

Biennial. Stem erect, 30-80 cm high, smooth, finely arachnoid-hairy
or almost glabrous, with wings in many rows throughout, wings crenate-
prickly-toothed 4—5 mm wide, branched in upper half; common
inflorescence paniculate, sometimes hemicorymbose. Leaves gray-
arachnoid on both sides; basal leaves petiolate, bipinnatisect, with lateral
segments narrow-lanceolate, almost entire; lower cauline leaves like basal,
but sessile, decurrent; middle and upper leaves pinnatipartite into narrow-
lanceolate lateral lobes, sessile, decurrent to stem wings.- Capitula on
peduncles, numerous, 16 mm long and 9-10 mm wide (excluding cusps),
ovoid, weakly arachnoid-hairy. Involucral bracts numerous (about 70),
all except inner lanceolate, 5—10 mm long, gradually acuminate in thin,
about 5 mm long, deflexed cusp; inner bracts narrow-lanceolate, 16 mm
long, erect, membranous above, gradually narrowed in thin, erect cusp.
Corolla yellowish; anther tube pink. Receptacular bristles smooth.
Achenes obovoid, almost triquetrous, weakly curved, with three teeth
above, 3.5 mm long and 1.5 mm wide. Flowering VII—VIII.

Semi-stabilized, small-hummocky sands.—Soviet Central Asia:
Balkhash Region, Aralo-Caspian Region, Syr-Darya, Kyzyl-Kum, Kara-
Kum. General distribution: Northern Iran. Described from the Arganaty
Mountains. Type in Leningrad.

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163

25. C. oxiana Tschern. in Bot. Mag. Gerb. Inst. Bot. Akad. Nauk
UzbSSR, XVII (1962) 100.—C. kuschkensis Kult. in Fl. Turkm. VII
(1960) 197, nomen.—C. bipinnata auct. Fl. As. Med. p. p—C. alata
auct. non Schrenk; Bornm. in Oest. Bot. Zeitschr. (1912) 9.—TIe.: FI.
Turkm. VII, Plate 30, 2.

Biennial. Stem erect, 20-40 cm high, floccose-tomentose, winged
throughout, with pinnatisect, spinescent, arachnoid-hairy wings, branched
in upper half; branches divaricate, branched furhter into numerous, short,
usually one-headed secondary branches. Leaves grayish-green, arachnoid-
hairy above, gray-tomentose beneath; basal leaves petiolate, pinnatisect,
lateral segments acuminate, entire or with one pair of small lobes at
base, less often pinnatilobate; lower cauline leaves like basal, but very
short-petiolate; middle and upper leaves sessile, small, decurrent to stem
wings, pinnatilobate, lobes narrow-deltoid, spinescent. Capitula numerous,
4-5 mm wide, oblong-globose, arachnoid-hairy, roundish at base.
Involucral bracts numerous, imbricate, terminating in weakly curved 1—
1.5 mm long cusp, ovate to lanceolate. Receptacular bristles smooth.
Corolla yellow; anther tube pink. Achenes obovoid, three or four-angled,
toothed above, brown, 3 mm long. Flowering V—VI; fruiting VI-VII.

On stabilized hummocky sands and loamy slopes in foothills
and deserts.—Soviet Central Asia: mountainous Turkmenia (central
Kopetdag, Badkhyz, Amu-Darya, Kyzyl-Kum, Pamiro-Alai). Endemic.
Described from the vicinity of Kushka. Type in Tashkent.

Note. It is a very widely distributed species, smiliar to C. bipinnata.
The characters vary somewhat. Apparently, C. oxiana Tschern. and C.
turkmenorum Bornm. also grow in central Kopetdag; forms intermediate
between these species are found here. It is essential to study large
collections from this region to determine the precise range boundaries
of these two species.

26. C. psammophila Kult. in Tr. Sredneaz. Gos. Univ., Ser. VIII,
VI (1929) 15; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 551.

Biennial. Stem 40-70 cm high, slender, erect, glabrous, glossy,
discontinuously narrow-winged, paniculately-branched from middle,
branches slender, short, one-headed; wings of stem narrow, 2—3(5) mm
wide, crenate-prickly-toothed, glabrous, finely arachnoid-hairy only at
base. Basal leaves sparsely gray-tomentose on both sides (more densely
beneath), petiolate, ovate-lanceolate, deeply bipinnatipartite, with linear-
lanceolate, entire, terminal lobe and deeply pinnatipartite lateral lobes,
weakly spinose-toothed; leaf axes usually narrow-winged, weakly spinose-
toothed; lower cauline leaves like basal, but sessile, decurrent to stem
wings; middle cauline leaves deeply bipinnatipartite, glabrous, sessile,
decurrent to narrow spinose-toothed wings; upper cauline leaves greatly

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164

reduced, sessile, long-decurrent, glabrous; uppermost leaves reduced in
subulate scales. Capitula 5-7 mm wide with approximately 20-florets,
oblong-cylindrical, globose or weakly truncate at base, more or less
arachnoid-hairy. Involucral bracts numerous (about 50), appressed,
imbricate, linear-lanceolate, acuminate in thin, short, 2—3 mm long erect
spine; inner bracts coriaceous, green, long, lanceolate, 1—1.2 cm long,
acuminate above in thin spine. Receptacular bristles smooth. Corolla
yellow; anther tube purple. Achenes obovoid, light brown, tetraquetrous,
very narrow-winged along margin. Flowering V—VI; fruiting VII-—VIII.
(Plate X, Fig. 3).

Sandy loam hillocks and desert sands.—Soviet Central Asia: Kyzyl-
Kum, Amu-Darya, southern Pamiro-Alai. Endemic. Described from the
Saman Mountain. Type in Tashkent.

27. C. schistoptera Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1,
3 (1936) 301; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 551.—C.
bipinnata auct. Fl. As. Med. p.p.

Biennial. Stem erect, 20-65 cm high, weakly arachnoid-hairy
tomentose or glabrous above, winged throughout; wings in many rows,
narrow or rather wide (3-15 mm), deeply (almost up to stem) incised
into frequent, oblong or narrow-deltoid segments, abruptly acuminate
above in stiff and rather long spine, larger segments crenate, prickly
toothed; stem branched from base of often from middle with long,
winged, three to five headed branches, peduncles often wingless. Basal
leaves few (one to three) by anthesis, 10-25 cm long, coriaceous, oblong-
lanceolate, pinnatipartite, usually with narrow-winged petiole, lateral
lobes large-pinnatilobate, spinescent. Leaves green and glabrous above,
grayish-green, finely tomentose beneath; cauline leaves decurrent to
wings, gradually reduced, usually like basal, but uppermost leaves almost
undivided, greatly reduced. Capitula numerous, 20—150, oblong-ovoid,
13-20 mm long, 5-8 mm wide (excluding cusps), finely arachnoid-hairy
with nearly 20 florets. Involucral bracts bright green; outer bracts rather
short, narrow-lanceolate, appressed at base, attenuate above into divergent
spine, almost as long as appressed part; middle bracts oblong, lanceolate,
attenuate above into erect squarrose cusp; inner bracts thin-coriaceous,
acuminate, scabrous along margin. Receptacular bristles smooth. Corolla
yellow; anther tube pink. Achenes obpyramidal, three to five ribbed,
sharp toothed above, light-gray, about 4 mm long. Flowering V—VI;
fruiting VI-VII.

Sands and in the foothills——Soviet Central Asia: Kyzyl-Kum, Kara-
Kum, mountainous Turkmenia (foothills of Kopetdag and Badkhyz).
General distribution: Afghanistan. Described from Repetek. Type in
Leningrad.

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165

28. C. turkmenorum Bornm. in Beih. bot. Centralbl. XXXIV, II
(1916) 153.—C. dissecta var. microcephala Tautv. in Tr. Peterb. Bot.
Sada, IX, 2 (1886) 456; Winkl. Ibid., XI, 135; XII, 209.—C. dissecta
var. simplicior Trautv. op. cit. IX, 2 (1886) 456; Winkl. Ibid., XI, 135;
XII 209.—C. stenoptera Juz. in Bot. Mat. Gerb. Glavn. Bot. Sada, II
(1921) 59; Kult. in Fl. Turkm. VII, 198.—C. simplicior Juz. op. cit. IV

(1923) 77; Kult. op. cit. 196.

Biennial. Root vertical, 8-12 mm thick at root collar. Stem erect,
25-32 cm high, arachnoid hairy-tomentose, narrow-winged throughout,
wings in many rows, 1—2 mm wide, crenate-prickly-toothed, stem usually
corymbosely branched in upper half, with long, very narrowly winged
branches, almost wingless below capitula, forming broad corymbose
inflorescence. Leaves grayish-green, finely arachnoid-hairy above; basal
leaves few (two to four), petiolate, 12-20 cm long, comparatively soft,
oblong-lanceolate, pinnatisect, with narrow-winged petiole, lateral lobes,
in turn, pinnatilobate or pinnatipartite, abruptly short-spinescent-
acuminate; cauline leaves decurrent to wings, gradually reduced upward;
lower cauline leaves like basal; middle leaves less incised; upper leaves
almost undivided, linear, sparsely-toothed, spinescent. Capitula numerous,
20-40, ovoid, 10-18 mm long and 7-10 mm wide, arachnoid, with
approximately 30-florets, truncate at base. Involucral bracts sparsely
glandular, broad-lanceolate, broadened at base, appressed, narrowed above
into short, falcate cusp; inner involucral bracts thin-coriaceous, lanceolate,
erect, acuminate above in short recurved cusp, scabrous along margin
and dorsally. Receptacular bristles smooth. Corolla yellow; anther tube
purple. Achenes obovoid, 3 mm long and 1.5 mm wide, almost three-
angled, irregularly toothed above, light gray. Flowering V—VI; fruiting
VI-VII.

Loamy and clayey slopes in the foothills—Soviet Central Asia:
mountainous Turkmenia (western Kopetdag, Bolshie Balkhany). Endemic.
Described from Kara-Kala. Type in Leningrad.

Section 6. Olgaeanthe Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 78.—Drepanophorae Winkl. in Tr. Peterb.
Bot. Sada, XII (1892) 206 p. p.; Juz. in Tr. Tadzh. Bazy Akad. Nauk
SSSR, 8, 550 p. p.—Capitula globose, terminal on branches; involucral
bracts numerous, gradually narrowed into three-angled falcate cusp;
corolla yellow; receptacular bristles smooth; achenes obovoid, obtuse.

Biennial herbs with soft, sessile, almost entire cauline leaves.

Type of species: C. olgae Rgl. and Schmalh.

29. C. olgae Rgl. and Schm. in Izv. Obshch. Lyubit Estestv. Antrop.
Etnogr. 34. 2 (1882) 48; Winkl. in Tr. Peterb. Bot. Sada, XII, 212;

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166

XIV, 221; Bornm. Beih. bot. Cerntralbl. XXXIV, 2, 155; Juz. in Tr.
Tadzh. Bazy Akad. Nauk SSSR, VIII, 550; Kult. in Fl. Turkm. VII,
210.—C. olgae var. cordata Winkl. op. cit. XII (1892) 212; XIV, 221.—
C. capusii Franch. in Ann. Sc. Nat. Bot. 7 sér. 16 (1883) 313; Winkl.
op. cit. XII, 230—C. aptera auct. non Aitch. and Hemsl.; Winkl. op.
cit. XIV (1897) 224 p. p—Arctium capusi O. Ktze. Rev. Gen. 1 (1891)
307.—A. olgae O. Ktze. Ibid., 308.

Biennial. Stem 10—70 cm high, erect, almost smooth, more or less
branched from base. Leaves very finely arachnoid-hairy above, gray-
tomentose beneath, almost undivided spinescent with short, thin spine;
basal leaves short-petiolate, oblong-linear-lanceolate, pinnatisect, lateral
segments sessile, alternate, linear-lanceolate, with two teeth at base; lower
cauline leaves like basal, but sessile; middle and upper leaves. sessile,
short, closely placed, oblong, with one or two pairs of teeth at base,
short acuminate; uppermost leaves reduced. Capitula globose, 13-15 mm
wide (excluding cusps), weakly arachnoid-hairy. Involucral bracts
numerous, squarrose, imbricate, almost all equal, carinate, acuminate
into thin, three-angled falcate spine; outermost bracts somewhat decurved,
7-10 mm long; outer and middle bracts 15-17 mm long; inner bracts
membranous, narrow linear, up to 20 mm long, acuminate into thin
cusp. Receptacular bristles smooth. Corolla yellow. Achenes obovoid,
obtuse, grayish, with dark spots. Flowering V—VIII; fruiting VII-VIII.
(Plate IX, Fig. 3).

Loamy-stony slopes in the foothills—Soviet Central Asia: Pamiro-
Alai (west), mountainous Turkmenia (Badkhyz). General distribution:
Afghanistan. Described from the Zeravshan River Valley, Oalyk Ravine.
Type in Tashkent.

Section 7. Chrysantha Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk. UzbSSR, XVII (1962) 79.—Jnermes Winkl. in Tr. Peterb. Bot.
Sada, XII (1892) 196 p. p.—Series Pygmaeae Juz. in Tr. Tadzh. Bazy
Akad. Nauk SSSR, 8 (1940) 544.—Capitula solitary on short branches,
with many florates, with four floral leaves; involucral bracts acuminate
in thin, arcuate and scabrous cusp; corolla grayish-yellow; receptacular
bristles scabrous; achenes obpyramidal, almost tetraquetrous, narrow-
winged, finely toothed above. An annual plant with soft leaves, attenuate
at both ends, and spinose-toothed along margin.

Type of section: C. pygmaea Wink.

30. C. pygmaea Winkl. in Tr. Peterb. Bot. Sada, X (1889) 467;
XII, 198; Bornm. in Beih. bot. Centralbl. XXXIV, 2, 145; Juz. in Tr.
Tadzh. Bazy Akad. Nauk SSSR, 8, (1940) 544.—Arctium pygmaeum O.
Ktze. Rev. Gen. 1 (1891) 308.

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167

Annual. Stem erect, 6-8 cm high, almost four-angled, arachnoid-
hairy, leafy, branched in upper half; branches spreading, slender, shorter,
usually leafless (except four floral leaves), one-headed. Leaves sessile,
soft, gray-tomentose on both sides, lanceolate, attenuate at both ends,
spinescent with short and thin spine; spinose-toothed. Capitula 6 mm
wide (excluding cusps), with many florets, ovoid, arachnoid-hairy,
surrounded by four floral leaves; floral leaves like others, but somewhat
smaller. Involucral bracts numerous, narrow, linear-subulate, attenuate
into thin arcuate, scabrous spine; inner bracts erect, linear, acuminate
into thin scabrous cusp. Receptacular bristles unequal, longest bristles
scabrous. Corolla grayish-yellow. Achenes obpyramidal, almost four-
angled, narrow-winged, 2 mm long, finely toothed above. Flowering
IV-V; fruiting V. (Plate X, Fig. 2).

Varicolored foothills ——Soviet Central Asia: Pamiro-Alai. Endemic.
Described from the Tashbulak Mountains, near the village of Akmechet
along the Kafirnigan River. Type in Leningrad.

Section 8. Actinia Bge. in Mém. Acad. Pétersb. 7 sér 9, 2 (1865)
44: Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 561.—/mbricatae
Boiss. Fl. or. III (1875) 461.—Spicatae Winkl. in Tr. Peterb. Bot. Sada,
XII (1892) 237 p. max. p.—Heteracanthae auct. non Winkl.: Juz. in Tr.
Tadzh. Bazy Akad. Nauk SSSR, 8, 551 p. p—Capitula less numerous,
solitary on long peduncles, or numerous in racemose, discontinuously
racemose or lax paniculate inflorescence. Involucral bracts gradually
acuminate; receptacular bristles scabrous; corolla yellow. Achenes
obpyramidal or obovoid, ribbed, narrow-winged along ribs, sharply
toothed above. Perennial herbs with strongly thickened woody root collar,
often covered with remains of old petioles, leaves sessile at base,
orbicular or cordate, petiolate.

Type of section: C. actinia Boiss.

Series 1. Albertoregeliae Tschern.—Capitula numerous, usually
aggregated in inflorescence; root collars with numerous remains of old
petioles.

31. C. albertoregelia Winkl. in Tr. Peterb. Bot. Sada, X (1887) 91;
XII, 239; XIV, 205; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 561.

Perennial. Roots woody, thick, many-headed; root collars long,
thickened, with remains of old petioles, erect, usually white-tomentose.
Stem 55—150(200) cm high, erect, finely arachnoid-hairy or minutely
glandular, leafy, branched above, branches short, bearing few capitula
each, aggregated in more or less racemose or paniculate inflorescence.
Leaves coriaceous, green, very finely glandular or arachnoid-hairy above

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168

white-tomentose beneath with prominent veins; basal leaves petiolate
with petioles thickened at base, pinnatilobate with deltoid lateral long-
spinescent lobe; cauline leaves like basal but short-petiolate; other leaves
sessile, semiamplexicaul, gradually reduced upward, like basal leaves.
Capitula oblong-cylindrical, 8-10 mm wide and about 20 mm long
(excluding cusps), finely arachnoid-hairy. Involucral bracts numerous,
outer and middle bracts narrow linear, subulately acuminate, outer about
15 mm and middle 30 mm long; inner bracts narrow lanceolate, about
20 mm long, somewhat flat, membranous, mucronate. Corolla yellow.
Receptacular bristles weakly scabrous. Achenes obovoid, ribbed, narrow-
winged along ribs, with two teeth above, about 5. mm long. Flowering
VIUI-VII; fruiting VIII-IX.

Stony-clayey slopes in middle mountain zone.—Soviet Central Asia:
Pamiro-Alai (Vakhsh Valley). Endemic. Described from the vicinity of
the village of Tutkaul. Type in Leningrad.

32. C. bobrovii Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3
(1936) 308; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 561.

Perennial. Root woody, with fibrous-gray bark, many-headed, root
collars globose, 2—3.5 cm wide, with numerous stiff remains of old
petioles, spreading on all sides, densely arachnoid hairy-tomentose. Stem
erect, 50-90 cm high, densely leafy, arachnoid hairy-tomentose near
base, very finely glandular in upper half, almost glabrous, sulcate,
sparsely branched; short branches with one to five densely aggregated
capitula in lax spicate inflorescence. Leaves coriaceous, very finely
glandular on both sides, green, very finely arachnoid-hairy beneath, with
prominent veins; basal leaves petiolate, petioles flattened at base,
semiamplexicaul, arachnoid-hairy and thickened; lamina broad oblong-
lanceolate, 15—20-24 cm long and about 10 cm wide (including lobes);
pinnatilobate, with five or six lateral lobes, lobes broadly deltoid, with
two lateral short spinescent teeth near base; lower cauline leaves like
basal but sessile or short-petiolate, slightly smaller than basal, 8-20 mm
long; middle leaves sessile, auriculate, smaller, otherwise like basal
leaves; upper leaves small sessile, narrow-lanceolate, spinose-toothed,
acuminate in very long and stiff spine. Capitula numerous (15-25), on
very short peduncles, or more or less sessile, broad ovoid, about 15 mm
long, and 8 mm wide (excluding cusps), very finely arachnoid-hairy.
Involucral bracts about 45, herbaceous, very finely glandular; outer bracts
narrow lanceolate, dorsally ribbed, spinulose; middle and subsequent
bracts attenuate from narrow ovate or ovate base into long spine, as
long as capitulum, almost horizontal; inner bracts usually membranous,
linear, short acuminate. Corolla pink (?). Receptacular bristles scabrous.
Achenes compressed-obpyramidal, usually triquetrous, 3.5 mm long and

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169

2 mm wide, dark brown to gray, spotted. Flowering VII—VIII; fruiting
VII.

Outcrops of varicolored rocks in middle mountain zone.—Soviet
Central Asia: Pamiro-Alai (Kugitang). Endemic. Described from the
Kugitang Range, in the vicinity of the village of Kundzhak. Type in
Leningrad.

33. C. oopoda Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3
(1936) 310; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 561.

Perennial. Root woody; blackish-fibrous; root collars very strongly
thickened, almost ovoid’ 3.5—-5 cm wide, with remains of petioles of
previous year’s leaves, white-tomentose: Stem erect, 60-120 cm high,
smooth, finely floccose-tomentose, especially in lower part, sparsely
branched; branches few, short, forming somewhat racemose or narrow
paniculate inflorescence. Leaves green, glabrous above, with prominent
veins, terminating into more or less short and stiff spine, finely gray-
tomentose beneath; basal leaves about 20-25 cm long, short-petiolate,
oblong-lanceolate, crenate-lobed, lobes broadly deltoid, sparsely and
finely spinose-toothed; lower cauline leaves like basal, but with shorter
petioles; middle leaves about 4-10 cm long, sessile, semiamplexicaul,
lanceolate, spinose-toothed, somewhat reduced; upper leaves greatly
reduced, 1.5—3 cm long, as a result upper half of stem very sparsely
covered with small leaves. Capitula small, 7-8 mm wide, almost globose,
rusty-arachnoid-hairy. Involucral bracts less numerous; outer bracts 4—5
mm long, carinate, mucronate; middle bracts linear-lanceolate, carinate,
attenuate into divergent cusp, usually not exceeding capitulum in length;
inner bracts lanceolate, coriaceous, acuminate in thin and short cusp.
Receptacular bristles somewhat broadened above, scabrous. Achenes
obovoid, about 5 mm long, triquetrous, with narrow wings along ridges.
Flowering VII—VIII; fruiting VIII.

Outcrops of varicolored rocks in middle mountain zone.—Soviet
Central Asia: Pamiro-Alai (south-west). Endemic. Described from the
Buryutakht Mountain 30 km south of Baisun. Type in Leningrad.

34. C. gontscharowii Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser.
1, 3 (1936) 311; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 562.—
C. catenata Rech. f. in Dan. Biol. Skr. 8, 2 (1955) 82.—Ie.: Rech. f.
Ibid., fig. 61, 66f.

Perennial. Root woody, strongly fibrous, many-headed, root collar
thick, covered with remains of petioles of previous year’s leaves,
arachnoid-hairy. Stem erect, 40-70 cm high, striate-sulcate, arachnoid-
hairy, white-tomentose below, repeatedly branched in upper half; branches
long, bearing numerous capitula aggregated in broad and lax panicles.

72

170

Leaves glabrous, green above, gray-tomentose beneath, with prominent
veins, terminating into strong spines; basal leaves petiolate, sometimes
petioles with one or two pairs of spines, linear-lanceolate, 18-27 cm
long, irregularly pinnatipartite or lobed, lobes narrowly deltoid or deltoid-
spinescent, with two smaller spinescent lobes at base; lower cauline
leaves like basal but short-petiolate; middle leaves lanceolate, sessile,
semiamplexicaul, spinose-dentate, one-fourth to one-third as long as lower
cauline leaves; upper leaves like middle but much smaller; uppermost
leaves more or less reduced to spines. Capitula 8-9 mm wide, ovoid-
globose, finely arachnoid-hairy. Involucral bracts less numerous; outer
bracts carinate, squarrose not exceeding capitulum; middle bracts also
carinate, strong, squarrose with cusp one and one-half times as long
as capitulum; inner bracts coriaceous, mucronate; innermost bracts
linear, membranous. Corolla yellow. Receptacular bristles scabrous.
Achenes compressed-pyramidal, almost triquetrous, with narrow wings
along angles, about 4 mm long. Flowering VII—VIII; fruiting VIII-—
IX.

Outcrops of varicolored rocks in middle mountain zone.—Soviet
Central Asia: Pamiro-Alai (south). General distribution: Afghanistan.
Described from the Gazimailik Mountains, near the village of Surkhat.
Type in Leningrad.

35. C. stricta Tschern. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
UzbSSR, 16 (1961) 60.

Perennial. Root woody, blackish-fibrous, many-headed; root collars
thick, up to 2.5 cm wide, covered with remains of petioles spreading
above. Stem erect, 40-70 cm high, with sessile glandular hairs, sparsely
branched in upper half; branches numerous, forming paniculate
inflorescence. Leaves coriaceous, green on both sides, with sessile
glandular hairs above, sparsely arachnoid-hairy and conspicuously
glandular-hairy beneath; basal leaves petiolate, with almost glabrous
petioles, oblong-lanceolate, pinnatipartite, lateral lobes approximate,
deltoid, terminating in strong, 2—-2.5 cm long spine, with one or two
spinescent lobes on both sides at base; cauline leaves gradually reduced,
sessile, with auricles. connate with stem, lanceolate, pinnatilobate, lobes
spinescent; upper leaves reduced to spines. Capitula 22 mm long and
7-8 mm wide (excluding cusps), sparsely rusty-arachnoid-hairy, oblong-
cylindrical, narrowed at base. Involucral bracts erect; outer bracts 10
mm long, lanceolate, subulately acuminate; middle bracts flat, broad-
lanceolate, 20-22 mm long, subulately acuminate; not exceeding
capitulum; inner bracts flat, narrow-lanceolate, terminating in short spine,
18 mm long, somewhat scabrous dorsally. Corolla yellow, prominent.
Receptacular bristles scabrous. Achenes oblong-ovoid, about 4.5—5 mm

171

Plate X.
1—Cousinia angusticeps Juz.; 2—C. pygmaea Winkl; 3—C. psammophila Kult.

17]

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172

long, longitudinally ribbed, with narrow-winged nibs, light gray, black-
spotted, finely toothed above. Flowering VIII; fruiting IX. (Plate XI,
Fig. 1).

Limestone and gypsum outcrops in the lower part of middle
mountain zone.—Soviet Central Asia: Pamiro-Alai (Babatag Range).
Endemic. Described from the Babatag Mountains, opposite the Garmaly
Spring. Type in Tashkent.

Series 2. Scalerophyllae Tschern.—Capitula relatively less
numerous, solitary, on long peduncles; root collars thick, with numerous
scars of old leaves, but without remains of petioles.

36. C. scalerophylla Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser.
1, 3 (1936) 308; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 551.

Perennial. Roots woody, with strongly fibrous blackish bark, and
1-2 cm thick collar. Stem solitary, 25-50 cm high, erect, white
arachnoid-hairy, branched in upper one-third with less numerous
branches, in turn branched further. Lower leaves petiolate, oblong,
narrowed toward both ends; cauline leaves sessile, oblong-ovate or
(uppermost) ovate, usually obtuse, usually cordate; all leaves deeply
lobed, with lobes broadly deltoid and spinose-toothed, terminating in
stiff spine, coriaceous, glabrous and lustrous above, densely white-
tomentose beneath, with prominent veins. Capitula solitary, on long
peduncles, about 25 mm long and 12 mm wide (excluding cusps), ovoid,
finely arachnoid-hairy with roundish or somewhat foveate base, usually
with 20-florets. Involucral bracts scabrous dorsally; outer bracts broadly
deltoid or broad-ovate, abruptly terminating into short spine; middle
bracts ovate, gradually attenuate into narrow-lanceolate, three-angled
oblique cusp; inner bracts membranous, oblanceolate, dark brown,
acuminate, more or less incised. Receptacular bristles flat, strongly
scabrous. Corolla straw-yellow; anther tube purple. Achenes obpyramidal,
three-angled, with winged angles, brown, with sharp teeth above, 6—6.5
mm long and 5 mm wide including wings. Flowering VI; fruiting VII.

Outcrops of red sandstones.—Soviet Central Asia: Pamiro-Alai.
Endemic. Described from the region between Naizabulak and Jidabulak.
Type in Leningrad.

Section 9. Pseudactinia Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 79.—Capitula aggregated in panicles or
racemes, solitary or in twos on very short branches. Involucral bracts
gradually acuminate into subulate cusps. Receptacular bristles smooth.
Corolla pink or brownish. Achenes obpyramidal, indistinctly ribbed,
obtuse above. Perennial herbs with thick and woody root collars usually

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173

covered with woody remains of old petioles and sessile semiamplexicaul
or short-decurrent cauline leaves.
Type of section: C. oreodoxa Bornm. and Sint.

37. C. oreodoxa Bornm. and Sint. in Mitt. Thir. Bot. Ver. 23 (1908)
99, nomen in Russk. Bot. Zhurn. 1, 2; Kult. in Fl. Turkm. VII, 205.—
Ic.: Bornm. and Sint. op. cit. tab. 2.

Perennial. Roots woody, blackish-fibrous, many-headed with caudices
covered by numerous woody deflexed petioles of old leaves. Stem erect,
40-70 cm. high, densely arachnoid-hairy, leafy, simple or very sparsely
branched above in very short one-headed branches, forming almost
spicate inflorescence; branches much reduced so that capitula almost
sessile in leaf axils on main stems. Leaves coriaceous, grayish-green,
finely gray-tomentose on both sides; basal and lower cauline leaves
petiolate, oblong, crisped-prickly-toothed, 8-12 cm long, 3—3.5 cm wide;
cauline leaves gradually reduced, sessile, semiamplexicaul, somewhat
attenuate towards both ends, uppermost leaves more or less orbicular,
with roundish auricles, short-decurrent. Capitula cylindrical, 10-14 mm
wide (excluding cusps), arachnoid-hairy. Involucral bracts numerous, all,
except inner, linear, weakly carinate dorsally, narrowed towards apex,
subulately pointed into erect, 5-17 mm long cusp; inner bracts
membranous, lanceolate, 20-25 mm long, long acuminate, scabrous along
margin. Receptacular bristles smooth. Corolla more or less pink; anther
tube pink. Achenes oblong-obpyramidal, 6 mm long, 2 mm wide,
inconspicuously ribbed, obtuse. Flowering VII; fruiting VIII. (Plate XI,
Fig. 2).

Stony slopes in middle mountain zone.—Soviet Central Asia:
mountainous Turkmenia (Kopetdag). Endemic. Described from Sulyukli.
Type in Leningrad.

38. C. apiculata Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad. Nauk
UzbSSR, XVIII (1962) 91.

Perennial. Roots woody, blackish-fibrous, many-headed; caudices
with numerous deflexed woody remains of old petioles. Stem erect, 80—
100 cm high, arachnoid hairy-tomentose, rather densely leafy, racemosely
branched above; branches short, one-headed. Leaves coriaceous, grayish-
green, glandular and arachnoid-hairy pubescent on both sides, some-
what more densely beneath; basal leaves 10-18 cm long and 4-6 cm
wide, oblong-lanceolate, long-petiolate, crenate-prickly-lobed, petioles
with one to three spines; cauline leaves gradually reduced, sessile,
semiamplexicaul, lower and middle leaves like basal; upper leaves 3
cm long, prickly-toothed. Capitula globose, 18-20 mm wide (excluding
cusps), densely arachnoid-hairy. Involucral bracts numerous (more than

175

174

100), all, except inner, linear-broadened at base, gradually attenuate into
divergent, subulately three-angled cusp; outer bracts decurved, 5-7 mm
long; middle bracts 18-20 mm long; inner bracts coriaceous, linear,
somewhat flat, scabrous along margin, weakly broadened above,
mucronate, about 20 mm long. Receptacular bristles smooth. Corolla
pink. Achenes about 5 mm long, obpyramidal, weakly compressed,
indistinctly three-angled, obtuse-roundish above, brownish with dark
spots. Flowering VII; fruiting VIII.

Stony-gravelly slopes in lower mountain zone.—Soviet Central Asia:
mountainous Turkmenia (Kopetdag). Endemic. Described from the
vicinity between the Kheirabad (meteorological) Station and Germab.
Type in Tashkent.

39. C. glochidiata Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIIIc, VI
(1929) 3; Kult. in Fl. Turkm. VU, 213.—Ie.: Fl. Turkm. VII, Plate 33.

Perennial. Roots woody, blackish-fibrous, many-headed; caudices
up to 5 cm wide, covered with remains of old petioles, white-tomentose.
Stem erect, 45-80 cm high, finely striate-sulcate, sparsely arachnoid-
hairy in upper half, floccose-white tomentose below, branched above;
branches short, one or two-headed. Leaves coriaceous, glandular on both
sides (more densely beneath), sparsely and finely arachnoid-hairy,
with prominent veins; lower cauline leaves 12 cm long, 4 cm wide,
short-petiolate, oblong-ovate, crenate-lobed with lobes deltoid-prickly-
acuminate, often with two prickly-acuminate teeth at base; middle leaves
7 cm-long and 4 cm wide, sessile, semiamplexicaul, auriculate, ovate-
oblong; upper leaves 5 cm long, 4 cm wide, semiamplexicaul, roundish-
auriculate at base, spinose-toothed, spinescent; uppermost leaves smaller,
reaching up to capitula. Capitula 4-6 cm wide (including cusps), 2—3
cm (excluding cusps), flat-globose, arachnoid hairy-tomentose. Involucral
bracts numerous, all except inner similar, basally appressed, scabrous
along margin, gradually narrowed into stiff linear-subulate cusp; outer
bracts falcate; middle 26-30 mm long, almost horizontal; inner
coriaceous, weakly scabrous along margin, finely and short acuminate,
slightly broadened, erect, 22-24 mm long. Receptacular bristles smooth.
Corolla brownish (?); anther tube pink. Achenes obovoid, more or less
ribbed. Flowering VII; fruiting VIII.

Turfy and stony slopes.—Soviet Central Asia: mountainous Turkmenia
(Kopetdag). Endemic. Described from the Aselma Range. Type in
Tashkent.

Section 10. Hoplophylla Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 79.—Nudicaules Winkl. in Tr. Peterb.
Bot. Sada, XII (1892) 191 p. p.—Capitula numerous, in paniculate

176

lei

175

inflorescence; involucral bracts somewhat flat, closely appressed, short

acuminate; receptacular bristles smooth. Corolla straw-yellow; anther

tube yellow, glabrous. Achenes obovoid, three-angled, toothed above.

Perennial plants with densely leafy stem and stiff coriaceous leaves,

usually with white spines scattered throughout (mainly along veins).
Type of section: C. raddeana Winkl.

40. C. raddeana Winkl. in Tr. Peterb. Bot. Sada, X (1887) 473 in
adnot.; XI, 132; XII, 196; Kult. in Fl. Turkm. VI, 200.—C. nobilis
Rech. f. in Dan. Biol. Skr. 8, 2 (1955) 132.—Arctium raddeanum O.
Ktze. Rev. Gen. 1 (1891) 308.—Ic.: Winkl. op. cit. (1889), Plate 1;
Rech. f. op. cit. fig. 90 and 94D.

Perennial. Stem solitary, erect, up to 2 m high, whitish, glabrous,
glossy, almost tetraquetrous, paniculately branched in upper half, densely
leafy below, leaves greatly reduced in upper part; branches numerous,
long, with numerous capitula. Leaves stiff-coriaceous, oblong, green
above, glabrous or with isolated multicellular hairs scattered throughout,
with sharply raised, white, glossy midrib, terminating in strong whitish
spine and less prominent, but also whitish and glossy lateral veins, usually
with few whitish spines scattered throughout and terminating into strong
white spine, densely white-tomentose beneath; basal leaves 20-40 cm
long, with short usually prickly petioles, usually prickly-toothed in lower
half, pinnatilobate in upper half with lanceolate finely prickly-toothed
lobes; cauline leaves gradually reduced, sessile, semiamplexicaul, sparsely
prickly-toothed; uppermost leaves strongly reduced, ovate or lanceolate,
prickly acuminate and with two spines at base. Capitula with less
numerous florets (12-15), ovoid, 8-9 mm wide, glabrous. Involucral
bracts green, closely appressed to each other, ovate to oblong-ovate,
somewhat flat, very short acuminate; inner bracts about 10 mm long,
lanceolate, weakly broadened above and short acuminate, scabrous
dorsally and along margin. Receptacular bristles smooth, slightly flattened
above. Corolla straw-yellow; anther tube straw-yellow. Achenes obovoid,
4.5—5 mm long and 3 mm wide, three-angled, indistinctly ribbed, toothed
above, grayish. Flowering VI—-VII; fruiting VII—VIII.

Loamy and sandy slopes.—Soviet Central Asia: Pamiro-Alai
(south, Koipitau Range), mountainous Turkmenia (Badkhyz). General
distribution: Afghanistan. Described from Ak-Rabot. Type in Leningrad.

Section 11. Nanarctium Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR,
8 (1940) 510, in adnot, nomen.; Bot. Mat. Gerb. Inst. Bot. Akad. Nauk
UzbSSR, XVII (1962) 80.—Lappaceae Bge. in Mém. Acad. Pétersb. 7
sér. 9, 2 (1865) 9 p. p. Uncinatae Winkl. in Tr. Peterb. Bot. Sada, XII
(1892) 186 p. p.—Capitula globose, solitary terminal; involucral bracts

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176

numerous, attenuate into long, subulate, uncinate cusp. Corolla whitish

or pale-yellow. Receptacular bristles smooth. Achenes narrow-pyramidal,

with very short apical corona. Perennial plants with pinnatipartite or

pinnatilobate leaves; leaves glandular-hairy above and with glandular

hairs and arachnoid-hairy pubescence of long, flexuous flattened hairs.
Type of section: C. arctioides Schrenk.

41. C. arctioides Schrenk in Bull. phys.-math. Ac. Petrop. 2 (1844)
115; Ldb. Fl. Ross. II, 679; Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2 10;
Winkl. in Tr. Peterb. Bot. Sada, XII, 187; Pavlov. Fl. Tsentr. Kazakh.
II, 295.—Arctium arctioides O. Ktze. Rev. Gen. 1 (1891) 307.

Perennial, plants with one stem. Stem erect, 35-70 cm high, finely
striate-sulcate, densely glandular, arachnoid-hairy pubescent with long,
flexuous flattened hairs, leafy, cymosely branched in upper half; branches
in turn branched into secondary branches, each bearing single capitulum.
Leaves soft, green, sparsely glandular above, densely glandular beneath
wih arachnoid-hairy pubescence of long, flexuous, flattened hairs; basal
leaves short-petiolate, pinnatipartite or pinnatilobate, lobes finely spinose-
toothed, with short and thin apical spine; cauline leaves reduced, sessile,
semiamplexicaul, attenuate towards base, with two or three pairs of
yellowish spines, like basal, leaves on branches reduced, spinose-toothed.
Capitula globose, 10-12 mm wide (excluding cusps). Involucral bracts
numerous, narrow lanceolate, attenuate into long, subulate uncinate cusp,
densely covered with fine glandular divergent hairs along margin; inner
bracts erect. Receptacular bristles smooth. Corolla whitish or pale yellow.
Achenes narrow pyramidal, 5 mm long and 1.5 mm wide, with short
corona above, dark. Flowering VI; fruiting VII. (Plate XII, Fig. 1).

In moist and alkaline meadow depressions in desert steppes, thickets
of shrubs and meadows, valleys of steppe rivers, less often in border
blow outs of sand dunes.—Soviet Central Asia: Aralo-Caspian (between
the city of Kyzyl-Orda and lower reaches of the Sarysu River), Balkhash
Region (Sarysu River Valley and western Betpakdal). Endemic. Described
from the Sarysu River Valley, between the Sarysu and Kara-Kingir rivers.
Type in Leningrad.

Section 12. Pseudarctium Juz. in Tr. Tadzh. Bazy Akad. Nauk
SSSR, 8 (1940) 512, 545—Lappaceae Bge. in Mém. Acad. Pétersb. 7
sér. 9, 2 (1865) 9 p. p.; Boiss. Fl. or. III, 459—Uncinatae Winkl. in
Tr. Peterb. Bot. Sada, XII, 186 p. p—Capitula solitary or aggregated at
branch tips; involucral bracts with uncinately incurved cusp. Corolla
purple or white. Receptacular bristles smooth. Achenes glabrous, smooth,
roundish above. Perennial plants with large, soft, somewhat pubescent
leaves.

Lectotype of section: C. umbrosa Bge.

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177

Series 1. Umbrosae Juz.—Capitula with 10—12-florets; involucral
bracts with one or two glandular hairs along each margin.

42. C. umbresa Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2 (1865) 10;
Boiss. Fl. or. III, 463; Winkl. in Tr. Peterb. Bot. Sada, XII, 188; XIV,
217; Bornm. in Beih. bot. Centralbl. XXXIV, 2, 135; Juz. in Tr. Tadzh.
Bazy Akad. Nauk SSSR, 8, 546.—C. uncinata Rgl. in Bull. Soc. Nat.
Mosc. XL. 2 (1867) 145; Winkl. op. cit. XI, 132.—C. umbrosa 8B.
virescens Bornm. |.c.—Arctium umbrosum O. Ktze. in Rev. Gen. 1 (1891)
308.—A. uncinatum O. Ktze. Ibid—Exs.: H.F.A.M. No. 489.

Perennial. Stem erect, 60-120 cm high, deeply sulcate, glossy,
branched in upper half; branches slender, elongate, with numerous
capitula in paniculate inflorescence. Leaves thin-coriaceous, soft, without
spines, green, glabrous above, gray-tomentose beneath, petiolate mainly
in lower half of stem; basal leaves very large, with thick, sulcate petioles,
oblong-ovate, cordate, irregularly toothed; cauline leaves gradually
reduced towards stem apex, ovate, almost ovate, otherwise like basal
leaves; uppermost leaves greatly reduced. Capitula on very short
peduncles, 5—6 mm wide (excluding cusps), with 10-12 florets, ovoid,
glabrous, weakly foveate at base. Involucral bracts oblong, closely
appressed at base, gradually acuminate into thin, about 3 mm_-long,
spreading uncinately incurved cusp, with one or two pairs of large
glandular hairs at transition of bracts to cusp; inner bracts linear,
gradually acuminate into very thin and short cusp. Receptacular bristles
smooth. Corolla pink. Achenes 6 mm long and 3 mm wide, obovoid,
roundish above, light brown with dark spots. Flowering VI—VII; fruiting
VII-VII. (Plate XII, Fig. 2).

Foothills and lower mountain zone, in shady places, growing as
weed.—Soviet Central Asia: Tien Shan (central and western), Pamiro-
Alai, mountainous Turkmenia. General distribution: Northern Iran,
Afghanistan. Described from the vicinity of village of Dshegar in the
Mashhad Province of northeastern Iran. Type in Leningrad.

Note. The hybrid Cousinia triflora Schrenk < Cousinia umbrosa
Bge., which goes by the name C. taschkentica Juz. in herb., is known
from the vicinity of Tashkent. This plant has large, thin-coriaceous,
exspinose, petiolate basal leaves that are glabrous on both sides; sessile,
coriaceous, finely spinose-toothed cuspidate cauline leaves; and capitula
in a corymbose-paniculate inflorescence. In contrast with C. triflora,
the involucral bracts of C. taschkentica have weakly uncinately incurved
cusps.

The hybrid C. umbrosa Bge. x C. pseudoarctium Bornm., described
as C. dualis Juz. (in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3 (1936)
297), is known from the Alai Range (Lyangar, Chigirchik River).

180

178

The hybrid C. alberti Rgl. and Schm. x C. umbrosa Bge., preserved
in the herbarium as C. triacantha Kult. (in Tr. Sredneaz. Gos. Univ.
Ser. VIlIc, VI (1929) 2), is reported from the Kelte-Moshat pit.

43. C. tomentella Winkl. in Tr. Peterb. Bot. Sada, X (1889) 469;
XII, 188; XIV, 191; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8,
546.—Arctium tomentellum O. Ktze. Rev. Gen. 1 (1891) 308.

Perennial. Species very similar to C. umbrosa Bge., from which it
differs only by densely arachnoid hairy-tomentose capitula. Distributed
only in the southern Pamiro-Alai, where it is sometimes found together
with C. umbrosa. However, I consider that these two species are
distinguishable since, under cultivation in the experimental plots, C.
tomentella maintained its characters and developed much faster than C.
umbrosa.

Central Asia: Pamiro-Alai (south—southern slope of the Gissar
Range, Aktau Mountains, Gazimailik, Terekli, Sarsarak Range, Khoja-
Kazyan). Endemic. Described from the Karatag River Valley, in the
vicinity of Khakimi. Type in Leningrad.

Series 2. Spuriae Juz—Capitula with 20—-45-florets; involucral bracts
without glandular hairs along margin.

44. C. pseudarctium Bornm. in Beih. bot. Centralbl. XXXIV, 2
(1916) 135; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 545.—C.
spuria Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3 (1936) 296;
Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR 8, 545.—C. pseudarctium B.
leiocephala Bornm. op. cit. 136.—C. amplissima auct. non Boiss.:
Franch. in Ann. Sc. Nat. Bot. 6 sér. 16 (1883) 92.

Perennial. Stem erect, 80-100 cm high, longitudinaly sulcate,
floccose-arachnoid hairy, leafy, sparsely branched in upper part; branches
elongate, with numerous capitula in paniculate inflorescence. Leaves
thin-coriaceous, soft, without spines, green, and glabrous above, finely
gray-tomentose beneath, petiolate, basal and lower cauline leaves very
large, lamina about 20-40 cm long and petioles thick, striate-sulcate,
20-40 cm long, ovate, crenate-lobed; other cauline leaves abruptly
reduced upward, ovate or orbicular-ovate, lobed, lobes roundish with
short, thin, apical spine; uppermost leaves greatly reduced. Capitula on
short peduncles or sessile, 10-12 mm wide (excluding cusps), with 20-
45-florets, oblong-cylindrical or almost ovoid, foveate at base. Involucral
bracts numerous, scabrous dorsally, all except inner, narrow lanceolate,
densely appressed at base, gradually attenuate into long, divergent
uncinate cusp; inner bracts lanceolate, somewhat flat, attenuate, short
acuminate. Receptacular bristles smooth. Corolla purple; anther tube pale

179

A
N\A
A \ Y
NN \
\\ |
\ |

\

181 Plate XI.

1—Cousinia stricta Tschern.; 2—C. oreodoxa Bormnm.

182

180

yellow. Achenes large, about 8 mm long and about 3 mm wide, obovoid,
glabrous, smooth, roundish above, light brown. Flowering VI—VII; fruiting
VII.

Turfy slopes.—Soviet Central Asia: Pamiro-Alai (Zeravshan Range,
Alai Range). Endemic. Described from the Yagnob River Basin, found
in the vicinity of the village of Varsout. Type in Leningrad.

Section 13. Lappaceae Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2
(1865) 9 p. p—Uncinatae Winkl. in Tr. Peterb. Bot. Sada, XII (1892)
186 p. p.—Xerarctium Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8
(1940) 510 in adnot. nomen.—Capitula small, numerous, sessile,
aggregated in heads, readily falling. Involcural bracts acuminate in short,
thin and uncinately incurved cusp; corolla purple, receptacular bristles
flat, smooth. Achenes glabrous, smooth, roundish above. Perennial plants,
usually with purple branches and soft, usually pubescent, leaves.

Lectotype of section: C. lappacea Schrenk.

45. C. lappacea Schrenk Enum. pl. nov. (1841) 42; Ldb. Fl. Ross.
II, 679; Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2 10; Winkl. in Tr.
Peterb. Bot. Sada, XII, 187.—Arctium lappaceum O. Ktze. Rev. Gen. 1
(1891) 308.

Perennial. Plants with many stems. Stem erect, strong, 40-70 cm
high and 2 cm thick at base, floccose-tomentose, striate-sulcate,
somewhat angular, usually pinkish or purple (especially branches and
branchlets), spreading, branches short, further branched, secondary
branches bearing less numerous sessile capitula aggregated in heads.
Leaves soft, finely gray arachnoid-hairy above, gray-tomentose beneath;
basal leaves petiolate, oblong, finely spinose-toothed; cauline leaves
gradually reduced, sessile, semiamplexicaul, rather dense, finely prickly-
toothed, attenuate toward apex, oblong-lanceolate; leaves on branches
greatly reduced. Capitula readily falling, ovoid, glabrous, 5-6 mm wide
(excluding cusps) and 12-13 mm long (excluding florets), with four or
five florets. Involucral bracts 25-30, all, except inner, sparsely glandular,
closely appressed, ovate, weakly carinate, with short glandular hairs along
margin, divergent above and acuminate into thin and short uncinately
incurved cusp; inner bracts scarious purple in upper half, obtuse, with
thin and uncinate spine. Receptacular bristles weakly broadened, flat,
smooth. Corolla purple. Achenes obovoid, glabrous, smooth, roundish
above, 6 mm long and 4 mm wide. Flowering VI; fruiting VII.

Stony slopes in middle mountain zone, apparently a semi-weed.—
Soviet Central Asia: Tien Shan (Central Tien Shan, Chu-I]i Mountains,
Aleksandrovskii Range). Endemic. Described from Alatau (Schrenk).
Type in Leningrad.

183

181

Section 14. Ctenarctium Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR,
8 (1940) 512, 546.—Uncinatae Winkl. in Tr. Peterb. Bot. Sada, XII (1892)
186 p. p.—Capitula small, on short peduncles, aggregated at branch
tips, readily falling; involucral bracts spinose-toothed, like apical spine
uncinately incurved; corolla yellow; receptacular bristles smooth; achenes
obovoid, smooth, roundish above. Perennial plants with coriaceous leaves.
Type of section: C. anomala Franch.

46. C. anomala Franch. in Ann. Sc. Nat. Bot. 7 sér. 16 (1883) 93;
Winkl. in Tr. Peterb. Bot. Sada, XII, 189; XIV, 218; Bornm. in Beith.
bot. Centralbl. XXXIV, 2, 138; Juz. in Tr. Tadzh. Bazy Akad. Nauk
SSSR, 8, 546.—Arctium anomalum O. Ktze. Rev. Gen. | (1891) 307.—
Exs.: H.F.A.M. No. 479.

Perennial. Stems few, with remains of petioles of previous year’s
leaves at base, erect, 20-40 cm high, slender, shallow-sulcate, arachnoid-
hairy, sparsely branched from middle; branches long, branched further
in upper part, bearing few capitula on each at tip. Leaves coriaceous,
entire, without spines, very finely arachnoid-hairy above, finely tomentose
beneath; basal leaves petiolate, oblong-lanceolate, attenuate toward both
ends, sometimes lyrate, attenuate into narrow apex; cauline leaves
gradually reduced upward, lower leaves short-petiolate, broad-lanceolate,
entire; middle leaves short-petiolate, ovate; upper leaves lanceolate,
sessile, with one or two pairs of teeth on both sides at base. Capitula on
very short peduncles, readily falling, 15 mm long and 6 mm wide
(excluding cusps), finely arachnoid-hairy. Involucral bracts somewhat
flat; outer bracts oblong-ovate, with membranous border at base, border
disrupted in upper one-third, and two or three pairs of horizontal, uncinate
spines, like apical spine, on both sides right up to apex, middle bracts
like outer, but lanceolate; inner more or less membranous, lanceolate,
gradually acuminate into uncinately incurved spine. Corolla yellow.
Receptacular bristles smooth. Achenes obovoid, smooth, about 2 mm
wide and 4 mm long, roundish above. Flowering VI—VI]J; fruiting VII—
VIII. (Plate XII, Fig. 2).

Clayey and stony slopes in middle mountain zone.—Soviet Central
Asia: Pamiro-Alai (the Zeravshan and Gissar ranges). Endemic. Described
from the vicinity of Voru. Type in Paris.

Section 15. Molles Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2 (1865)
13.—IJnermes Winkl. in Tr. Peterb. Bot. Sada, XII, 196 p. p.—
Malacoloma Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8 (1940) 513,
546.—Stem winged. Capitula small, in groups of few at tips of branches,
on short peduncles or sessile, with few florets; involucral bracts abruptly
attenuate, cusp erect or divergent; corolla yellow with purple anther tube

184

182

or yellowish or purple; receptacular bristles smooth. Achenes obovoid,

smooth, roundish or almost three-angled above, and then toothed in upper

part. Biennial or perennial herbs with soft, sessile cauline leaves.
Type of section: C. mollis Schrenk.

Series 1. Pseudomolles Juz.—Capitula with 6—10-florets, like entire
plant weakly pubescent, greenish; corolla yellow, anther tubes purple.
Perennial plants with almost undivided or pinnatilobate basal and lower
cauline leaves.

47. C. pseudomollis Wink]. in Tr. Peterb. Bot. Sada, X (1889) 470
(exch. spec.” e* Karat. )- Xi "199. XIVe 2195" yuz. in Tr. Padzh"Bazy
Akad. Nauk SSSR, 8, 546 p. p—C. pseudomollis var. platyacantha
Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916) 149.—C. dilatata
Juz. op. cit. (1940) 547.—Exs.: H.F.A.M. No. 32.

Perennial. Plant with many stems. Stems inclined, 50-60 cm high,
arachnoid-hairy, winged throughout, with 2-3 mm wide wings, weakly
branched in upper half; branches short with capitula aggregated at tips
in heads. Leaves soft, without spines, sparsely pubescnet above, gray-
tomentose beneath, all leaves lanceolate, gradually reduced upward; basal
leaves petiolate, cauline leaves sessile, decurrent to wings, sometimes
with two pairs of teeth along margin at base. Capitula gn very short
peduncles, readily falling, narrow-lanceolate, 15 mm long, 4—6 mm wide
(excluding cusps), with 6—10-florets, pubescent. Involucral bracts 25—
30, carinate; outer bracts lanceolate, 5 mm long, rather abruptly
acuminate in short spine basally, appressed, divergent up to 2 mm in
upper half; middle bracts lanceolate, 2 mm wide and 5—7 mm long,
appressed at base, divergent above, acuminate in very short spine,
divergent part 2-3 mm long; inner bracts scarious, linear, spinescent.
Receptacular bristles smooth. Corolla yellow; anther tube violet-purple.
Achenes obovoid, almost three-angled, 3-4 mm long, toothed above.
Flowering VI-VI; fruiting VII—VIII. (Plate XIII, Fig. 1).

Loamy slopes in the foothills and lower mountain zone.—Soviet
Central Asia: Tien Shan (western), Pamiro-Alai. Endemic. Described
from Ganchulag near Tashkent. Type in Leningrad.

48 C. leptacantha (Bornm.) Juz. in Tr. Tadzh. Bazy Akad. Nauk
SSSR, 8 (1940) 547.—C. pseudomollis Wink. in Tr. Peterb. Bot. Sada,
X (1889) 470 (quoad sp. e Karatag.)—C. pseudomollis B. leptacantha
Bornm. in Beith. bot. Centralbl. XXXIV, 2 (1916) 149.

Perennial. Plants with many stems. Stem inclined, slender, 15—40
cm high, with narrow, 1-2 mm wide wings, weakly branched; branches
bearing few capitula at their tips in heads. Leaves soft, without spines,

185

183

sparsely pubescent above, gray-tomentose beneath; basal leaves petiolate,
lanceolate, undivided or sometimes pinnatipartite into narrow, oblong,
lateral lobes; cauline leaves gradually reduced toward stem apex,
lanceolate, sessile, decurrent into stem wings. Capitula on very short
peduncles, readily falling, narrow-ovoid, 12 mm long and 4 mm wide,
with few (up to 10) florets, pubescent. Involucral bracts 30—40, carinate,
imbricate, basally appressed, narrow-lanceolate, divergent in upper half,
gradually and long acuminate, divergent part 2-3 mm long and 1-1.5
mm wide; inner bracts membranous, linear, spinescent. Receptacular
bristles smooth. Corolla yellow; anther tube purple. Achenes obovoid,
somewhat triquetrous, 3.5—4 mm long, finely toothed above. Flowering
VI-VII; fruiting VII-VIII.

Loamy slopes in lower mountain zone.—Soviet Central Asia: Pamiro-
Alai (south). Endemic. Described from the vicinity of Faizabad. Type
in Leningrad.

Series 2. Fasciculares Juz.—Capitula with 8—10-florets, greenish
like other plant parts, sparsely pubescent; corolla purple. Perennial plants
with pinnatisect basal and lower cauline leaves.

49. C. fascicularis Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1,
3 (1936) 299; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 547.—C.
gillii Rech. f. in Dan. Biol. Skr. 8, 2 (1955) 99.—Ie.: Rech. f. Ibid.
figs. 71 and 78D.

Perennial. Plants with many stems. Stems erect, 30-60 cm high,
arachnoid-hairy, narrow-winged throughout, wings entire, 2-3 mm wide,
branched in upper one-third; branches long, slender, further branched
into short branchlets forming paniculate inflorescence. Leaves soft,
without spines, green and sparsely arachnoid-hairy above, gray-tomentose
beneath; basal leaves petiolate, oblong, attenuate toward both ends,
sometimes pinnatipartite into few oblong lateral lobes, then terminal
lobes much larger than laterals; cauline leaves gradually reduced towards
stem apex; all leaves lanceolate, attenuate toward both ends, sessile,
decurrent into stem wings. Capitula numerous, aggregated in groups of
four to seven into compact heads at branch tips, on short peduncles,
divergent, 12 mm long and 4 mm wide, ovoid, pubescent, with few (8—
10) florets. Involucral bracts about 30, imbricate, closely appressed at
base, lanceolate, gradually acuminate into falcate, cusp; inner bracts
membranous, often slightly broadened above, purple, spinescent.
Receptacular bristles smooth. Corolla pink. Achenes glabrous, smooth,
roundish above, 2.5 mm long. Flowering V—VI; fruiting VI-VII.

On the weathering products of varicolored rocks in the foothills —
Soviet Central Asia: Pamiro-Alai (south). General distribution:

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184

Afghanistan. Described from the Sarsaryak Range, near the village of
Margak. Type in Leningrad.

Series 3. Molles Juz.—Capitula with five or six florets, very densely
white-tomentose like other plant parts. Biennial plants.

50. C. mollis Schrenk in Bull. phys.-math. Ac. Pétersb. 3 (1845)
108; Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2, 13; Winkl. in Tr. Peterb.
Bot. Sada, XII, 200; XIV, 219; Bornm. in Beih. bot. Centralbl. XXXIV,
2, 149; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 547; Kult. in FI.
Turkm. VII, 180.—C. mollis var. latifolia Wink]. op. cit. (1892) 200.—
C. mollis B. squarrosa Bornm. op. cit. (1916) 150.—Arctium molle O.
Ktze. Rev. Gen. 1 (1891) 308.—Exs.: H.F.A.M. No. 484.

Biennial. Plants with solitary stem. Stem erect, 15-80 cm high,
densely tomentose, narrow-winged throughout, with 2-4 mm wide wings,
strongly branched; branches white-tomentose, with few (four or five)
almost sessile capitula at tips, aggregated in heads; inflorescence
corymbose-paniculate. Leaves soft, without spines, sparsely arachnoid-
hairy above, white-tomentose beneath; basal leaves petiolate, lanceolate,
pinnatisect into narrow lanceolate-linear lateral segments; cauline leaves
gradually reduced toward stem apex; lower cauline leaves like basal,
but with shorter winged petioles; middle and upper leaves sessile,
lanceolate. Capitula 15 mm long and 3-4 mm wide, narrow oblong,
sparsely arachnoid-hairy, with five or six florets. Involucral bracts 25,
white-tomentose, erect or somewhat divergent, linear, acuminate; inner
bracts membranous, deltoidly-broadened at apex, with thin, up to 1 mm
long, spine. Receptacular bristles smooth. Corolla yellowish, later pink.
Achenes obovoid, glabrous, smooth, roundish above, whitish, 2.5 mm
long. Flowering V—VI; fruiting VI-VII.

Debris slopes and loess hillocks in the foothills.—Soviet Central
Asia: Tien Shan (central and western), Pamiro-Alai, Kyzyl-Kum,
Balkhash Region. Endemic. Described from Khantau. Type in Leningrad.

Section 16. Sciadocousinia Tschern. in Bot. Mat. Gerb. Inst. Bot.
Akad. Nauk UzbSSR, XVII (1962).—Congestae Bge. in Mém. Acad.
Pétersb. 7 sér. 9, 2 (1865) 11 p. min. p.—Calcitrapae Boiss. FI. or. III
(1875) 460 p. min. p.—Xiphacanthae Winkl. in Tr. Peterb Bot. Sada,
XII (1892) 232 p. min. p.—Capitula in more or less unbellate or
paniculate inflorescences; involucral bracts with grooved, divergent or
acuminate, three-angled cusp; receptacular bristles smooth or weakly
scabrous. Corolla pink. Achenes obpyramidal, ribbed, toothed above.
Perennial monocarpic plants with large basal and smaller sessile cauline
leaves.

Type of section: C. eryngioides Boiss.

187

185

51. C. eryngioides Boiss. in Pl. Kotschy Pers. bor. (1846) No. 623,
nomen and Diagn. Pl. or. ser I, X, 101; Bge. in Mém. Acad. Pétersb. 7
sér., 9, 2, 12; Boiss. Fl. or. III, 490; Winkl. in Tr. Peterb. Bot. Sada,
XII, 236; XIV, 224; Kult, in Fl. Turkm. VII, 194.—Arctium eryngioides
O. Ktze. Rev. Gen. 1 (1891) 307.

Monocarpic perennial. Stem erect, 75-120 cm high, with remains
of old leaves at base, whitish, glossy, finely arachnoid-hairy pubescent,
sometimes glabrous, repeatedly divaricately branched in upper half,
terminating in umbellate branching of two to eight short, usually one-
headed branches; branching itself with few modified leaves at base.
Leaves on both sides, especially beneath, appressed-arachnoid-hairy; basal
leaves large, 30 cm long and 6-8 cm wide, petiolate, with thickened
petioles bearing spinescent lobes, broad-lanceolate, pinnatisect into
unequal broad-lanceolate or almost ovate, uncinate lateral lobes; cauline
leaves abruptly reduced toward stem apex, sessile, sometimes weakly
decurrent; lower cauline leaves like basal, but somewhat shorter; middle
leaves 10 cm long and 5 cm wide, pinnatipartite into deltoid, spinose-
toothed, cuspidate, lateral lobes; upper leaves 3 cm long and 2.5 cm
wide, spinose, lobed; leaves at base of branches reduced, sharply
divergent, lanceolate, with two to four spinescent teeth at base. Capitula
cylindrical, 6-7 mm wide (excluding cusps), glabrous. Involucral. bracts
few (12-14), stiff, glabrous; outer bracts 5-7-9 mm long, lanceolate,
somewhat flat, divergent or decurved; middle bracts broad-lanceolate,
somewhat flat below, graudally narrow toward apex, sulcate-deltoid,
divergent, 35-40 mm long; inner bracts coriaceous-membranous,
lanceolate, 17-20 mm long, terminating in short, thin, cusp, scabrous
along margin. Receptacular bristles unequal, longest slightly broadened
above, scabrous. Corolla pink. Achenes 4.5—5 mm long, 1.5—2 mm wide,
angular, grayish, dull, with three or four teeth above. Flowering VI-—
VU; fruiting VIJ—VIII. (Plate XIV, Fig. 2).

Loamy-gravelly slopes in the coachgrass and sheep’s fescue vegetation
zone.—Soviet Central Asia: mountainous Turkmenia (Kopetdag). General
distribution: Northern Iran, Afghanistan. Described from northern Iran,
from the Demavend Mountains. Type in Geneva.

52. C. margiana Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3
(1936) 303.

Monocarpic perennial. Stem erect, 70-75 cm high, with the remains
of old petioles, whitish, glabrous, glossy, repeatedly dichotomously
branched in upper half; branches spreading, forming lax inflorescence.
Leaves arachnoid-hairy on both sides, especially beneath; basal leaves
not known, petiolate, with thick, spiny petioles; middle cauline leaves
coriaceous, 10 cm long and 4 cm wide, petiolate, with short and slightly

188

186

flattened petioles, oblong or ovate, narrowed or roundish toward both
ends, with prominent veins, pinnatisect into unequal, broad-ovate, ovate
or orbicular, spinose-toothed lobes overlapping each other, as a result
leaf margin crisped; upper leaves reduced, decurved; leaves on
inflorescence branches strongly reduced, decurved, with one or two pairs
of teeth near base. Capitula numerous (about 100), sessile at base of
dichasium, terminal capitula on peduncles, narrow ovoid, 14-16 mm
long, 4-5 mm wide (excluding cusps), pauciflorate, finely and sparsely
arachnoid-hairy. Involucral bracts less numerous, about 25; outer bracts
ovate, appressed at base, narrowed in upper half into short, falcate, three-
angled spine; middle bracts lanceolate, 15 mm long, appressed in lower
part, weakly carinate dorsally, almost horizontal in upper one-third part,
subulately three-angled; inner bracts thin-coriaceous, lanceolate,
acuminate into short and erect cusp. Receptacular bristles smooth, slightly
flattened above. Corolla pink, exserted. Achenes narrow obpyramidal,
44.5 mm long, finely ribbed, with sharp teeth above. Flowering VIII;
fruiting IX.

Outcrops of white clay.—Soviet Central Asia: mountainous
Turkmenia (Badkhyz). Endemic. Described from the Kushka area, village
of Morgunovskii, near Sinii Spring. Type in Leningrad.

Section 17. Cousinia——Congestae Bge. in Mém. Akad. Pétersb. 7
sér 9, 2 (1865) 11 p. p—Carduiformes Bge. Ibid, 14 p. p—Pugioniferae
Bge. ibid, 30 p. p.—Rectispina Boiss. Fl. or. HI (1875) 459 p. p.—
Orthacanthae Winkl. in Tr. Peterb. Bot. Sada, XII (1892) 214 p. p.;
Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 552.—Capitula solitary
terminal on branches or crowded in inflorescences; involucral bracts
gradually acuminate into short cusp or slightly constricted in middle;
receptacular bristles smooth, weakly scabrous or scabrous. Corolla yellow,
less often whitish; anther tube yellow or purple, glabrous. Achenes
obpyramidal, obovoid, or ovoid, ribbed, obtusely roundish or with sharp
teeth above. Monocarpic perennial plants with strongly thickened, often
resinous-sticky root collar, and decurrent, strongly spinose cauline leaves.

Type of section: Type species.—C. orientalis (Adams) C. Koch.

Subsection 1. Yrano-armeniacae Takht. in Tr. Arm. Fil. Akad. Nauk
SSSR, Biol. Ser. 2 (1937) 179.—Capitula ovoid or globose, solitary;
involucral bracts narrow-lanceolate, attenuate into erect or arcuate
decurved cusp; receptacular bristles smooth.

Series 1. Chlorocephalae Takht.—Capitula oblong-ovoid or ovoid,
truncate at base, with 15—25-florets; involucral bracts green, subulately
acuminate, with erect cusp.

189

187

53. C. chlorocephala C.A.M. in DC. Prodr. VI (1837) 553; Ldb.
Fl. Ross. Il, 677; Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2, 16; Boiss. FI.
or. III, 474; Winkl. in Tr. Peterb. Bot. Sada, XII, 221; Grossh. Fl. Kavk.
IV, 166; Takht. in Tr. Arm. Fil. Akad. Nauk SSSR, Biol. Ser., 2, 180;
Fl. Erev. 319; Fl. Azerb. VIII, 363.—C. intermedia C.A.M. in DC. op.
cit. VI (1837) 553; Winkl. op. cit. XII, 222.—Arctium chlorocephalum
O. Ktze. Rev. Gen. 1 (1891) 307.—A. intermedium O. Ktze., Ibid, 308.

Perennial (sometimes monocarpic). Root woody, many-headed; root
collars white-tomentose, with remains of old petioles. Stem 15-20 cm
high, erect, finely arachnoid hairy-tomentose, glabrescent, or almost
glabrous and dark, cymosely branched in upper half. Leaves coriaceous,
sparsely pubescent and sparsely glandular above, white-tomentose
beneath; basal leaves petiolate, pinnatisect, lateral lobes unequal, linear-
lanceolate, sparsely prickly-toothed; cauline leaves gradually reduced
toward stem apex, sessile, short- or rather long-decurrent, pinnatilobate
with narrow lanceolate, spinescent lobes. Capitula oblong-ovoid, solitary
7-8 mm wide (excluding cusps), glabrous. Involucral bracts numerous
(70-100), all except inner linear, gradually subulately narrowed in erect,
squarrose, cusp, dorsally glabrous, smooth, scabrous along margin; inner
bracts narrow-lanceolate, scabrous along margin, short-acuminate.
Receptacular bristles smooth. Corolla yellow; anther tube purple. Achenes
4 mm long, obovoid, usually five-angled, toothed above. Flowering VI—
VI; fruiting VII-VIII.

Dry stony slopes in lower and middle mountain zone.—Caucasus:
Southern Transcaucasia. General distribution: Northern Iran. Described
from the vicinity of Nakhichevan. Type in Leningrad.

54. C. daralaghezica Takht. in Tr. Arm. Fil. Akad. Nauk SSSR,
Biol. Ser. 2 (1937) 181; Fl. Azerb. VIII, 363.

Perennial (or monocarpic). Root woody, many-headed; root collar
pubescent, with remains of old petioles. Stem erect, 30-40 cm high,
finely arachnoid-hairy, densely leafy, strongly cymosely branched above.
Leaves coriaceous, green above, inconspicuously glandular, with fine
glandular hairs, scattered throughout, white-tomentose beneath; basal
leaves petiolate; pinnatisect, with narrow lanceolate lateral segments with
two basal lobes; cauline leaves sessile, short-decurrent, pinnatilobate,
with narrow-lanceolate, decurved, spinescent lobes. Capitula solitary
terminal on branches, aggregated in corymbose inflorescence, ovoid, 10
mm wide (excluding cusps) and 20 mm long (with florets), glabrous.
Involucral bracts numerous, more than 100, all, except inner linear,
sparsely scabrous along margin, gradually subulately acuminate into
smooth, erect cusp; inner bracts narrow-lanceolate, gradually short-
acuminate, dorsally smooth, scabrous along margin. Receptacular bristles

190

188

smooth. Corolla yellow; anther tube purple. Achenes obovoid, 3.54
mm long, usually 10-ribbed, toothed above. Flowering VII—VIII; fruiting
VITI-IX.

Dry slopes.—Caucasus: Southern Transcaucasia. Endemic. Described
from the Daralegez, in the vicinity of Azizbekov. Type in Erevan.

Note. This species is very similar to C. chlorocephala, from which it
differs only by larger capitula and a more densely leafy and more sparsely
branched stem. However, these differences are well marked only in
extreme forms. Possibly, further study of C. chlorocephala will permit a
critical assessment of the specific significance of the distinguishing
characters of these two similar species.

Series 2. Brachypterides Takht.—Capitula ovoid or globose-ovoid,
with 30—35-florets; involucral bracts numerous, carinate, with divergent
or arcuately decurved cusp.

55. C. armena Takht. in Tr. Arm. Fil. Akad. Nauk SSSR, Biol.
Ser. 2 (1937) 182; Takht. in Fl. Erev. 318; Fl. Azerb. VIII, 366.—C.
tenuifolia auct. Fl. cauc. non C.A.M.—Exs.: Pl. or. exs. No. 222.

Perennial. Roots woody, blackish-fibrous, many-headed; root collar
with remains of old leaves. Stem erect, about 20-30 cm high, finely
arachnoid-hairy, usually light purple, divaricate almost night from base,
cymosely branched, with almost horizontal one-headed branches. Leaves
coriaceous, grayish green and finely arachnoid-hairy above, finely white-
tomentose beneath; basal leaves petiolate, pinnatisect, with narrow-
lanceolate lobes, at base usually lobed or weakly crenate-toothed
throughout; cauline leaves graudally reduced, like basal, but sessile, short-
and narrow-decurrent; uppermost cauline leaves usually pinnatilobate,
very short-decurrent. Capitula ovoid, about 9-10 mm wide (excluding
cusps) and 17-18 mm long (with florets), very slightly pubescent.
Involucral bracts numerous, all, except inner lanceolate to oblong-
lanceolate, scabrous along margin, attenuate from middle into almost
linear, weakly carinate, S—6 mm long and arcuately decurved cusp; inner
bracts narrow-lanceolate, scabrous mainly along margin, acuminate into
thin and slightly divergent cusp. Receptacular bristles smooth. Corolla
yellow; anther tube purple. Achenes obovoid, about 5 mm long,
longitudinally ribbed, with five or six ribs, irregularly toothed above.
Flowering VII-—VIII; fruiting VIII-IX. (Plate XII, Fig. 1).

Dry stony slopes.—Caucasus: Southern Transcaucasia. Endemic.
Described from the vicinity of Arzni. Type in Erevan.

Note. The plants from the Alagyoz Region (Aragats) differ somewhat
from typical C. armena by more pubescent capitula and less divergent
cusps of the involucral bracts. Considering that this is the most

189

191 Plate XII.

1—Cousinia arctioides Schrenk; 2—C. umbrosa Rg.

192

190

northwestern locality of the species, additional material and further study
of plants from this region are essential.

56. C. fedorovii Takht. in Tr. Arm. Fil. Akad. Nauk SSSR, Biol.
Sey 21937) 83.

Perennial. Root woody, blackish-fibrous, many-headed, root collar
with remains of old petioles. Stem erect, 16-25 cm high, finely
arachnoid-hairy, somewhat divaricately branched in upper part; branches
short, densely leafy, one-headed. Leaves coriaceous, grayish green, finely
arachnoid-hairy above, white-tomentose beneath; basal leaves petiolate,
12-15 cm long, pinnatisect, with unequal lateral lobes, usually finely
spinose-toothed or pinnatilobate, petiole narrow-winged, spinose-toothed;
cauline leaves gradually reduced, sessile, short-decurrent, like basal;
uppermost leaves reduced, reaching base of capitulum. Capitula globose-
ovoid, 9-10 mm wide (excluding cusps) sparsely arachnoid-hairy.
Involucral bracts numerous, ovate, narrowed from middle into linear,
sulcate, 4-5 mm long, erect, divergent cusp. Receptacular bristles smooth.
Corolla yellow; anther tube purple. Achenes obovoid, 4 mm long, narrow-
oblong, ribbed. Flowering VIIJ—VIII.

Stony dry slopes—Caucasus: Southern Transcaucasia. Endemic.
Described from the Novo-Bayazet area between Gilli and Satanakkhach.
Type in Erevan, isotype in Tbilisi.

57. C. brachyptera DC. Prodr. VI (1837) 554; Bge. in Mém. Acad.
Pétersb. 7 sér. 9, 2, 31; Boiss. F. or. III, 488; Winkl. in Tr. Peterb. Bot.
Sada, XII, 212; Grossh. Fl. Kavk. IV, 167; Takht. in Tr. Arm. Fil.
Akad. Nauk SSSR, Biol. Ser. 2, 184.—C. gundelia C. Koch in Linnaea,
XXVI (1851) 387.—C. squarrosa auct. non Boiss.: C. Koch, Ibid, (1851)
387.—Ic.: Jaub. and Spach. Ill. Pl. or. 2 (1844-1846), tab. 159.—Exs.:
Pl. or. exs. No. 47.

Perennial. Root woody, blackish-fibrous, many-headed. Stem erect,
15-40 cm high, finely arachnoid-hairy, moderately leafy, branched in
upper half; branches short, one-headed. Leaves coriaceous, green and
finely arachnoid-hairy above, white-tomentose beneath; basal leaves
petiolate, oblong-lanceolate, pinnatisect into lanceolate, acuminate-
spinescent, recurved lateral lobes; cauline leaves gradually reduced,
sessile, very short-decurrent, pinnatipartite; uppermost leaves
pinnatilobate. Capitula ovoid, 12-13 mm wide (excluding cusps), sparsely
arachnoid-hairy, all involucral bracts except inner broad-lanceolate at
base, gradually attenuate in linear, arcuate or somewhat horizontal cusps;
inner bracts narrow lanceolate, scabrous dorsally and along margin,
usually dark colored in upper part and short acuminate-spinescent.
Receptacular bristles smooth. Corolla purple; anther tube purple. Achenes

193

191

obovoid, 4.5 mm long, with narrow, longitudinal nbs, with sharp fine °
teeth above. Flowering VII; fruiting VIII.

Dry slopes in middle mountain zone.—Caucasus: Eastern and
southern Transcaucasia. General distribution: Armenia-Kurdistan.
Described from the vicinity of Erzerum. Type in Geneva.

Series 3. Decipientes Takht—Capitula ovoid, with 15—25 florets,
glabrous; involucral bracts terminating in erect cusp.

58. C. araxena Takht. in Tr. Arm. Fil. Akad. Nauk SSSR, Biol.
Ser. 2 (1937) 185; Fl. Azerb. VIII, 363.

Perennial. Roots woody, blackish-fibrous, many-headed; root collar
with remains of old leaves. Stem erect, whitish, glabrous, moderately
leafy, sparsely branched in upper part. Leaves coriaceous, green on both
sides; basal leaves petiolate, broad lanceolate, crenate-lobed, with broadly
deltoid, acute lobes, green and glabrous above, usually finely white-
tomentose beneath; cauline leaves gradually reduced, sessile, broadly
short-decurrent lanceolate or ovate-lanceolate, crenate to prickly-toothed
along margin; uppermost leaves strongly reduced. Capitula ovoid, 9 mm
wide (excluding cusps) and 18-20 mm long (with florets), glabrous.
Involucral bracts numerous, all except inner ovate to oblong-ovate, short-
ciliate along margin, distinctly attenuate from middle into narrow-linear,
almost subulate, erect cusp; inner bracts narrow-lanceolate, dorsally
scabrous, with short and thin apical spine. Receptacular bristles smooth.
Corolla yellow; anther tube creamish. Achenes obovoid, 5.5 mm long,
transversely rusose, longitudinally ribbed, with three to five ribs, toothed
above. Flowering VIII; fruitng IX.

Dry stony slopes.—Caucasus: Southern Transcaucasia. Endemic.
Described from the Nakhichevan ASSR, outskirts of the village of
Vanand. Type in Erevan.

Series 4. llicifoliae Takht.——Capitula ovoid or globose, glabrous or
pubescent; involucral bracts attenuate into subulately three-angled,
arcuately decurved or erect cusp.

59. C. erivanensis Bornm. in Vestn. Tifl. Bot. Sada, No. 30 (1913)
16; Grossh. Fl. Kavk. IV, 167; Takht. in Tr. Arm. Fil. Akad. Nauk
SSSR, Biol. Ser. 2, 187; Fl. Azerb. VIII, 368.—C. meghrica Takht.
Ibid. 186.—Ic.: Bornm. op. cit., Plate 1; Fl. Azerb. VIII, Plate 35.

Perennial. Root woody, many-headed. Stem erect, 30-60 cm high,
usually gray arachnoid-hairy, leafy, spreadingly branched above; branches
and branchlets one-headed, less often stem glabrous. Leaves coriaceous,
with network of prominent veins, gray-arachnoid hairy-tomentose on

19

&

192

both sides (more densely beneath), less often glabrous; basal leaves short-
petiolate, oblong, pinnatilobate, lobes narrow-deltoid, spinescent; cauline
leaves gradually reduced toward stem apex, sessile, short-decurrent, like
basal. Capitula globose, 15-17 mm wide (excluding cusps) or 30-35
mm wide (including cusps), densely arachnoid-hairy, narrow-lanceolate,
attenuate almost from base into subulate, erect or weakly arcuate cusp;
inner bracts narrow-lanceolate, scabrous along margin, short-acuminate.
Receptacular bristles smooth. Capitula yellow; anther tube light pink.
Achenes 5 mm long, almost tetraquetrous; toothed above. Flowering V—
VI; fruiting VII.

Dry stony slopes in the lower mountain zone.—Caucasus: Southern
and eastern Transcaucasia. General distribution: Northern Iran. Described
from the vicinity of the village of Karababa (Nakhichevan). Type in
Tbilisi.

Note. In the Megri Region, entirely glabrous plants are found together
with pubescent ones; their involucral bracts are usually arcuate. Attempts
to split off these non-pubescent plants as a separate species seem to me
unsound at present, as both pubescent and non-pubescnet forms occur
together in a population. The degree of divergence of the involucral
bracts also varies. Moreover, forms intermediate between C. lomekinii
Winkl., growing here but at a somewhat lower level, and C. erivanensis
Bornm., whose identification in many cases is difficult, are found in
these regions. I believe that this series of forms deserves detailed study
in nature.

60. C. iljinii Takht. in Tr. Arm. Fil. Akad. Nauk SSSR, Biol. Ser.
2 (1937) 187; Fl. Azerb. VIII, 368.

Perennial. Root woody, many-headed; root collar gray-tomentose,
with numerous remains of old leaves. Stem erect, finely arachnoid-hairy,
cymosely branched from middle. Leaves coriaceous, green, very finely
arachnoid-hairy above, gray-tomentose beneath; basal leaves petiolate,
oblong, 2.5—3 cm wide and 6—7 cm long, crenate-lobed, lobes broadly
deltoid, crenate-toothed with spinescent teeth; cauline leaves like basal,
but sessile, short-decurrent, gradually reduced; uppermost cauline leaves
reaching base of capitulum. Capitula ovoid, 7-8 mm wide (excluding
cusps) and 17 mm long (with corolla), finely arachnoid-hairy. Involucral
bracts numerous, narrow-lanceolate, scabrous along margin, gradually
attenuate into linear, weakly arcuate and subulate cusp; inner bracts
scabrous dorsally, lanceolate, acuminate into short cusp. Receptacular
bristles smooth. Corolla yellow; anther tube light purple. Achenes
obovoid, 6 mm long (young). Flowering VI.

Gypsic slopes in lower mountain zone.—Caucasus: Southern
Transcaucasia. Endemic. Described from the vicinity of Ordubad.

195

193

Note. The type of the species could not be found; only plants from
the Dzhulfa Region are known. These plants actually differ from all
known species, but their differences are insignificant; additional material
is necessary to understand some of the morphological characters and
the range of the species.

61. C. lomakinii Winkl. in Tr. Peterb. Bot. Sada, XIV (1897) 220;
Grossh. Fl. Kavk. IV, 166; Takht. in Tr. Arm. Fil. Akad. Nauk SSSR,
Biol. Ser. 2, 188; Fl. Azerb. VIII, 369.

Perennial. Root woody, blackish-fibrous, many-headed, root collar
white-tomentose, with remains of old petioles. Stem erect, glabrous,
somewhat purple, divaricately branched, leafy. Leaves coriaceous, green
on both sides, with fine network of prominent veins, finely glandular;
basal leaves oblong, petiolate, crenate-lobed, lobes deltoid, sparsely
spinose; cauline leaves gradually reduced upward, sessile, decurrent,
lanceolate, crenate-toothed, prickly-toothed. Capitula ovoid, 8-10 mm
wide (excluding cusps), glabrous. Involucral bracts numerous, basally
appressed, gradually attenuate into subulate, usually divergent or almost
horizontal cusps, with short and stiff hairs on dorsal surface and along
margin, sometimes glandular; inner bracts narrow-lanceolate, densely
covered with short and stiff cilia along margin, or with weakly divergent
cusps. Receptacular bristles smooth. Corolla yellow; anther tube purple.
Achenes obovoid, 4 mm long, narrowly longitudinally ribbed, truncate
above, finely toothed. Flowering VI-—VII; fruiting VIII.

Dry slopes in lower mountain zone.—Caucasus: Southern
Transcaucasia. Endemic. Described from the Araks Valley. Type in
Leningrad.

62. C. hohenackeri Fisch. and Mey. Ind. sem. hort. Petrop. 2 (1835)
34; DC. Prodr. VI, 555; Ldb. FI. Ross. II, 677; Bge. in Mém. Acad.
Pétersb. 7 sér. 9, 2, 21; Boiss. Fl. or. III, 481; Winkl. in Tr. Peterb.
Bot. Sada, XII, 211; Grossh. Fl. Kavk. IV, 166; Takht. in Tr. Arm. Fil.
Akad. Nauk SSSR, Bicol. Ser. 2, 188; Fl. Azerb. VIII, 369.—Arctium
hohenackeri O. Ktze. Rev. Gen. 1 (1891) 307.

Perennial. Root woody, blackish-fibrous, many-headed. Stem about
25-30 cm high, finely arachnoid-hairy, usually purple at least in upper
part, divaricately branched, moderately leafy. Leaves coriaceous, finely
arachnoid-hairy and green above, gray-tomentose beneath; basal leaves
petiolate, pinnatisect in narrow-lanceolate, decurved lateral lobes; petiole
narrow-winged; cauline leaves sessile, very short-decurrent or almost
nondecurrent, semiamplexicaul, almost horizontal, pinnatilobate, or
pinnatipartite, gradually reduced toward stem apex. Capitula ovoid, 9—
10 mm wide (excluding cusps), sparsely arachnoid-hairy. Involucral bracts

196

194

numerous, all except inner lanceolate, weakly carinate, dorsally smooth,
scabrous along margin, gradually attenuate in falcate cusps; inner bracts
lanceolate, dorsally scabrous, gradually short-acuminate. Receptacular
bristles smooth. Corolla yellow; anther tube pink. Achenes obovoid,
about 4 mm long, longitudinally mbbed, transversely pitted, and with
sharp teeth above. Flowering VI-—VII; fruiting VII.

Dry slopes in middle mountain zone.—Caucasus: Eastern
Transcaucasia, Talysh. General distribution: Iran. Described from Talysh.
Type in Leningrad.

Subsection 2. Irano-turkestanicae Takht. in Tr. Arm. Fil. Akad.
Nauk SSSR, Biol. Ser. 2 (1937) 178.—Capitula ovoid or cylindrical,
solitary or aggregated on branch apices. Involucral bracts lanceolate;
receptacular bristles scabrous or smooth.

Series 5. Carduiformes Takht.—Plants medium high. Capitula
ovoid-oblong or oblong-cylindrical, usually aggregated on branch apices,
glabrous; involucral bracts narrow-lanceolate, acuminate into erect cusp;
receptacular bristles smooth.

63. C. orientalis (Adams) C. Koch in Linnaea, XXIV (1851) 386;
Fl. Azerb. VIII, 364; Fl. Gruzi, VIII, 444.—Carduus orientalis Adams
in Weber and Mohr. Beitr. 1 (1805) 63; MB. FI. taur.-cauc. 2, 270.—
Cousinia carduiformis Cass. in Dict. sc. nat. XLVII (1826) 503; DC.
Prodr. VI, 552; Ldb. FI. Ross. II, 676; Bge. in Mém. Acad. Pétersb. 7
sér. 9, 2, 18; Boiss. Fl. or. III, 472; Winkl. in Tr. Peterb. Bot. Sada,
XII, 224; Takht. in Tr. Arm. Fil. Akad. Nauk SSSR, Biol. Ser. 2, 179.—
C. orientalis Grossh. Fl. Kavk. IV, 166.—Arctium carduiforme O. Ktze.
Rev. Gen. 1 (1891) 307.—Exs.: Pl. or. exs. No. 248.

Perennial. Roots woody, blackish-fibrous. Stem erect, white arachnoid-
hairy, 10-25 cm high, usually cymosely branched; branches bearing two
to five capitula aggregated at tip, usually slightly enclosed by apical leaves.
Leaves coriaceous, green above, glabrous or inconspicuously sparsely
glandular, white-tomentose beneath; basal and lower cauline leaves
petiolate, oblong, pinnatilobate, lobes broadly deltoid, spinescent; cauline
leaves sessile, broadly short-decurrent, slightly reduced toward stem apex.
Capitula ovoid-oblong, 9-10 mm wide and 20-25 mm long (with
projecting corolla), almost without arachnoid-hairy pubescence. Involucral
bracts 40-45, narrow-lanceolate, gradually acuminate into short and stiff,
erect, cusp, weakly carinate dorsally, densely scabrous; inner bracts
scarious, acuminate in thin cusp. Receptacular bristles smooth. Corolla
yellow; anther tube purple. Achenes obovoid, about 5 mm long,
longitudinally ribbed, transversely rugose, toothed above. Flowering VII-—
VIII; fruiting VITI-IX.

197

195

Dry slopes from lower to middle mountain zone.—Caucasus:
Ciscaucasia, eastern and southern Transcaucasia. General distribution:
Turkey. Described from the vicinity of Tbilisi. Type in Leningrad.

64. C. microcephala C.A.M. in DC. Prodr. VI (1837) 552; Bge. in
Mém. Acad. Pétersb. 7 sér., 9, 2, 16; Boiss. Fl. or. III, 472; Winkl. in
Tr. Peterb. Bot. Sada, XII, 219; Grossh. Fl. Kavk. IV, 166; Takht. in
Tr. Arm. Fil. Akad. Nauk SSSR, Biol. Ser. 2, 179; Fl. Azerb. VIII,
365.—Arctium microcephalum O. Ktze. Rev. Gen. 1 (1891) 308.

Perennial. Root woody, blackish-fibrous. Stem numerous, slender,
20-30(40) cm high, finely arachnoid-hairy or almost glabrous. Leaves
coriaceous, green, and very finely arachnoid-hairy above, white-tomentose
beneath; basal leaves short-petiolate, lanceolate, crenate-pinnatilobate or
crenate-pinnatipartite, with broadly deltoid, spinose-toothed lobes; cauline
leaves gradually reduced, sessile, short-decurrent, oblong or lanceolate,
shallow crenate-prickly toothed. Capitula oblong-cylindrical, 4-5 mm
wide (excluding cusps) and 12-17 mm long, finely arachnoid-hairy, with
fewer florets, solitary or in twos on short peduncles; involucral bracts
coriaceous, somewhat flat, scabrous along, lanceolate, gradually short-
acuminate, with erect cusp; inner bracts thinner, membranous, scabrous
dorsally and along margin, acuminate in thin and short cusp. Receptacular
bristles smooth. Corolla yellow; anther tube purple. Achenes obovoid,
44.5 mm long, longitudinally narrow-ribbed, transversely rugose, with
sharp teeth above. Flowering VI; fruiting VI-VII.

Dry stony or limestone slopes in lower and middle mountain zone.—
Caucasus: Eastern and southern Transcaucasia. General distribution:
Northern Iran. Described from northern Iran. Type in Leningrad.

Series 6. Eudolicholepides Juz—Short or often medium high plants.
Capitula solitary terminal on branches, broadly ovoid; involucral bracts
dorsally smooth; receptacular bristles scabrous.

65. C. dolicholepis Schrenk in Bull. phys.-math. Ac. sc. Pétersb. 3
(1845) 107; Ldb. FI. Ross. II, 681; Bge. in Mém. Acad. Pétersb. 7 sér.,
9, 2, 18; Winkl. in Tr. Peterb. Bot. Sada, XII, 232 p. p.; XIV, 224;
Bornm. in Beih. bot. Centralbl. XXXIV, 2, 158 p. p.; Juz. in Tr. Inst.
Kauch. i Gutt. 4, 11.—C. semenovii Rgl. in Bull. Soc. Nat. Mosc. XI, 3
(1867) 147; Winkl. op. cit. (1892) 232.—Arctium dolicholepis O. Ktze.
Rev. Gen. 1 (1891) 307.—A. semenowii O. Ktze. Ibid. 308.

Perennial. Roots vertical, with thick, 1-2 cm in diameter, collar
covered by remains of old petioles. Stems one to four, erect or weakly
divergent, 15-35 cm long, rather densely arachnoid-hairy scabrous,
somewhat oblong-sulcate, repeatedly divaricately branched from base or

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196

middle, forming broad corymbose-paniculate inflorescence. Basal leaves
short-petiolate, with 1—2 cm long petioles, broad-lanceolate, lamina 6—
20 cm long and 1.5—9 cm wide (including lobes), slightly attenuate
toward both ends, sinuate, pinnatilobate or pinnatisect with oblique,
almost ovate, spinescent-acuminate, almost entire lobes or lobes with
one or two teeth at base; lower cauline leaves like basal, but sessile;
middle and upper leaves gradually reduced, sessile, short-decurrent;
uppermost leaves narrow ovate, with few teeth; all leaves arachnoid
hairy-tomentose on both sides, more densely beneath, with prominent
veins. Capitula ovoid or broad-ovoid, 15-25 mm long and 6-17 mm
wide (excluding cusps), usually with foveate base, arachnoid-hairy or
tomentose. Involucral bracts about 40, ciliate, dorsally, except inner
bracts, smooth; outer bracts lanceolate, divergent; middle abruptly
narrowed from ovate base into long, almost horizontal, deltoid cusp;
inner bracts linear-lanceolate, erect, membranous, gradually acuminate
with thin and short spine at apex. Receptacular bristles scabrous. Corolla
yellow. Achenes 3—6 mm long, 1.5—2.5 mm wide, obpyramidal, four or
five-angled, ribbed, rugose-pitted, and with fine teeth above. Flowering
V-VI; fruiting VII.

Sands, alkaline valleys, wormwood steppes, wastelands and weedy
places.—Soviet Central Asia: Balkhash Region, Tien Shan (central),
Dzhungaria-Tarbagatai (Dzhungarian Alatau). Endemic. Described from
Khantau. Type in Leningrad.

Series 7. Scabridae Juz—Plants medium high. Capitula solitary
terminal on branches, ovoid to almost globose; involucral bracts more
or less scabrous dorsally; receptacular bristles smooth.

66. C. vicaria Kult. in Tr. Turk. Nauchn. Obshch. 1 (1923) 113;
Juz. in Tr. Inst. Kauch. i Gutt. 4, 15—C. pskemensis Juz. nomen Ibid.
(1932) 16 and Herb.—C. decurrens f. tschimganica O. Fedtsch. in Perech.
Rast. Turkm. IV (1911) 250.—C. polycephala auct. non Rupr.: Bornm.
in Beth. bot. Centralbl. XXXIV, 2 (1916) 156.—Ie.: Kult. op. cit. 115.—
Exs.: H.F.A.M. No. 50.

Perennial. Root vertical, woody, blackish-fibrous, with thickened
collar. Stem solitary, erect, 60-75 cm high, somewhat striate-sulcate,
arachnoid-hairy, strongly branched from base, with long, spreading-erect
branches, forming corymbose-paniculate inflorescence. Basal leaves
coriaceous, long petiolate (with petioles 5-8 cm long), oblong-lanceolate,
pinnatilobate, with narrow deltoid, acuminate-spinescent lobes, broadened
at base, bearing one or two large, acute prickly teeth along margin on
both sides; lower cauline leaves like basal, but sessile, long-decurrent;
middle and upper leaves ovate, decurrent. Capitula numerous, 8-13 mm

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197

wide (excluding cusps) and 17-23 mm long, ovoid, arachnoid-hairy.
Involucral bracts scabrous dorsally; outer bracts deltoid, acuminate in
small spine; middle bracts ovate, 2.5-3 mm wide at base, gradually
narrowed into short, almost deltoid, erect cusp; inner bracts membranous,
linear-lanceolate, usually dark colored above, mucronate. Receptacular
bristles smooth. Corolla yellow. Achenes obpyramidal, almost triquetrous.
Flowering VI-VII; fruiting VII—VIII.

Loessic slopes in middle mountain zone.—Soviet Central Asia: Tien
Shan (western). Endemic. Described from the vicinity of Khumsan. Type
in Leningrad.

67. C. angreni Juz. in Tr. Inst. Kauch. 1 Gutt. 4 (1932) 17.

Perennial. Root vertical, woody, blackish-fibrous, with thickened,
somewhat oblong collar. Stem usually solitary, erect, 30-60 cm high,
striate-sulcate, weakly arachnoid-hairy, branched in upper half, with
spreading branches bearing solitary capitulum. Basal leaves petiolate,
oblong-lanceolate, pinnatisect into unequal, divergent lateral segments,
in turn prickly-lobed, lower cauline leaves like basal but sessile; middle
leaves oblong, short-decurrent, pinnatipartite into relatively narrow-
deltoid, spinescent-acuminate lateral lobes; upper leaves ovate, broadly
short-decurrent, prickly-lobed. Capitula less numerous, almost globose,
18-20 mm wide (excluding cusps), 25 mm long, arachnoid-hairy.
Involucral bracts somewhat divergent, even somewhat falcate, dorsally
carinate, scabrous; outer bracts oblong-ovate, with thin, up to 3 mm
long cusp; middle bracts oblong, gradually acuminate into thin up to 6
mm long cusp; inner bracts almost membranous, linear, with thin and
short cusp. Receptacular bristles smooth. Corolla yellow; anther tube
purple. Achenes 6 mm long and 4 mm wide, with strongly prominent
ribs, narrow-winged along ribs, with four teeth above. Flowering VI-—
VII; fruiting VII—VIII.

Loamy-gravelly slopes in lower mountain zone.—Soviet Central
Asia: Tien Shan (western). Endemic. Described from the Angren River
Basin between lIertash and Lachenkiya. Type in Leningrad.

68. C. scabrida Juz. in Tr. Inst. Kauch. i. Gutt. 4 (1932) 13.—C.
dolicholepis auct. non Schrenk: Winkl. in Tr. Peterb. Bot. Sada, XII
(1892) 232 p.p.

Perennial. Root vertical, fibrous, with thickened collar about 2 cm
wide. Stems two, erect, 25-35 cm high, densely arachnoid-hairy, shallow
sulcate, branched; branches erect and spreading, in turn branched further.
Basal leaves petiolate, with 2—3.5 cm long petioles, oblong-lanceolate,
lamina 10-18 cm long, 3-4.5 cm wide, pinnatisect into narrow deltoid
lobes, cuspidate, with one or two large teeth at base; lower and middle

200

202

198

cauline leaves like basal but sessile, short and narrow decurrent, gtadually
reduced toward stem apex; upper leaves narrow-ovate or oblong-ovate,
almost undivided, cuspidate, crenate-lobulate or strongly toothed. Capitula
less numerous, solitary on peduncle, broadly ovoid or ovoid, 15-25 mm
long and 10-15 mm wide, foveate at base, weakly arachnoid-hairy.
Involucral bracts about 50, strongly or weakly scabrous in lower part on
dorsal surface, outermost bracts narrowed in erect cusp from appressed
ovate base; middle bracts with longer, almost three-angled, erect or
slightly recurved cusp, not exceeding capitulum; inner bracts slightly
shorter than or as long as middle, membranous, slightly broadened toward
apex, with almost roundish apex, finely spinescent. Receptacular bristles
smooth. Corolla yellow. Achenes 6 mm long and 3 mm wide,
obpyramidal, ribbed, ribs usually winged, toothed above. Flowering VI-—
VU; fruiting VII-IX.

Dry gravelly slopes in lower and middle mountain zone.—Soviet
Central Asia: Tien Shan (Talass Alatau, Karatau, Tashkent Alatau).
Endemic. Described from the Koktal River Basin. Type in Leningrad.

Note. The type was not designated in the description of the species;
therefore, | propose to consider for the lectotype the plant: “Near Kcoktal
Brook. Knorring. 1909, Vol. 26, No. 376.”

69. C. minkwitziae Bornm. in Beih. bot. Centralbl. XXXIV, 2
(1916) 158; Juz in Tr. Inst. Kauch. 1 Gutt, 4, 14.

Perennial. Koot vertical, blackish-fibrous, with thickened collar, 2—
2.5 cm wide. Stem erect, 30-60 cm high, finely arachnoid-hairy, striate-
sulcate, usually cymosely branched in upper part, branches and branchlets
one-headed. Leaves green, finely arachnoid-hairy above, gray-tomentose,
with sharply prominent veins beneath; basal leaves petiolate, with 2—5
cm long petioles, oblong-lanceolate, with lamina 18-25 cm long and 4—
5.5 cm wide, pinnatisect into narrow-deltoid, spinose lateral lobes, in
turn, pinnatilobate; lower cauline leaves like basal but with shorter
petioles; middle leaves somewhat reduced, morphologically like basal
but sessile, semiamplexicaul and very short-decurrent; upper cauline
leaves ovate, cuspidate, spinose-toothed. Capitula pedunculate, broadly
cylindrical, 25-35 mm long and (12)17-20 mm wide, weakly foveate at
base, very slightly arachnoid-hairy. Involucral bracts about 60, dorsally
somewhat scabrous only at base; outer bracts oblong-ovate, acuminate,
divergent; middle bracts oblong-lanceolate, flat, dorsally carinate,
cuspidat, usually longer than inner bracts and also corolla; inner bracts
membriunous; somewhat acuminate, scabrous along margin. Corolla
yellow. anther tube light pink. Receptacular bristles smooth. Achenes
5-7 mm long and 3 mm wide, obpyramidal, triquetrous or almost
tetraquetrous, usually winged along edges, toothed above. Flowering VI-—
VI; firutiing VIL.

199

Plate XIII.
1 (2a)—Cousinia pseudomollis Winkl; 2—C. anomala Franchet.

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200

Loamy-gravelly meadow-steppe slopes in middle mountain zone.—
Soviet Central Asia: Tien Shan (Talass Range; Aksu-Dzhebogly Reserve).
Endemic. Described from Aksai. Type in Leningrad.

70. C. tenuisecta Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3
(1936) 306.

Perennial. Root vertical, blackish-fibrous, with thickened collar. Stem
erect, white-pubescent, (18)25—45 cm high, leafy, cymosely branched in
upper part. Leaves green, finely arachnoid-hairy above, white-tomentose
beneath, basal and lower cauline leaves petiolate, oblong, bipinnatisect,
with primary segments narrow-lanceolate, spinescent, on narrow-winged
petioles, secondary segments almost linear, spinescent-acuminate,
decurved; cauline leaves gradually reduced, sessile, semiamplexicaul,
narrow and short-decurrent. Capitula ovoid, 12-15 mm wide (excluding
cusps), arachnoid-hairy, with somewhat foveate base. Involucral bracts

_ about 40; outer bracts oblong-ovate, attenuate into divergent strong cusp,

203

very inconspicuously scabrous below on dorsal surface; middle bracts
narrow-lanceolate, erect-spreading, dorsally carinate, weakly scabrous
below; inner bracts membranous, lanceolate, short-acuminate, scabrous
along margin. Receptacular bristles smooth. Corolla yellow; anther tube
pink. Achenes three or four-angled, narrow-winged along edges, weakly
rugose, 4-5 mm long, toothed above. Flowering VI—VII; fruiting VIII.

Loamy slopes.—Soviet Central Asia: Tien Shan (Kirgizian Alatau,
Ichkeletau, Karatau). Endemic. Described from the Botomainak Ravine.
Type in Leningrad.

Series 8. Turkestanicae Juz—Plants short or medium high; basal
leaves petiolate, without spines in lower part. Capitula more or less
aggregated in heads, narrow ovoid; receptacular bristles smooth.

71. C. syrdariensis Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIlIc,
III (1928) 106; Juz. in Tr. Inst. Kauch. i Gutt. 4, 32 p. p—Exs.:
H.F.A.M. No. 488.

Perennial. Root vertical, slender, with thickened collar. Stem usually
solitary, erect, 10-45 cm high, arachnoid-hairy, striate-sulcate, branched
from middle; branches forming dense, corymbose-paniculate
inflorescence. Basal leaves petiolate, coriaceous, lanceolate, pinnatipartite
or sometimes pinnatisect into deltoid prickly-toothed lateral segments,
attenuate-spinescent, finely arachnoid-hairy, grayish-green, above, white-
tomentose beneath with prominent veins; lower cauline leaves like basal
but sessile, short- and broad-decurrent; middle and upper leaves oblong-
ovate, sessile, short-decurrent, crenate-lobed, with acuminate-spinescent:
lobes; uppermost leaves almost deltoid, with less numerous large teeth

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201

on each side. Capitula numerous, compactly aggregated in groups of 4—
10 on tips of primary and secondary branches, almost sessile, narrow-
ovoid, with roundish or somewhat narrowed base, small, 10-13 mm
long and 3-4 mm wide (excluding cusps), finely arachnoid-hairy.
Involucral bracts about 20, squarrose, weakly carinate; outer bracts
lanceolate, gradually acuminate into thin short cusp; middle bracts two
times as long as outer, narrow-lanceolate, gradually acuminate-spinescent
with thin spine; inner bracts membranous, linear, with finely spinescent.
Receptacular bristles smooth. Corolla pale yellow; anther tube pink.
Achenes flat-obpyramidal, almost three or four-angled, with four teeth
above, somewhat truncate, 3 mm long and 1.5 mm wide. Flowering
VI-VII; fruiting VII—VIII.

In foothills—Soviet Central Asia: Syr-Darya, Tien Shan (western).
Endemic. Described from the vicinity of Tashkent. Type in Tashkent.

72. C. turkestanica (Rgl.) Juz. in Tr. Inst. Kauch. 1 Gutt. 4 (1932)
35.—C. decurrens y. turkestanica Rgl. in Tr. Peterb. Bot. Sada, IV (1880)
316; Winkl. Ibid, XII, 221.

Perennial. Plants very similar to C. syrdariensis Kult., from which
it differs by larger, 17 mm long and 6 mm wide (excluding cusps)
capitula, broader, at base 2.5 mm wide involucral bracts, larger, 4—4.5
mm long, achenes and almost solitary pedunculate capitula. Flowering
VI; fruiting VII.

In foothills—Soviet Central Asia: Tien Shan (Karatau). Endemic.
Described from Karatau, Turtchi. Type in Leningrad.

Note. Apparently, it hybridizes with C. scabrida Juz.; a large number
of intermediate forms from the Karatau Range can be found in herbaria.

Series 9. Pungentes Juz.—Plants medium high; basal leaves with
unarmed petioles. Capitula solitary, broadly ovoid; involucral bracts
smooth; middle bracts acuminate into carinate, strong, divergent cusp;
receptacular bristles scabrous.

73. C. juzepezukii Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 95.

Perennial. Root woody, blackish-fibrous, with strongly thickened,
almost globose, white-tomentose collar. Stem erect, 40-60 cm high,
white-tomentose, divaricately branched almost from base, branches, in
turn, further branched, into short one-headed secondary branches. Leaves
coriaceous, finely arachnoid-hairy above, finely tomentose beneath, with
prominent veins; basal leaves short-petiolate, with unarmed petiole,
oblong, pinnatilobate, lobes broadly deltoid, spinescent, spinose-toothed,
somewhat approximate, as a result leaf margin more or less crisped;

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202

cauline leaves gradually reduced, sessile; lower cauline leaves like basal
but somewhat shorter; middle leaves sessile, semiamplexicaul, with broad
roundish auricles, spinose-toothed; upper leaves strongly reduced.
Capitula globose, 20 mm wide (excluding cusps), slightly foveate at
base. Involucral bracts about 50, all densely scabrous dorsally, outer
bracts broad-lanceolate, somewhat flat, gradually acuminate in carinate,
falcate, and rather broad cusp with thin spine, 22-25 mm long; middle
bracts erect, like outer, 25-30 mm long; inner bracts narrow lanceolate,
membranous, with thin and short cusp, 20-22 mm long. Receptacular
bristles scabrous. Corolla yellow; anther tube pink. Achenes obovoid,
usually three or four-angled, narrow-winged along edges, with three or
four teeth above, 5 mm long, 2.5—3 mm wide. Fruiting X.

Stony slopes of monadnock mountains.—Soviet Central Asia: Kyzyl-
Kum. Endemic. Described from Kyzyl-Kum, Aktau Mountains. Type in
Leningrad.

74. C. umbilicata Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1,
3 (1936) 306.

Perennial. Plants rather tall. Stem erect, white-arachnoid-hairy,
sparsely branched; branches short, one-headed. Leaves coriaceous, finely
arachnoid-hairy, green above, white-tomentose beneath, upper leaves
ovate, broadly short-decurrent, crenate-prickly toothed; uppermost leaves
strongly reduced, reaching almost to base of capitula. Capitula broad
ovoid or almost globose, 25 mm long (with corollas) and 12-15 mm
wide (excluding cusps), arachnoid-hairy, with rather distinctly foveate
base. Involucral bracts 25-27; outer bracts oblong-ovate, somewhat flat
at base, short-acuminate, with erect-deflected cusp; middle bracts broad-
lanceolate, somewhat flat in lower half, 5-6 mm wide, narrowed from
middle in carinate, acuminate-spinescent, erect-delfected cusp, usually
17-25 mm long; inner bracts membranous, lanceolate, short acuminate
into fine cusp, cusp erect, 17-18 mm long. Receptacular bristles unequal,
longest slightly broadened above, scabrous. Corolla yellow; anther tube
purple. Achenes obovoid, 5 mm long, almost three-angled, with sharp
teeth above. Flowering VI; fruiting VII.

On sandy compacted slopes.—Soviet Central Asia: Kyzyl-Kum.
Endemic. Described from the vicinity of Besh-Bulak. Type in Leningrad.

75. C. pungens Juz. in Tr. Inst. Kauch. 1 Gutt. 4 (1932) 19; Juz. in
Tr. Tadzh. Bazy Akad Nauk SSSR, 8, 552.—C. dolicholepis auct. non
Schrenk; Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916) 158 p. p.

Perennial. Root vertical, blackish-fibrous, with globose thickened
collar. Stem erect, 20-40 cm high, shallow-sulcate, rather densely
arachnoid hairy-lanate, with few weakly branched, erect branches, bearing
less numerous solitary capitula. Basal leaves petiolate with unarmed

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203

petioles, coriaceous, lanceolate, lobed, with broadly deltoid acuminate-
spinescent lobes, finely arachnoid hairy-tomentose, grayish-green above,
densely white-tomentose beneath, with prominent veins; lower cauline
leaves like basal but somewhat shorter and sessile; middle and upper
leaves ovate, short-decurrent, large prickly-toothed; uppermost leaves
strongly reduced. Capitula less numerous (about 20), broadly ovoid or
almost globose, about 20 mm long and 12 mm wide (excluding cusps),
somewhat roundish at base, finely arachnoid-hairy. Involucral bracts less
numerous, wide; outer bracts short, gradually acuminate from wide
appressed base into short cusp; middle bracts two to three times as long
as outer, abruptly acuminate from broad ovate appressed base or almost
attenuate into long, almost three-angled, strong, erect or weakly recurved
cusp; inner bracts linear, scarious above, with thin cusp. Receptacular
bristles broadened above, scabrous. Corolla yellow. Achenes obpyramidal,
almost four-angled, with sharp teeth above, about 4 mm long. Flowering
VI-VII; fruiting VII-VIU.

In foothills and lower mountain zone.—Soviet Central Asia: Pamiro-
Alai (northern slope of the Alai Range). Endemic. Described from the
Shahimardan River Basin, Ankhor gorge. Type in Leningrad.

76. C. scleracantha Kult. in Bot. Mat. Gerb. Inst. Bot. Akad. Nauk
UzbSSR, XVII (1962) 104.

Perennial. Root vertical, blackish-fibrous, with thickened collar. Stem
solitary, erect, 30-60 cm high, densely white-tomentose, branched from
base, branches convergent, bearing capitula at tips in semicorymbose
inflorescence. Cauline leaves sessile, broadly long-decurrent, broad
oblong-ovate to ovate and even orbicular, large-spinose-toothed or even
prickly-lobed, very finely arachnoid-hairy above, white-tomentose beneath
with prominent veins. Capitula large, 13-14 mm wide (excluding cusps),
arachnoid-hairy. Involucral bracts carinate; outer bracts lanceolate, short-
acuminate; middle bracts 35 mm long, lanceolate, somewhat carinate,
gradually broadly acuminate into grooved, almost falcate cusp; inner
bracts finely coriaceous, almost membranous, gradually acuminate.
Receptacular bristles scabrous. Corolla yellow, not exserted from
involucre. Achenes obovoid, 5 mm long, indistinctly ribbed. Flowering
VII; fruiting VIII.

In varicolored lower hills——Soviet Central Asia: Tien Shan (foothills
of the Fergana Range). Endemic. Described from the vicinity of Turpak-
Bel Pass. Type in Leningrad.

Series 10. Polycephalae Juz.—Plants tall; petioles of basal leaves
spinose along sides throughout. Two or three capitula aggregated on
branch tips, others solitary, usually ovoid; receptacular bristles usually
smooth.

204

77. C. polycephala Rupr. in Mém. Acad. Pétersb. 7 sér. 14, 4 (1869)
54; Winkl. in Tr. Peterb. Bot. Sada, XII, 219; XIV, 199; Juz. in Tr.
Inst. Kauch. i Gutt. 4, 21; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR 8,
552.—Arctium polycephalum O. Ktze. Rev. Gen. 1 (1891) 308—C.
decurrens auct. Winkl. op. cit. (1892) 220 p. p.; XIV, 222 p. p—C.
decurrens var. congesta auct. non Rgl.: O. and B. Fedtsch. Perech. Rast.
Turkm. IV (1911) 249.—C. decurrens B. oligocephala auct. non Rgl.:
Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916) 157 p. p.

Perennial. Root vertical, woody, fibrous, with thickened collar 25—
40 mm wide. Stem solitary, erect, up to 100 cm high, more or less
arachnoid-hairy, branched from middle or above with long, erect
branches, in turn branched forming a rather wide inflorescence, usually
wider than height of secondary branches. Basal leaves usually withering
by anthesis and fruiting, 7—20 in sterile rosette, 15-50 cm long and 5—
11 cm wide (with spines), coriaceous, oblong-lanceolate, slightly crisped
along margin, pinnatisect, lobes not approximate, deltoid, large toothed,
attenuate into strong spine, green and sparsely arachnoid-hairy above,
more densely along midrib, densely and finely white-tomentose beneath,
petiolate, with petioles thickened at base, bearing erect spreading spines
on sides; cauline leaves reduced towards stem apex; lower cauline leaves
like basal but sessile; middle and upper cauline leaves narrow-ovate or
broad-lanceolate, sessile, narrowly long-decurrent, prickly-toothed,
attenuate into strong cusp. Capitula numerous; lower capitula solitary,
upper in groups of two to four on branch tips, short- or rather long-
pedunculate, ovoid or narrowly ovoid, with roundish base, 18-22 mm
long and 6—9 mm wide, pubescent. Involucral bracts about 25, herbaceous
or almost coriaceous; outer bracts ovate, dorsally carinate, acuminate in
short cusp; adjacent bracts gradually transitional into middle bracts in
shape and size, three times or more exceeding outer bracts, narrowed
from narrow ovate base in rather long, green, strong, almost three-angled
cusp, erect or weakly curved, as long as or shorter than florets; inner
bracts herbaceous, lanceolate; innermost membranous, straw-yellow,
linear, slightly broadened toward apex, acuminate into long and thin
spine. Receptacular bristles whitish, almost smooth or with one or two
serrations. Corolla yellow; anther tube purple. Achenes 4-5 mm
long, obpyramidal, flattened three or four-angled, sinuate-ribbed, with
somewhat sharp three or four teeth above. Flowering VIJ—VIII; fruiting
Vil.

In foothills and lower mountain zone, on stony and clayey-stony
slopes, in wormwood and grass-wormwood steppes and on wastelands.—
Soviet Central Asia: Tien Shan, Pamiro-Alai (Alai Range, southern
Tadzhikistan). General distribution: Dzhungaria-Kashgaria. Described
from the Chu River Basin. Type in Leningrad.

208

205

78. C. finitima Juz. in Tr. Inst. Kauch. i Gutt. 4 (1932) 24; Juz.
in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 552.—C. decurrens B.
oligocephala auct. non Rgl.: Bornm. in Beih. bot. Centralbl. XXXIV, 2
(1916) 157 p. p.

Perennial. In all respects plants similar or C. polycephala Rupr.,
differing from the latter by slightly flexuous inflorescence branches;
longer, somewhat curved cusps of middle involucral bracts, exceeding
florets by 5 mm; and almost spatulate inner involucral bracts. Flowering
VII; fruiting VIJ-VIII.

Valleys of mountain rivers—Soviet Central Asia: Pamiro-Alai.
Endemic. Described from the middle reaches of the Kshtut River Valley
(“Vorukh”). Type in Leningrad.

79. C. sporadocephala Juz. in Tr. Bot. Inst. Akad. Nauk SSSR,
Ser. 1, 3 (1936) 307.

Perennial. Roots vertical, woody, grayish-fibrous, with globose,
thickened collar. Stem solitary, erect, striate-sulcate, tomentose, 40—60
cm high, with less numerous long branches, in turn, branched forming
lax paniculate inflorescence; basal leaves petiolate, with unarmed petioles,
oblong-lanceolate, coriaceous, unequally narrow lobed, with acuminate-
spinescent, spinose-toothed lobes, weakly arachnoid-hairy above, gray-
tomentose beneath, with prominent veins; lower cauline leaves like basal
but somewhat shorter, sessile and long-decurrent; middle and upper leaves
ovate, broadly decurrent, crenate-prickly toothed, pubescent like basal
leaves. Capitula comparatively less numerous, with distinct peduncles,
broadly ovoid, pubescent, roundish or somewhat narrowed at base, 20
mm long and 10 mm wide (excluding cusps). Involucral bracts 20,
squarrose; outer bracts broad-ovate, appressed, with short divergent cusp;
middle bracts oblong-ovate, appressed, attenuate in somewhat carinate,
strong and divergent cusp, almost as long as appressed base; inner bracts
linear, membranous above, with thin short cusp. Receptacular bristles
weakly flattened above, almost smooth. Corolla yellow. Achenes
obpyramidal, with four teeth above, 4 mm long. Flowering VII—VIII;
fruiting VIII-IX.

Loamy slopes in foothills.—Soviet Central Asia: Tien Shan.
Endemic. Described from Maili-Sai. Type in Leningrad.

Series 11. Resinosae Juz.—Plants mostly medium tall; basal leaves
with petioles without spines in lower part. Capitula more or less clustered,
ovoid, narrow-ovoid or almost cylindrical; receptacular bristles scabrous,
weakly scabrous or smooth.

80. C. spiridonovii Juz. in Tr. Inst. Kauch. 1 Gutt. 4 (1932) 29;

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206

Juz. in Tr. Tadzh. Bazy Akad Nauk SSSR, 8, 552.—C. decurrens y.
oocephala Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916) 157.

Perennial. Root vertical, woody, blackish-fibrous, thickened into
globose resinous-sticky collar. Stem usually solitary, erect, 30-40 cm
high, densely white-tomentose, branched in upper half, with branches
forming corymbose-paniculate inflorescence. Basal leaves petiolate,
oblong-lanceolate, lobed, with broadly deltoid lobes acuminate into small
spine, coriaceous, densely white-tomentose beneath, finely arachnoid-
hairy above; lower cauline leaves sessile, otherwise like basal leaves;
middle and upper leaves ovate or almost orbicular, sessile, broadly short-
decurrent, large crenate-prickly-toothed, white-tomentose beneath, finely
arachnoid-hairy above. Capitula numerous, broadly ovoid or ovoid, 8—
10 mm wide (excluding cusps) and 17-22 mm long, densely arachnoid
hairy-lanate. Involucral bracts erect-spreading, carinate, with thin and
rather strong short cusp; outer bracts oblong-ovate, with short thin cusp,
sometimes with two pairs of acicular spines along margin; middle bracts
oblong, gradually acuminate into stronger, up to 3 mm long cusp; inner
bracts lanceolate, with thin and very short cusp. Receptacular bristles
weakly flattened above, scabrous. Corolla yellow; anther tube purple.
Achenes obovoid, 5 mm long and 3 mm wide. Flowering VI-—VII; fruiting
VU-VIl.

Clayey deserts—Soviet Central Asia: Syr-Darya, Pamiro-Alai?
Endemic. Described from the vicinity of the village of Nikolskii. Type
in Leningrad.

81. C. transoxana Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 105.

Perennial. Root vertical, woody, blackish-fibrous with globose,
resinous-sticky, weakly pubescent collar. Stem solitary, erect, 20-50 cm
high, densely arachnoid hairy-tomentose, branched in upper half; branches
short, bearing one or two capitula at tips of very short peduncles, forming
almost corymbose-paniculate inflorescence. Basal leaves short-petiolate,
oblong-lanceolate, prickly-lobed, with narrow deltoid strongly acuminate-
spinescent lobes, coriaceous, finely arachnoid-hairy above, white-
tomentose beneath; lower cauline leaves like basal but sessile, excurrent
over stem; middie and upper cauline leaves oblong-ovate or almost
orbicular, large spinose-toothed with cuspidate teeth, broadly decurrent.
Capitula relatively less numerous, broadly ovoid, 7-9 mm wide
(excluding cusps) and 15—18 mm long, arachnoid hairy-lanate. Involucral
bracts erect-spreading or slightly recurved, somewhat flat at base,
narrowed toward apex in strong, carinate cusp; outer and middle bracts
6-17 mm long; inner bracts membranous, lanceolate, short-acuminate.
Receptacular bristles almost smooth. Corolla yellow; anther tube purple.

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207

Achenes obovoid, longitudinally ribbed, almost three-angled. Flowering
VI-VII; fruiting VII.

Clayey deserts and foothills —Soviet Central Asia: Amu-Darya,
Kyzyl-Kum, Pamiro-Alai. Endemic. Described from the vicinity of the
village of Talimardzhan. Type in Tashkent.

82. C. ambigens Juz. in Tr. Inst. Kauch. 1 Gutt. 4 (1932) 17.

Perennial. Root vertical, woody with globose collar. Stem erect,
30-60 cm high, white arachnoid-hairy, divaricately branched almost from
base. Basal leaves 25-30 cm long, oblong, petiolate, with unarmed
petioles, pinnatipartite, with unequal, acuminate-spinescent lateral lobes,
green, finely arachnoid-hairy above, finely tomentose beneath with
prominent veins; cauline leaves gradually reduced, oblong-ovate to almost
orbicular, sessile, broadly short-decurrent, crenate-prickly-toothed.
Capitula broadly ovoid or almost globose, 17 mm long and 12 mm
wide (excluding cusps), arachnoid-hairy, pedunculate. Involucral bracts
somewhat flat, dorsally smooth; outer bracts about 10 mm long,
squarrose, acuminate; middle bracts 17 mm long, lanceolate, slightly
attenuate in middle, terminating into carinate, deltoid, erect cusp; inner
bracts lanceolate, 15 mm long, membranous,. mucronate. Receptacular
bristles unequal, longest slightly flattened above, scabrous. Corolla
yellow; anther tube pink. Achenes obovoid, 4.5—5 mm long and 2 mm
wide, almost three-angled, with sharp teeth above. Flowering VI-—VII;
fruiting VII—VIII.

In foothills —Soviet Central Asia: Tien Shan (foothills of the
Kuraminskii and Chatkal ranges). Endemic. Described from Dzhambulak.
Type in Leningrad.

83. C. resinosa Juz. in Tr. Inst. Kauch. i Gutt. 4 (1932) 26; Juz. in
Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 552.—C. decurrens auct. non
Rgl.: Winkl. in Tr. Peterb. Bot. Sada, XII (1892) 220 p. p.: XIV, 222. —
C. decurrens var. congesta auct. non Rgl.: Winkl. op. cit. (1892) 220 p.
p-, Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916) 156 p. max p.—
Ic.: Sornye Rast. SSSR, 4, Fig. 472.

Perennial. Root vertical, woody, blackish-fibrous, with globose,
resinous-sticky, weakly arachnoid-hairy collar. Stem solitary, 15-60 cm
high, erect, densely white-tomentose, branched in upper half or sometimes
almost from base, with erect-spreading branches, forming corymbose-
paniculate inflorescence. Basal leaves petiolate, oblong-lanceolate,
coriaceous, grayish green above due to thin, appressed, white tomentum,
densely white-tomentose beneath with strongly raised midrib and lateral
veins, lobed with deltoid spinescent lobes, lobes somewhat approximate,
as a result leaf margin appearing crisped; lower cauline leaves like basal

PMN

208

but sessile, decurrent; middle and upper leaves ovate or broad-ovate,
sessile, broadly long-decurrent, large and prickly-crenate-toothed,
pubescent like basal leaves. Capitula numerous, almost sessile or on
very short peduncles, in groups of two or three at branch tips, ovoid,
medium-sized, 18-23 mm long and 7-8 mm wide (excluding cusps),
densely pubescent. Involucral bracts about 25, herbaceous, squarrose;
outer bracts oblong-ovate, with short, up to 1 mm long, strong cusp;
middle bracts two times as long as outer, lanceolate, with strong, up to
10 mm long cusp; inner bracts linear, membranous above, with thin and
short cusp. Receptacular bristles unequal, at least some scabrous above.
Corolla yellow; anther tube purple. Achenes obovoid, dorsally
compressed, finely pitted, with sharp teeth above, 4.5 mm long and 2
mm wide. Flowering VIIJ—VIII; fruiting VIII-IX. (Plate XV, Fig. 2).

Loamy slopes in foothills and lower mountain zone.—Soviet Central
Asia: Pamiro-Alai, Tien Shan (foothills of western Tien Shan). Endemic.
Described from the vicinity of the village of Kuropatkino. Type in
Leningrad.

Note. A highly polymorphic species, whose range apparently is being
extended by human activity. Further study of this complex of species
on extensive material is essential. Undoubtedly, numerous hybrids of
these species are present in the belt of contact with C. maracandica
Juz.

84. C. kazachorum Juz. ex Tschern. in Bot. Mat. Gerb. Inst. Bot.
Akad. Nauk UzbSSR, XVII (1962) 96.

Perennial. Root vertical, woody, with globose, arachnoid-hairy collar.
Stem usually solitary, erect, 25-40 cm high, flocculose-tomentose,
cymosely branched from middle with short branches bearing two or
three capitula at tips, usually short peduncles. Leaves coriaceous, green,
sparsely arachnoid-hairy above, white-tomentose beneath, with prominent
veins; basal leaves short-petiolate, with thick and unarmed petioles,
oblong-lanceolate, pinnatilobate, with somewhat approximate broadly
deltoid, spinescent lobes; cauline leaves gradually reduced, lower leaves
like basal but sessile; middle and upper cauline leaves oblong-ovate or
broad lanceolate, sessile, broadly long-decurrent along stem, prickly-
toothed; uppermost leaves strongly reduced. Capitula broadly ovoid, 8—
10 mm wide (excluding cusps), densely arachnoid-hairy. Involucral bracts
about 25-30; outer and middle bracts ovate or oblong-ovate, somewhat
flat, from middle attenuate into slightly carinate, divergent cusp, from
7-20 mm long; inner bracts scarious, narrow-lanceolate, gradually
acuminate into short and thin cusp, 17 mm long. Receptacular bristles
somewhat scabrous. Corolla yellow; anther tube pink. Achenes obovoid,
5.5-6 mm long, longitudinally mbbed, almost three-angled. with sharp
teeth above, rugose-pitted. Flowering VI-—VII; fruiting VIII-IX.

209

Plate XIV.

1—Cousinia egregia Juz.; 2—C. eryngioides Boiss.

211

213

210

Loamy slopes in foothills.—Soviet Central Asia: Tien Shan
(Kirgizian Alatau, Ichkeletau, Karatau), Balkhash Region (Muyunkums).
Endemic. Described from the Muyunkums, region of the Dyuksembai
well. Type in Tashkent.

85. C. horridula Juz. in Tr. Inst. Kauch. i Gutt. 4 (1932) 30; Juz.
in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 553.—C. decurrens var.
scabriseta Winkl. in Tr. Peterb. Bot. Sada, XIV (1897) 222.

Perennial. Root vertical, fibrous, with somewhat thickened collar.
Stem solitary, erect, shallow-sulcate, tomentose-woolly, branched from
base; branches erect-spreading, in turn, branched forming broad
corymbose-paniculate inflorescence. Basal leaves petiolate, oblong-
lanceolate, coriaceous, light green above, with scarcely visible
pubescence, densely white-tomentose beneath, deeply crenate-lobed, with
numerous narrow deltoid lobes, terminating in long, stiff spine; lower
cauline leaves like basal but sessile, decurrent; middle and upper cauline
leaves oblong-ovate or broad-ovate, broadly and rather long-decurrent,
with large teeth and narrow deltoid teeth terminating in long spine;
leaves light green above, very finely arachnoid-hairy, almost glabrous,
white-tomentose beneath. Capitula numerous, almost sessile, aggregated
at branch tips, narrow ovoid, 6 mm wide and 20 mm long. Involucral
bracts somewhat squarrose, herbaceous, except outer, weakly and very
sparsely arachnoid-hairy, gradually narrowed from ovate appressed base
to rather wide, dorsally carinate, smooth cusp; outermost bracts with
two or three pairs of acicular spines along margin, with strong, almost
three-angled cusp; middle bracts lanceolate, somewhat attenuate in
middle, gradually acuminate in strong cusp; inner bracts membranous,
linear, with very short and thin cusp. Receptacular bristles weakly serrate
above. Corolla yellow, anther tube pink. Achenes obovoid, somewhat
three-anged, toothed above, 4 mm long. Flowering VI-VII; fruiting VII—
VII.

In foothills and lower mountain zone.—Soviet Central Asia: Pamito-
Alai (northern slope of the Turkestan Range). Endemic. Described from
Uratyube. Type in Leningrad.

86. C. maracandica Juz. in Tr. Inst. Kauch. i Gutt. 4 (1932) 30;
Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 9, 553.—C. decurrens var.
subcabriseta Winkl. ex. O. and B. Fedtsch. Perech. Rast. Turk. V (1911)
250.

Perennial. Root vertical, woody, covered with fibrous-black bark,
with thickened resinous-sticky, weakly pubescent collar. Stem solitary,
25-80 mm high, erect, densely white-tomentose, branched in upper half
with branches forming corymbose-paniculate inflorescence. Basal leaves
petiolate, oblong-lanceolate, light green above, very finely pubescent,

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white-tomentose beneath with prominent veins, irregularly large spinose-
toothed or lobed, with acuminate-spinescent deltoid lobes; lower cauline
leaves like basal but sessile, decurrent; middle and upper cauline leaves
oblong-ovate, long-decurrent, large toothed with acuminate-spinescent
teeth. Capitula numerous, on very short peduncles, densely aggregated
at branch tips, narrow cylindrical, 5 mm wide and 17 mm long,
pubescent. Involucral bracts about 25, herbaceous, somewhat flat, closely
appressed, with erect cusp; outer bracts half as long as middle, ovate,
with thin glabrous cusp; middle bracts oblong-ovate, gradually
acuminate-spinescent; inner bracts linear, membranous, finely cuspidate.
Receptacular bristles smooth, sometimes with only one or two pairs of
serrations above. Corolla yellow; anther tube purple. Achenes cylindrical,
almost three-angled, toothed above, 4 mm long. Flowering VI—VII;
fruiting VII-VIII. .

Outcrops of varicolored rocks in middle mountain zone.—Soviet
Central Asia: Pamiro-Alai. Endemic. Described from the Zeravshan
Valley. Type in Leningrad.

Note. Numerous hybrids of C. maracandica and C. resinosa Juz.
are known from the regions where they grow together.

87 C. platystegia Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 103.

Perennial. Root vertical, woody, with thick collar. Stem few, erect,
20-25 cm high, arachnoid-hairy, leafy, slightly branched at apex, with
short branches bearing two to five capitula at tips. Basal leaves usually
petiolate, with broad bases; cauline leaves sessile, broadly long-decurrent,
oblong or lanceolate, spinose-toothed, with short-acuminate teeth, finely
pubescent above, white-tomentose beneath, gradually reduced upwards.
Capitula cylindrical, 6 mm wide (excluding cusps), very finely arachnoid-
hairy. Involucral bracts less numerous (10-15); outer and middle bracts
broad lanceolate, 3-3.5 mm wide at base, somewhat boat-shaped,
carinate, broadly acuminate, strongly divergent in upper half, 15-23 mm
long; inner bracts linear-lanceolate, membranous, short-acuminate, 15—
17 mm long. Receptacular bristles entirely smooth. Corolla yellow, not
exserted from capitulum. Achenes obovoid, almost three-angled, shallow
pitted-rugose, 5 mm long. Flowering VI-—VII; fruiting VII.

Outcrops of varicolored rocks in lower hills——Soviet Central Asia:
Pamiro-Alai (Shirabad Valley). Endemic. Described from the Shirabad
Valley, Boz-Rabat-Zagon. Type in Tashkent.

Note. An entirely isolated species of the section, known only from
a single specimen.

Series 12. Egentes Juz.—Plants short or medium high. Capitula
ovoid or cylindrical, aggregated at tips of branches; involucral bracts

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212

somewhat flat, smooth, short and finely acuminate; receptacular bristles
smooth.

88. C. dichromata Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIIIc, VI
(1929) 14; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 553.—C.
cisdarvasica V. Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XIX (1959) 385.—Ie.: Kult. op. cit. Fig. 13.

Perennial (Monocarpic). Root vertical, woody, covered with blackish-
fibrous bark; collar thickened. Stem usually solitary, erect, 30-80 cm
high, white-tomentose, leafy throughout, branched in upper half; branches
with closely aggregated capitula at tips (10 each). Leaves coriaceous,
white-tomentose beneath, green and glossy above, pubescent only along
veins; basal leaves petiolate, oblong-ovate, broadly deltoid-lobed, lobes
spinose-toothed, acuminate-spinescent; all cauline leaves sessile, broadly
long-decurrent, ovate or almost orbicular, deltoid-spinose-toothed.
Capitula numerous, 18-25 mm long and 4—6 mm wide, almost sessile,
cylindrical, gray-tomentose, with fewer florets (four or five). Involucral
bracts coriaceous, imbricate, closely appressed at base, with short,
recurved stiff cusp; inner bracts membranous, linear, distinctly broadened
at apex, with very thin and short cusp. Receptacular bristles smooth.
Corolla whitish; anther tube purple. Achenes ovoid, indistinctly ribbed,
roundish above, with two teeth, whitish, 4 mm long and 2 mm wide.
Flowering VI-—VII; fruiting VII—VIII. (Plate XV, Fig. 2).

Clayey and gravelly slopes, on outcrops of varicolored rocks in
foothills ——Soviet Central Asia: Pamiro-Alai (south). Endemic. Described
from Baisuntau, between Vakhshur and Denau. Type in Tashkent.

89. C. egens Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3
(1936) 304; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 553.

Perennial (Monocarpic). Roots vertical, woody, blackish-fibrous; root
collar thick, up to 2 cm in diameter; stem usually solitary, erect, 30-35
cm high, finely sulcate, densely arachnoid-hairy, discontinuously winged
with long-decurrent leaves, repeatedly branched in upper half; branches
spreading with capitula aggregated at tips, forming rather wide,
semicorymbose inflorescence. Leaves coriaceous, green and lustrous
above with floccose pubescence mainly along veins, white-tomentose
beneath; basal leaves petiolate, with 3-4 cm long somewhat thick
petioles, white arachnoid-hairy, oblong-lanceolate, with 16—20 cm long
and 5—8 cm wide lamina, narrowed toward both ends, prickly lobed,
lobes deltoid, spinose-toothed; cauline leaves like basal but sessile,
broadly long-decurrent on stem, gradually reduced towards apex. Capitula
on short peduncles, ovoid, somewhat truncate at base, strongly narrowed
at apex, with fewer florets (three to five) with 20 mm long and about 6

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213

mm wide florets, densely arachnoid-hairy. Involcural bracts about 20,
all except inner closely appressed, ovate to oblong-ovate, acuminate
into short erect spine; inner bracts membranous, linear-lanceolate or
linear, erect, short-acuminate, scabrous along margin. Receptacular
bristles smooth. Corolla yellow; anther tube purple. Achenes obovoid,
slightly flattened, 5 mm long, 2 mm wide, indistinctly ribbed, toothed
above, forming very small corona. Flowering VI—VII; fruiting VII.

Clayey dry slopes in foothills—Soviet Central Asia: Pamiro-Alai
(south). Endemic. Described from the vicinity of the Shagaon Post. Type
in Leningrad.

Series 13. Congestae Tschern.—Plants high; basal leaves with almost
unarmed petioles. Capitula narrow cylindrical, numerous, aggregated in
heads; involucral bracts somewhat flat, short and finely acuminate;
receptacular bristles smooth; achenes without pappus.

90. C. congesta Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2 (1865) 12;
Boiss. Fl. or. III, 469; Winkl. in Tr. Peterb. Bot. Sada, XI, 132; XII,
221; XIV, 222; Kult in Fl. Turkm. VII, 203.

Perennial. Roots vertical, woody, blackish-fibrous, with globose
collars, often with many stems from base or stem solitary, floccose-
finely arachnoid-hairy, longitudinally-sulcate, 80-100 cm high. Leaves
coriaceous, almost glabrous and green above, densely white-tomentose
beneath with prominent veins; basal leaves about 40 cm long, 10 cm
wide, oblong-lanceolate, crenate-lobed, lobes broadly deltoid or almost
orbicular, spinose-toothed, somewhat approximate, short-petiolate, with
thick unarmed petioles; lower cauline leaves like basal but smaller; other
cauline leaves gradually reduced, sessile, broadly short-decurrent, oblong
or almost ovate, crenate-prickly-toothed, sparsely glandular; uppermost
leaves strongly reduced, reaching up to capitula, but not enclosing them.
Capitula terminal on branches in groups of 15-40, sessile, 20 mm long
and 4—5 mm wide, oblong-cylindrical, usually with five florets, very
finely arachnoid-hairy or almost glabrous. Involucral bracts green, flat,
all erect; outer bracts oblong-ovate, somewhat acuminate, scabrous along
margin, closely appressed; middle bracts oblong-lanceolate, about 18
mm long, gradually and short acuminate, about 3 mm wide at base,
scabrous along margin; inner bracts coriaceous, whitish, lustrous, 18-19
mm long, linear, acuminate-spinescent. Receptacular bristles white,
smooth. Corolla yellow; anther tube glabrous, pink. Achenes obpyramidal,
5—6 mm long, 2—2.5 mm wide, with three or four ribs, narrowly winged
along ribs, with small toothed corona above, white, without pappus.
Flowering V—VI; fruiting VII. 3

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Loamy slopes in foothills and plains at the foot of mountains,
often as a ruderal weed.—Soviet Central Asia: mountainous Turkmenia
(Kopetdag, Badkhyz, Bolshie [Great] Balkhany). General distribution:
Iran. Described from northern Iran. Type in Leningrad.

91. C. agelocephala Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 89.

Perennial. Root vertical, woody, blackish-fibrous, with strongly
thickened, 3-6 cm wide, usually tomentose collar. Stems several, often
solitary, erect, white-tomentose, 30-80 cm high, divaricately branched
from middle; branches short, bearing numerous (up to 40) capitula at
tips aggregated in heads. Leaves coriaceous, green and finely arachnoid-
hairy above, white-tomentose beneath; basal leaves petiolate, with almost
unarmed petioles, oblong-lanceolate, pinnatilobate, with unequal,
acuminate-spinescent lobes; lower cauline leaves like basal, but with
shorter petioles or almost sessile; middle leaves broad-lanceolate, sessile,
very short-decurrent or not decurrent, semiamplexicaul, crenate-prickly
toothed, upper leaves almost orbicular, sessile, semiamplexicaul, with
broad roundish auricles, spinescent, crenate-prickly-toothed, reaching up
to head, appearing as common involucre, slightly exceeding heads in
length. Capitula narrow-cylindrical, 20-22 mm long and 5—6 mm wide,
usually with 10-11 florets, sessile, finely arachnoid-hairy, usually with
floral leaves modified into coriaceous, lanceolate toothed leaflets.
Involucral bracts green, flat, erect; outer bracts oblong, somewhat
acuminate, scabrous along margin, closely appressed; middle bracts
oblong-lanceolate, 17-20 mm long, gradually and short-acuminate,
scabrous along margin; inner bracts coriaceous, whitish, glossy, roundish
above, with thin short cusp, 17 mm long. Receptacular bristles white,
smooth. Corolla yellow; anther tube glabrous, pink. Achenes obpyramidal,
4-5 mm long, ribbed, narrow-winged along ribs, with small toothed
corona, without pappus. Flowering VI; fruiting VII.

Sandy and gravelly stony slopes.—Soviet Central Asia: Pamiro-Alai
(Buryutau Plateau). Endemic. Described from Buryutau. Type in
Leningrad.

92. C. androssovii Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1,
3 (1936) 305.

Perennial (Monocarpic?). Apparently very tall plant, with thick,
almost 2 cm thick, striate-sulcate, floccose-arachnoid hairy, sparsely
branched stem; branches long, further branched and bearing numerous
capitula aggregated in lax paniculate inflorescence. Leaves coriaceous,
green above, white-tomentose beneath, with prominent veins; cauline
leaves large, 20-50 cm long, 12-13 cm wide, pinnatilobate, with broadly

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deltoid, short-acuminate lateral lobes, sessile, broadly short-decurrent;
upper cauline leaves reduced. Capitula ovoid, in groups of five to seven
at tips of secondary branches, pedunculate, glabrous, 7-8 mm wide,
slightly attenuate at base or roundish; involucral bracts ovate to oblong-
ovate, closely appressed with each other, with only slightly divergent
short cusp; all bracts flat, green, scabrous along margin, sparsely glandular
dorsally; inner bracts membranous, white above with short cusp.
Receptacular bristles smooth, white. Corolla yellow, anther tube purple.
Achenes obovoid, ribbed, narrowly winged along ribs. Flowering VI.

On the slopes of hills—Soviet Central Asia: mountainous Turkmenia
(Badkhyz). Endemic. Described from the vicinity of Kushka. Type in
Leningrad.

Note. A very unique species, known only from a single location.

Section 18. Abolinia Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 81.—Capitula aggregated at branch tips;
involucral bracts gradually acuminate into stiff, falcate or thin, erect
cusp, and then inner bracts with membranous appendages; receptacular
bristles smooth; corolla whitish or pale yellow; achenes obovoid, smooth,
roundish above. Perennial plants with large pinnatisect or pinnatipartite
basal leaves.

Type of section: C. abolinii Kult.

93. C. abolinii Kult. ex Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 88.

Perennial. Roots vertical, woody, with thickened woody collar. Stem
erect, apparently tall, striate-sulcate, white arachnoid-hairy, up to 15
mm thick at base, branched almost from base. Capitula terminal on
stem and branches aggregated into compact racemose inflorescence.
Leaves finely arachnoid-hairy above, white-tomentose beneath with
prominent veins; basal leaves long-petiolate, with thick and weakly
sulcate 12-15 cm long petioles, oblong, about 30 cm long and 12 cm
wide (including spines), pinnatisect into oblong, prickly-lobed lateral
segments; lower leaves like basal; middle leaves gradually reduced,
sessile; upper leaves strongly reduced. Capitula ovoid, sessile, with
slightly narrowed base, 12-15 mm wide and 25 mm long (excluding
projecting corollas), with many florets, glabrous. Involucral bracts
numerous, all except inner coriaceous; outer bracts narrow-lanceolate,
closely appressed at base, gradually attenuate toward apex into thin,
almost falcate cusp, scabrous along margin; middle bracts cuspidate,
somewhat erect-spreading, scabrous along margin; inner bracts in many
rows, membranous, broad-lanceolate, acuminate in fine cusp.
Receptacular bristles short, white, smooth. Corolla whitish or pale yellow,

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216

exserted. Achenes large, obovoid, roundish above, 7 mm long. Flowering
VI.

Stony slopes.—Soviet Central Asia: Tien Shan (Naryn Basin).
Endemic. Described from the Nizhnii Naryn Basin, from Garpsai. Type
in Tashkent.

94. C. egregia Juz. in Bot. Mat. Gerb. Glavn. Bot. Sada, V (1924)
110.

Perennial. Root vertical, woody, stem erect, 1—1.20 mm high, striate-
sulcate, floccose-tomentose, up to 15 mm thick at base, weakly branched;
branches weakly divergent from main stem, each with two or three
capitula forming narrow paniculate inflorescence in upper half of stem.
Basal leaves long-petiolate, with triquetrous, floccose-tomentose, 10—15
cm long petioles, oblong-lanceolate, 27 cm long and 11 cm wide (including
spines), coriaceous, with sharply prominent midrib, pinnatipartite,
sometimes even pinnatisect on lower side into oblong, prickly-toothed
lateral lobes, white-tomentose beneath, finely arachnoid-hairy above; lower
and middle cauline leaves like basal, but slightly shorter, middle leaves
also sessile and gradually reduced upward; upper leaves sparse, oblong,
sessile, prickly-lobed. Capitula 15—25, ovoid, somewhat foveate at base,
13-17 mm wide (excluding cusps) and 30 mm long, with many florets,
on rather long peduncles, sometimes almost sessile, glabrous. Involucral
bracts numerous, except inner coriaceous, oblong-lanceolate, gradually
acuminate; all bracts erect, imbricate, closely appressed to each other,
acuminate into short, thin, and weakly squarrose cusp, glabrous, glossy,
scabrous along margin; inner bracts membranous, exceeding middle and
appearing to form collar, lanceolate, almost roundish above, with very
thin and short cusp. Receptacular bristles short, smooth. Corolla white,
exserted from involucre. Achenes obovoid, dorsally compressed, roundish
above, dark brown, 8—9 mm long. Flowering VI; fruiting VII. (Plate XIV,
Fig. 1).

Stony slopes in middle mountain zone.—Soviet Central Asia: Tien
Shan (Angren River Valley). Endemic. Described from Nurek-Ata in
the Chirchik River Basin. Type in Leningrad.

95. C. dolichophylla Kult. in Tr. Turk. Nauchn. Obshch. 1 (1923)
112.—Ie.: Kult. Ibid., tab. (2).

Perennial. Root vertical, woody, many-headed. Stem erect, 60—90
cm tall, indistinctly striate-sulcate, arachnoid-hairy, almost tomentose
above, sparsely branched; branches erect-spreading, slender, with several
terminal capitula forming lax inflorescence. Basal leaves long-petiolate,
with triquetrous, floccose-tomentose, up to 10 cm long, petioles, oblong-
lanceolate, usually 25-30 cm long and 5—8 cm wide, coriaceous, very

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217

indistinctly arachnoid-hairy above, green, white-tomentose beneath,
pinnatipartite, almost pinnatisect in lower half into lanceolate, prickly-
toothed, sometimes prickly-lobed, lateral lobes short acuminate in thin
spine; lower cauline leaves like basal, but slightly shorter; middle leaves
like lower, but sessile gradually reduced towards stem apex; uppermost
leaves sparse, sessile, prickly-lobed, small. Capitula numerous, narrow
ovoid, sometimes with somewhat foveate base, 10-12 mm wide
(excluding cusps) and 25 mm long, with many florets, on short peduncles,
sometimes sessile, arachnoid-hairy. Involucral bracts numerous, all except
inner coriaceous, narrow lanceolate, slightly carinate, short-acuminate,
all bracts erect; middle bracts narrowly scarious-fimbriate, scabrous along
margin; inner bracts with membranous, finely toothed, lobed, spatulate,
their appendages usually divided in two halves in middle and slightly
recurved, forming small collar. Receptacular bristles smooth. Corolla
exserted from involucre, whitish (?). Achenes obpyramidal, compressed,
roundish above, 6-7 mm long and 3 mm wide. Flowering VII-VIII;
fruiting VITI-IX.

Slopes in middle mountain zone.—Soviet Central Asia: Tien Shan
(western). Endemic. Described from Korzhantau, vicinity of Khumsan.
Type in Tashkent.

Section 19. Eriocousinia Tschern. in Bot. Mat. Gerb. Inst. Bot.
Akad. Nauk UzbSSR, XVII (1962) 82.—Odontocarpae Winkl. in Tr.
Peterb. Bot. Sada, XII (1892) 253 p. p.; Juz. in Tr. Tadzh. Bazy Akad.
Nauk SSSR, 8, 562.—Heteracanthae auct. non Winkl.: Juz. in Tr. Tadzh.
Bazy Akad. Nauk SSSR, 8 (1940) 551 p. p—Capitula solitary terminal
on stem or branches; involucral bracts gradually acuminate in stiff
subulate cusp, or somewhat flat with membranous appendages, irregularly
toothed or coriaceous ciliate-toothed; receptacular bristles smooth or
scabrous. Corolla pink or purple. Achenes obovoid, glabrous, smooth,
roundish above, or with scarcely visible filiform longitudinal ribs and
in that case with scarcely visible teeth. Perennial herbs with many-headed
woody root and coriaceous, pinnatisect or pinnatipartite basal leaves.

Type of section: C. pannosa Wink.

Series 1. Pannosae Juz.—Involucral bracts lanceolate, acuminate
into stiff cusp, without appendages; receptacular bristles scabrous. Cauline
leaves short-decurrent.

96. C. pannosa Winkl. in Tr. Peterb. Bot. Sada, X (1889) 578; XII,
257; XIV, 229; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 562.—C.
franchetii auct. non Winkl.: Bornm. in Beih. bot. Centralbl. XXXIV, 2
(1916) 184 p. p—C. resupinata Juz. in Tr. Bot. Inst. Akad. Nauk SSSR,
Ser. 1, 3 (1936) 318; Juz. op. cit. 562—C. pannosa var. microcephala

—

218

Winkl. op. cit. (1897) 229.—C. franchetii 8. drepanophora Bornm. op.
cit. (1916) 184.

Perennial. Root woody, many-headed; caudices with less numerous
remains of previous year’s leaves at base. Stem numerous, white-
tomentose, one-headed, leafy, erect, 25-60 cm high. Leaves oblong,
white or gray-tomentose on both sides, sometimes with thinner tomentum
above; basal leaves petiolate, irregularly pinnatipartite in deltoid, usually
approximate, acuminate-spinescent lateral lobes, with two lanceolate
acuminate-spinescent lobes at base, lobes 15—18 mm long in middle
part of leaf and 10-15 mm wide near base (broadest place); cauline
leaves gradually reduced, like basal, but sessile, short-decurrent,
auriculate. Capitula almost globose, 15-20 mm wide (excluding cusps),
very densely rusty- or white-tomentose. Involucral bracts about 70-80;
outer bracts horizontal or decurved, 15-25 mm long, lanceolate;
acuminate in stiff spine; middle bracts (15)20 mm long, erect or
divergent, acuminate; inner bracts membranous, 18-20 mm long, narrow-
linear, with very thin and short apical spine. Corolla purple. Receptacular
bristles scabrous. Achenes obovoid, about 6 mm long and 3 mm wide,
indistinctly longitudinally ribbed. Flowering VIII-IX; fruiting IX.

Stony slopes in upper mountain zone.—Soviet Central Asia: Pamiro-
Alai (Gissar Range, Peter the First Range, Darvaz, Shugnan, southern
slopes of the Alai Range, western part of the Trans-Alai Range).
Endemic. Described from Sagyrdasht Ravine. Type in Leningrad.

Note. The plant from the Alai Range has smaller capitula and is
more sparsely pubescent; it is necessary to study additional material.

97. C. franchetii Winkl. in Tr. Peterb. Bot. Sada, X (1889) 578;
XII, 257; XIV, 208; Bornm. in Beih. bot. Centralbl. XXXIV, 2, 184 p.
p.; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 562.—C. canescens
auct. non DC.; Franch. in Ann. Sc. Nat. Bot. 6 sér. 16 (1883) 323.—C.
pannosa var. subviridis Winkl. op. cit. (1897) 229.—Arctium
franchetianum O. Ktze. Rev. Gen. 1 (1891) 307.

Perennial. Root woody, many-headed; caudices with remains of
previous year’s leaves at base. Stems numerous, white-tomentose, one-
headed, leafy, erect, 15-30 cm high. Leaves oblong, grayish-green above
due to appressed-arachnoid-hairy pubescence, gray-tomentose beneath;
basal leaves short-petiolate, irregularly pinnatipartite into narrow-
lanceolate, often sparse attenuate-spinescent lateral lobes, usually with
two lateral spines at base, about 4-5 mm wide at base (broadest place)
and about 15 mm long (in middle part of leaf); cauline leaves gradually
reduced towards apex, like basal but sessile, short-decurrent. Capitula
cylindrical, 15—18 mm wide (excluding cusps), sparsely rusty-arachnoid-
hairy. Involucral bracts 50-60, erect, sometimes outermost bracts

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219

horizontal; outer and middle bracts lanceolate, about 15 mm long,
acuminate-spinescent; inner bracts membranous, linear, with thin apical
spine, usually dark colored on dorsal surface, 16-17 mm long. Corolla
purple. Receptacular bristles scabrous. Achenes obovoid, about 6 mm
long, 3 mm wide, indistinctly longitudinally ribbed. Flowering VI-—VII;
fruiting VII-VIII.

Stony-gravelly slopes in upper mountain zone.—Soviet Central Asia:
Pamiro-Alai (Zeravshan Range, northern slopes of the Gissar Range).
Endemic. Described from the Zeravshan Valley “Kirschlak” (Capus).
Type in Paris.

Note. A species with a rather narrow range extending from Lake
Iskander-Kul to the upper reaches of the Yagnob and seemingly
penetrating the range of the closely related species C. pannosa Winkl.,
with which it apparently forms hybrids especially in the western part of
the range. This species differs from C. pannosa mainly by the much
more sparsely pubescent capitula, erect involucral bracts, and lateral
lobes of the leaves that approach narrowly lanceolate. However, this
species varies slightly in its own characters within the limits of its range.

98. C. pannosiformis Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 101.

Perennial. Root woody, many-headed; caudices with numerous
remains of previous year’s leaves. Stems erect, erect-spreading upward,
one-headed, white arachnoid-hairy, 15—40 cm high, sparsely leafy. Leaves
gray-arachnoid-hairy on both sides, more densely beneath; basal leaves
petiolate, irregularly pinnatipartite in deltoid, acuminate-spinescent lateral
lobes with two small spines at base; cauline leaves like basal but sessile,
short-decurrent, gradually reduced towards stem apex. Capitula ovoid,
10-15 mm wide (excluding cusps), white arachnoid-hairy. Involucral
bracts about 60—70; outer and middle bracts broad-lanceolate, flat, short-
acuminate, divergent, 10-12 mm long; inner bracts membranous, linear,
very short acuminate, 15-18 mm long, erect. Corolla pink. Receptacular
bristles scabrous. Achenes obovoid, glabrous, smooth, about 7 mm long.
Flowering VII—VIII; fruiting VIII.

Stony slopes in upper mountain zone.—Soviet Central Asia: Pamiro-
Alai (the Alai and Turkestan ranges). Endemic. Described from the
Shauvali Pass. Type in Tashkent.

Series 2. Auriculatae Juz.—Involucral bracts lanceolate, short
acuminate-spinescent, without appendages; receptacular bristles scabrous.
Cauline leaves broadly long-decurrent.

99. C. semidecurrens Winkl. in Tr. Peterb. Bot. Sada, X (1889)
574; XII, 259; XIV, 209; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8,

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220
562.—C. pamirica Bornm. in Bejh. bot. Centralbl. XXXIV, 2 (1916)
182.—C. chionophila Rech. f. and Koie in Dan. Biol. Skr. 8, 2 (1955)
84.—C. auriculata auct. non Boiss.: O. and B. Fedtsch. Perech. Rast.
Turkm. IV (1911) 259.—Ie.: Rech. f. and Koie, op. cit. 85, fig. 62.

Perennial. Root woody, many-headed; caudices elongate, with less
numerous membranous remains of previous year’s leaves. Stems
numerous, erect, branched in upper half, densely leafy, branches one-
headed. Leaves grayish on both sides due to thin, closely appressed
tomentum; basal leaves oblong, pinnatipartite in deltoid lateral lobes,
usually with two acuminate-spinescent lateral lobes at base, acuminate-
spinescent, with thick petioles; cauline leaves like basal but sessile,
broadly long-decurrent on stem, gradually reduced toward stem apex;
uppermost leaves almost pinnatilobate, with acuminate-spinescent lobes
or even spinose-toothed. Capitula cylindrical, about 20 mm wide
(excluding cusps), white-tomentose. Involucral bracts 50-60; outer and
middle bracts coriaceous, erect-spreading, or outer bracts almost
horizontal, 15—20 mm long, lanceolate, gradually attenuate toward apex,
short acuminate-spinescent, carinate dorsally; inner bracts membranous,
narrow-lanceolate or almost linear, scabrous dorsally and along margin,
about 20 mm long. Corolla pink. Receptacular bristles scabrous. Achenes
obovoid, almost smooth, about 5 mm long, with scarcely noticeable
small teeth above. Flowering VII—VIII; fruiting VIII.

Stony slopes in upper mountain zone.—Soviet Central Asia: Pamiro-
Alai (Shugnan, Pamir). General distribution: Afghanistan. Described from
the region of the Karasu River (Kushakevich). Type in Leningrad.

Series 3. Pterolepideae Juz.—tInvolucral bracts somewhat flat, with
membranous or coriaceous appendages; receptacular bristles smooth.
Cauline leaves sessile.

100. C. fedtschenkoana Bornm. in Beih. bot. Centralbl. XXXIV, 2
(1916) 199; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 562.

Perennial. Root thick, woody. Stem erect-spreading, 40-60 cm high,
with less numerous remains of previous year’s leaves at base,
longitudinally sulcate, arachnoid-hairy, usually simple, one-headed, less
often sparsely branched in upper part; branches short, one-headed. Leaves
green and finely arachnoid-hairy above, white-tomentose beneath, oblong;
basal and lower cauline leaves about 17-25 cm long and 3.5 cm wide,
petiolate, pinnatipartite into deltoid, approximate lateral lobes, with one
or two pairs of acuminate-spinescent lateral lobes at base acuminate
into long stiff spine; other leaves like basal but sessile, semiamplexicaul,
gradually reduced toward stem apex. Capitula broadly ovoid, 22-28 mm
wide (excluding cusps), glabrous. Involucral bracts numerous, about 100,

22M

Plate XV.

1—Cousinia dischromata Kult.; 2—C. resinosa Juz.

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/ 222

imbricate, broadened above in coriaceous (membranous in inner bracts)
ciliate-toothed (irregularly incised in inner bracts), broad lanceolate
appendages, usually acuminate-spinescent in short and stiff spine, erect
or (in outer bracts) horizontal, 5-7 mm wide and 12-15 mm long. Corolla
light pink. Receptacular bristles smooth. Achenes obovoid, about 6 mm
long and 4 mm wide, smooth above: Flowering VII-VIII; fruiting VIII.

Gravelly slopes in upper mountain zone.—Soviet Central Asia:
Pamiro-Alai (Gissar Range). Endemic. Described from the Kafaraga Pass,
between the villages of Zimarl and Anzob (Yagnob River). Type in
Leningrad.

101. C. korshinskyi Winkl. in Tr. Peterb. Bot. Sada, XIV (1897)
236.

Perennial. Root thick, woody. Stems with remains of previous year’s
leaves at base, thick, up to 10 mm at base, 100-120 cm high, erect,
striate-sulcate, floccose-arachnoid hairy, glabrescent, sparsely corymbosely
branched in upper half; branches long, erect, almost leafless, one-headed.
Leaves green and glabrous above, white-tomentose beneath, broad-oblong;
basal and lower cauline leaves long-petiolate, about 40-60 cm long and
6 cm wide, pinnatipartite into broad lanceolate, about 3—3.5 cm long
lateral segments, in turn pinnatilobate into short acuminate-spinescent
lobes; cauline leaves like basal but sessile, semiamplexicaul, gradually
reduced toward stem apex, sparse; uppermost leaves pinnatilobate, with
acuminate-spinescent lobes. Capitula almost globose, 16-28 mm wide
(excluding cusps), glabrous; involucral bracts about 80, imbricate,
broadened above in broad-lanceolate, divergent (horizontal in outer bracts)
acuminate-spinescent appendages, about 7 mm wide and 15—17 mm long;
inner bracts with membranous, unequal, brownish appendages, with very
short spine at apex. Corolla pink. Receptacular bristles smooth. Achenes
obovoid, 7-8 mm long and 4-5 mm wide, glabrous, with two or three
scarcely noticeable longitudinal ribs, and two or three indistinct small
teeth above, or smooth. Flowering VII—VIII; fruiting VIII. (Plate XVI,
Fig. 2).

Clayey slopes in the upper part of middle mountain zone.—Soviet
Central Asia: Tien Shan (Fergana Range). Endemic. Described from the
Kugart Pass. Type in Leningrad.

102. C. pterolepida Kult. in Tr. Tadzh. Bazy Akad. Nauk SSSR,
VUlc, VI (1929) 27.—Ie.: Kult. Ibid.

Perennial. Root woody, many-headed; caudices with less numerous
scaly remains of petioles or previous year’s leaves. Stems several, 35—
45 cm high, erect, arachnoid hairy-tomentose, simple, one-headed, less
often sparsely branched; branches erect, one-headed. Leaves coriaceous,

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223

grayish-green, finely arachnoid-hairy above, white-tomentose beneath;
basal and lower cauline leaves about 14-25 cm long and 4—5 cm wide,
petiolate, pinnatipartite in deltoid prickly-lobed lateral lobes; other leaves
like basal but sessile, broadly short-decurrent, gradually reduced toward
stem apex; uppermost leaves more or less prickly-lobed, with toothed
acuminate-spinescent lobes. Capitula cylindrical, about 20 mm wide
(excluding cusps), glabrous. Involucral bracts numerous, closely imbricate,
outermost bracts dorsally white arachnoid-hairy, about 15 mm long,
horizontal, lanceolate, acuminate-spinescent, ciliate-toothed; other bracts
broad-lanceolate, broadened in upper half in membranous, usually
transparent appendage brownish in inner bracts, irregularly toothed,
acuminate into somewhat short stiff spine, and erect appendage. Corolla
pink. Receptacular bristles smooth. Achenes obovoid, glabrous, smooth,
about 8 mm long and 4 mm wide. Flowering VIII; fruiting VIII-IX.

Clayey slopes in subalpine mountain zone.—Soviet Central Asia:
Tien Shan (western). Endemic. Described from the Ugam Range, Kurum-
Dzhul Pass. Type in Tashkent.

Series 4. Macilentae Tschern—Involucral bracts lanceolate, scabrous
dorsally, without appendages; receptacular bristles smooth. Cauline leaves
sessile.

103. C. macilenta Winkl. in Tr. Peterb. Bot. Sada, XIV (1897)
222; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 551.

Perennial. Root woody, many-headed; caudices densely covered with
remains of previous year’s leaves. Stems few, erect, arachnoid hairy-
tomentose, 30—6C cm high, usually with very short branches in upper
half, sometimes branches reduced to 6 mm, always one-headed. Leaves
coriaceous, grayish-green above due to arachnoid-hairy pubescence,
appressed gray-tomentose beneath; basal and lower cauline leaves
petiolate, 2-3 cm wide, 15—20 cm long, irregularly pinnatipartite into
narrow deltoid, prickly-lobed lateral lobes, gradually attenuate into long
spine; cauline leaves like basal but sessile, semiamplexicaul, gradually
reduced upward; uppermost leaves almost spinose-pinnatilobate. Capitula
ovoid, 13-15 mm wide (excluding cusps), finely arachnoid-hairy.
Involucral bracts about 60, all lanceolate, dorsally scabrous, often dark
colored; outer bracts divergent, 8-10 mm long; weakly carinate, attenuate
into short cusp; middle bracts with weakly divergent cusp, 10-16 mm
long, short-acuminate; inner bracts erect, 18 mm long, short-acuminate
into thin spine. Corolla pink. Receptacular bristles smooth. Achenes:
obovoid, glabrous, smooth, about 8 mm long and 4 mm wide. Flowering
VU-VIUI; fruiting VIII.

Gravelly slopes in middle mountain zone.—Soviet Central Asia:

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224

Pamiro-Alai (Gissar Range). Endemic. Described from the vicinity of
the village of Zarchob. Type in Leningrad.

Section 20. Homalochaete Juz—in Tr. Tadzh. Bazy Akad. Nauk
SSSR, 8 (1940) 522, 550.—Homalochaetae Winkl. in Tr. Peterb. Bot.
Sada, XII (1892) 200 p. p.—Capitula solitary terminal on branches,
arachnoid-hairy or tomentose; involucral bracts gradually long acuminate
into almost horizontal attenuate cusp or short-acuminate; corolla pink;
receptacular bristles smooth; achenes glabrous, smooth or weakly angular,
obovoid. Perennial herbs with leaves grayish-green on both sides.

Lectoype of section: C. corymbosa Winkl.

104. C. leptacma Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, 15 (1959) 35.

Perennial. Stem erect, arachnoid-hairy, smooth, 25-30 cm high,
branched above; branches one-headed, short, forming dense, sometimes
corymbose inflorescence. Leaves coriaceous, densely white-tomentose
on both sides; basal leaves petiolate (petioles one-third as long as lamina,
with or without small remote lobes), oblong-elliptical, flat, finely lobed,
lobes broadly deltoid, gradually attenuate in short, rather weak spine;
cauline leaves reduced upward; lower cauline leaves like basal but sessile,
auriculate; middle and upper leaves oblong-ovate, otherwise like basal
leaves. Capitula cylindrical-oblong, 12-15 mm wide (excluding cusps),
arachnoid-hairy. Involucral bracts numerous, deltoid, acicular, carinate;
outer bracts decurved, 5-7 mm long; middle bracts arcuate, up to 15
mm long; subsequent bracts erect, straight, flat, 12-15 mm long; inner
bracts almost membranous, linear, 20 mm long, small toothed, very
long and narrowly acuminate. Receptacular bristles smooth. Achenes
oblong-obovoid, gray, spotted. Flowering VII—VIII; fruiting VIII.

Limestone deposits in middle mountain zone.—Soviet Central Asia:
Pamiro-Alai (Samarkand Mountains). Endemic. Described from the
vicinity of the village of Sarykul. Type in Tashkent.

105. C. litvinovii Kult. ex Juz. in Tr. Tadzh. Bazy Akad. Nauk
SSSR, 8 (1940) 522, 550; Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, 15, 36.—C. amoena auct. non Winkl.: Bornm. in Beih.
bot. Centralbl. XXXIV, 2 (1916) 150.—Exs.: GRF No. 1422.

Perennial. Stem 25-50 cm high, erect, striate-sulcate, appressed,
branched from middle, branches short, one-headed. Leaves coriaceous,
white-tomentose on both sides, with prominent veins, terminating in
stiff spines; basal leaves petiolate, lanceolate or oblanceolate, lobed,
lobes large, deltoid, spinescent, approximate, as a result margin appearing
crisped; lower cauline leaves like basal but sessile, crenate-toothed below,

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225

lobed above; middle leaves oblong-ovate, with acuminate-spinescent
lobes; upper leaves ovate, smaller than middle, uppermost leaves oblong.
Capitula ovoid, 18-20 mm wide (excluding cusps), arachnoid-hairy.
Involucral bracts relatively short, divergent, with strong spine at apex;
outer bracts 5-7 mm long, decurved; middle bracts up to 15 mm long,
divergent, carinate, acuminate into three-angled spine; subesquent bracts
smaller than middle, 10-12 mm long, more flat, with small spine at
apex; inner bracts coriaceous, up to 20 mm long, with small teeth, with
very short spine. Receptacular bristles smooth. Corolla pink. Achenes
oblong-obovoid, dark. Flowering VIJ—VIII; fruiting VIII-IX.

In middle mountain zone.—Soviet Central Asia: Pamiro-Alai
(western spurs of the Zeravshan Range). Endemic. Described from Aman-
Kutan. Type in Leningrad.

106. C. amoena Winkl. in Tr. Peterb. Bot. Sada, XIII (1894) 237;
XIV, 220; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 550; Tschern.
in Bot. Mat. Gerb. Inst. Bot. Akad. Nauk UzbSSR, 15, 36.

Perennial. Stem 25-60 cm high, erect, appressed-arachnoid-hairy,
striate-sulcate, usually branched from middle, branches short, one-headed.
Leaves coriaceous, white-tomentose on both sides, but tomentum thinner
above, with prominent veins, terminating in stiff spines, large toothed
basal leaves short-petiolate, oblanceolate, lobed, lobes sharply incised;
lower cauline leaves like basal but sessile, semiamplexicaul; middle
leaves smaller than lower, oblong-ovate, sessile, semiamplexicaul, large-
toothed; upper leaves ovate, sessile, semiamplexicaul, acuminate-
spinescent; uppermost leaves reduced. Capitula ovoid-globose, 18—25
mm wide (excluding cusps), weakly arachnoid-hairy. Involucral bracts
squarrose, stiff, carinate, terminating in a three-angled spine; outer bracts
up to 10 mm long, somewhat decurved; middle bracts up to 20 mm
long, squarrose, three-angled acuminate; subsequent bracts somewhat
flat, up to 20 mm long, up to 3 mm wide at base, short spinescent;
inner bracts coriaceous, linear, up to 2 mm wide and 25 mm long, with
small teeth terminating in very small spine. Receptacular bristles smooth.
Corolla pink. Achenes obovoid, light gray. Flowering VI-VII; fruiting
VU-VII.

Stony screes in middle mountain zone.—Soviet Central Asia: Pamiro-
Alai (Zeravshan Range). Endemic. Described from the Sangydzhuman
Pass. Type in Leningrad.

107. C. campyloraphis Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, 15 (1950) 37.

Perennial. Stem erect, striate-sulcate, arachnoid-hairy-tomentose, 30—
50 cm high, branched from middle; branches more or less short, one-

229

230

226

headed, forming lax, corymbose-inflorescence. Leaves coriaceous,
grayish-tomentose on both sides, more sparsely above; basal leaves
petiolate (petiole half as long as lamina, with or without small, remote,
prickly lobes), oblong-elliptical, crisped and almost lobed-prickly-toothed,
lobes deltoid, sharply incised, approximate; cauline leaves reduced
upward; lower leaves like basal but with smaller petioles; middle leaves
sessile, auriculate, with up to 1 cm long spines; upper and uppermost
leaves reduced. Capitula broadly ovoid or globose, 15—17(20) mm wide
(excluding cusps), arachnoid-hairy. Involucral bracts numerous, rather
weak, dorsally carinate; outer bracts divergent, up to 10 mm long; middle
bracts falcate, up to 25 mm long, subsequent bracts almost erect, more
or less flat, up to 23 mm long; inner bracts almost membranous, linear,
25(30) mm long, finely toothed, acuminate, very sharp. Receptacular
bristles smooth. Corolla pink. Achenes oblong-obovoid, gray, smooth, 6
mm long and 2.5 mm wide. Flowering VII—VIII; fruiting VIII.

Middle mountain zone.—Soviet Central Asia: Pamiro-Alai. Endemic.
Described from the mountains south of Yakkabag. Type in Tashkent.

108. C. proxima Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3
(1936) 301; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 550; Tschern.
in Bot. Mat. Gerb. Inst. Bot. Akad. Nauk UzbSSR, 15, 38.

Perennial. Stem 30-80 cm high, erect, floccose-arachnoid hairy,
striate-sulcate, branched from base; branches long, one-headed. Leaves
coriaceous, finely arachnoid-hairy, grayish-green above, finely gray-
tomentose beneath, with prominent veins, large-prickly dentate or lobed;
basal leaves petiolate, oblong-lanceolate or lanceolate, large-toothed or
lobed (only one-third lamina incised), lobes spinescent, remote; lower
cauline leaves like basal, petiolate or sessile; middle leaves oblong-
ovate, sessile, auriculate; upper leaves smaller than middle; uppermost
leaves reduced. Capitula globose. 12-20 mm wide (excluding cusps),
densely arachnoid-hairy. Involucral bracts comparatively small, finely
short-spinescent, somewhat flat; outer bracts 3-6 mm long, decurved;
middle bracts up to 10 mm long, divergent, weakly carinate dorsally;
subsequent bracts somewhat flat, finely spinescent, not longer than middle
bracts; inner bracts membranous, linear-lanceolate, up to 2 mm wide,
acuminate, somewhat finely serrate. Receptacular bristles smooth. Corolla
light pink. Achenes obovoid, gray. Flowering VI-VII; fruiting VIJ-VIII.

Middle mountain zone on rocks and stony-gravelly mountain
slopes.—Soviet Central Asia: Pamiro-Alai (Gissar Range). Endemic.
Described from rocks along the bank of the Sanga-Osiiya River. Type
in Leningrad.

109. C. subcandicans Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, 15 (1959) 41.

227

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Plate XVI.

1—Cousinia podophylla Tschern.; 2—C. korshinskyi Winkl.

231

232

228

Perennial. Stems erect, arachnoid hairy-tomentose, striate-sulcate,
40-100 cm high, branched above; branches long, one-headed, forming
lax inflorescence. Leaves coriaceous, gray-tomentose on both sides; basal
leaves very short-petiolate, oblanceolate or oblong-oblanceolate, less often
elliptical, weakly crisped-lobed, lobes deltoid, attenuate into short cusp,
prickly-lobed; lowermost leaves reduced, distant; cauline leaves reduced
upward; lower leaves like basal but sessile, auriculate; middle leaves
oblong-ovate or ovate; upper leaves broad-ovate, with longer (up to 1
cm) more stiff spines; leaves on branches strongly reduced, closely
placed. Capitula broadly ovoid, arachnoid-hairy, 12-15 mm wide
(excluding cusps). Involucral bracts numerous, three-angled-subulate, stiff;
outer bracts weak, decurved, 5—7 mm long; middle bracts stiff, horizontal,
up to 15 mm long; subsequent bracts somewhat flat, erect, shorter; inner
bracts almost membranous, linear, 20-22 mm long, finely serrate,
acuminate, very sharp. Receptacular bristles smooth. Corolla pink;
achenes oblong-obovoid, dark gray, spotted. Flowering VII—-VIII; fruiting
VU.

Rocks, screes, and clayey slopes in middle mountain zone.—Soviet
Central Asia: Pamiro-Alai (upper reaches of the Tankhas River).
Endemic. Described from the Kashka-Darya River Basin, above the
village of Chopukh, along the road to Tamshush. Type in Tashkent.

110. C. candicans Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1,
3 (1936) 300; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 550;
Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad. Nauk UzbSSR, 15, 42.

Perennial. Stems 25-70 cm high, erect, appressed-arachnoid-hairy,
almost smooth, branched from base, branches slender, long, one-headed.
Leaves coriaceous, gray-tomentose on both sides, with prominent veins
terminating in stiff spines, large-toothed; basal leaves short-petiolate,
almost sessile, oblong or oblong-lanceolate, large-toothed or lobed, lobes
sharply incised, with stiff spine; lower cauline leaves like basal; middle
and upper leaves broad-ovate or almost orbicular, sessile, auriculate,
spinescent; uppermost leaves closely placed, reduced. Capitula ovoid,
17-20 mm wide (excluding cusps), densely arachnoid-hairy. Involucral
bracts squarrose, stiff, three-angled, spinescent; outer bracts up to 10
mm long, decurved; middle bracts somewhat falcate, divergent, 25 mm
long, with stiff 3-angled spine; subsequent bracts somewhat flat, up to
15 mm long, gradually acuminate into short stiff spine; inner bracts
coriaceous, linear, up to 2 mm wide and 25 mm long, finely toothed,
with very short spine. Receptacular bristles smooth. Corolla purple;
achenes oblong-obovoid, spotted, almost three-angled. Flowering VI-—
VII; fruiting VII—-VIII.

Limestone slopes in middle mountain zone.—Soviet Central Asia:

233

229

Pamiro-Alai (Babatag Range). Endemic. Described from the central part
of Babatag, from Bish-Archa Peak. Type in Leningrad.

111. C. podophylla Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, 15 (1959) 43.

Perennial. Stem erect, arachnoid hairy-tomentose, striate-sulcate, 30—
40 cm high, branched in upper part, branches long, one-headed, very
densely and leafy, forming lax inflorescence. Leaves coriaceous, gray-
tomentose on both sides; basal leaves long-petiolate (petioles two-thirds,
less often half, as long as lamina, with less numerous small prickly
lobes), obovate or broad-elliptical, crisped to large-toothed, teeth broadly
deltoid, abruptly attenuate in rather short spine and with lateral spines;
cauline leaves reduced toward stem apex; lower leaves like basal but
more short-petiolate or almost sessile; middle and upper leaves oblong-
ovate or ovate, short-decurrent. Capitula ovoid, arachnoid-hairy, 12—17
mm wide (excluding cusps). Involucral bracts numerous; outer bracts
decurved or almost horizontal, 5 mm long, middle bracts stiff, horizontal
or more or less horizontal, carinate, about 15 mm long; subsequent bracts
erect, smaller; inner bracts almost membranous, linear, 18—20 mm long,
very finely serrate, attenuate, acute. Corolla purple. Receptacular bristles
smooth. Achenes oblong-obovoid, almost three-angled, gray with dark
spots, 6 mm long and 2 mm wide. Flowering VII; fruiting VIII. (Plate
XVI, Fig. 1).

Gypsic and limestone slopes in middle mountain zone.—Soviet
Central Asia: Pamiro-Alai (Babatag, Gazimailik, Aktau and Gardani-
Ushti). Endemic. Described from the Babatag Mountains (vicinity of
the village of Dzhidda-Bulak. Type in Tashkent.

112. C. leptocladoides Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, 15 (1959) 45.

Perennial. Stem erect, slender, arachnoid hairy-tomentose, striate-
sulcate, 25-50 cm high, branched from base; branches slender, long,
one-headed, forming lax inflorescence. Leaves coriaceous on both sides,
gray arachnoid hairy-tomentose, more sparsely and finely above; basal
leaves short-petiolate (petioles one-sixth as long as lamina, with very
narrow lobes, almost reduced to spines), crisped, oblong-oblanceolate,
pinnatipartite, with lateral deltoid lobes from broadly deltoid prickly-
deltoid lobed base, attenuate into long and stiff spine; cauline leaves
reduced above; lower cauline leaves sessile or subsessile, otherwise like
basal; middle leaves elliptical-oblong, sessile, auriculate, with smaller
spines; upper leaves ovate, sessile, auriculate, deltoid-lobed, lobes
terminating into spine, terminal lobe stiff; leaves on branches strongly
reduced, approximate, sessile, mainly in middle. Capitula oblong-ovoid,

234

230

10-12 mm wide (excluding cusps), arachnoid-hairy. Involucral bracts
relatively less numerous, carinate; outer bracts decurved, weak, short,
up to 7 mm long; middle bracts horizontal, up to 20 mm long, three-
angled-subulate, terminating in long spine; subsequent bracts flat, erect,
12-15 mm long, short-spinescent; inner bracts almost membranous,
very narrow-linear, finely toothed, long attenuate, 15-17 mm long.
Receptacular bristles smooth; corolla pink. Achenes oblong-obovoid,
almost three-angled, light gray, spotted. Flowering VI-VIII; fruiting VIII-
IX.

Gypsic slopes in middle mountain zone——Soviet Central Asia:
Pamiro-Alai (southwest). Endemic. Described from the upper reaches of
the Katta-Uru River. Type in Tashkent.

Note. The material for this species is very heterogeneous; therefore,
additional collections are necessary to determine the precise boundary
of its range.

113. C. gnezdilloi Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, 15 (1959) 47.

Perennial. Stem erect, white arachnoid hairy-tomentose, glabrescent,
striate-sulcate, 30-60 cm high, weakly branched above; branches long,
one-headed, often sparsely leafy, forming lax inflorescence. Leaves
coriaceous, very finely gray arachnoid hairy-tomentose on both sides;
basal leaves short-petiolate (petioles one-fourth to one-third as long as
lamina, smooth or almost smooth), oblong-obovate or obovate-elliptical,
crisped, prickly-lobed, lobes deltoid, gradually attenuate-spinescent,
repeatedly prickly deltoid-lobed; cauline leaves rather abruptly reduced
above; lower cauline leaves short-petiolate or sessile, otherwise like basal;
middle and upper leaves distant, ovate or broad-ovate, sessile, auriculate,
large prickly-toothed: Capitula ovoid, 12-15 mm wide (excluding cusps),
finely arachnoid-hairy. Involucral bracts relatively less numerous, deltoid-
subulate, carinate, stiff; quter bracts decurved, up to 10 mm long; middle
bracts horizontal, almost two times as long as involucral diameter, 25—
30 mm long; subsequent bracts flat, erect or slightly recurved, with
very short spine, 15—17 mm long; inner bracts almost membranous, linear,
finely toothed, very long attenuate, 20-22 mm long. Receptacular bristles
smooth. Corolla pink; achenes oblong-obovoid, light colored. Flowering
VI-VII; fruiting VII—VIII.

Gypsic slopes.—Soviet Central Asia: Pamiro-Alai (southwest).
Endemic. Described from the vicinity of Lake Kurgantash. Type in
Tashkent.

114. C. leptoclada Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIlIc, VI
(1929) 15; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 550; Tschern.

235

231

in Bot. Mat. Gerb. Inst. Bot. Akad. Nauk UzbSSR, 15, 49; Kult. in FI.
Turkm. VII, 206.

Perennial. Stem 30-60 cm high, slender, white-tomentose, striate-
sulcate, moderately branched above; branches long, slender, usually one-
headed. Leaves gray-tomentose on both sides, but tomentum thinner
above, coriaceous, strongly spinose; basal leaves short-petiolate (petioles
one-fifth as long as lamina), narrow oblong-lanceolate, about 45 mm
wide (with spines), pinnatisect, strongly acuminate into long stiff spine,
lateral segments numerous and strongly spinose; lower cauline leaves
like basal but sessile; middle leaves oblong, pinnatisect, lateral segments
narrow, uniformly spaced, spinescent; uppermost leaves sessile, densely
spinose, small, remote on floriferous branches. Capitula oblong-ovoid,
10-12 mm wide (excluding cusps), arachnoid-hairy. Involucral bracts
fewer, green, subulate, dorsally carinate, squarrose; outer bracts decurved,
up to 7 mm long; middle bracts horizontal, two or three times exceeding
capitulum diameter (20-30 mm long), subulately acuminate in thin, long,
three-angled spine; subsequent bracts somewhat flat, 12-15 mm long,
erect, with short spine at apex; inner bracts membranous, narrow-linear,
1-2 mm wide, finely serrate. Receptacular bristles smooth. Corolla purple.
Achenes obovoid, light colored. Flowering VI—VII; fruiting VII-VIU.

Stony slopes in the upper part of middle mountain zone.—Soviet
Central Asia: Pamiro-Alai (Kugitang Range). Endemic. Described from
the Kugitang Range, near Kyzylalma. Type in Tashkent.

115. C. hoplopylla Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, 15 (1959) 49.

Perennial. Stem erect, strong, 50-80 cm high, arachnoid hairy-
tomentose, sulcate, branched from middle; branches long, one-headed,
forming lax inflorescence. Leaves coriaceous, green on both sides, gray-
tomentose, basal leaves small, reduced; cauline leaves reduced above,
lower leaves very short-petiolate or sessile, oblanceolate, crisped,
pinnatipartite, lateral segments triangular, remote and spiny-triangular,
attenuate into strong, long spine; middle leaves sessile, auriculate, with
weakly prickly lobes, otherwise like lower leaves; upper leaves ovate,
auriculate, long and hard-prickly lobed, terminal lobe with more stiff
spine; leaves on branches strongly reduced, spinescent, less numerous.
Capitula globose, arachnoid-hairy, 18-20 mm wide (excluding cusps).
Involucral bracts numerous, triangular, all bracts erect-spreading; outer
more or less decurved, about 7 mm long; middle bracts 20-22 mm
long; subsequent bracts shorter and more flat. Receptacular bristles
smooth. Flowering VII.

Stony slopes in upper mountain zone.—Soviet Central Asia: Pamiro-
Alai (Chulbair Mountains). Endemic. Described from Khodzha-Barku.
Type in Tashkent.

Ie 92

116. C. corymbosa Winkl. in Tr. Peterb. Bot. Sada, X (1887) 89;
Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 550.

Perennial. Stem erect, floccose-arachnoid hairy, 80-100 cm high,
cymosely branched in upper half; branches leafy, long, one-headed.
Leaves coriaceous, finely arachnoid-hairy on both sides, more densely
beneath; basal leaves petiolate, broad elliptical, prickly-lobed, 10—12
cm wide (with spines), with broad deltoid, acuminate-spinescent lobes;
cauline leaves gradually reduced, sessile, semiamplexicaul, narrow
deltoid-lobed, with acuminate-strong spinescent lobes, oblong-ovate;
leaves on branches reduced, more finely prickly-lobed, more or less
approximate. Capitula globose, 17-20 mm wide (excluding cusps),
arachnoid-hairy. Involucral bracts numerous, horizontal, gradually long-
acuminate into long subulate cusp; outer bracts weakly decurved, 12-15
mm long; middle bracts horizontal, 20-25 mm long; inner bracts narrow-
lanceolate, dorsally scabrous, thin and short acuminate, 20 mm long.
Receptacular bristles smooth. Corolla light pink. Achenes obovoid,
glabrous, smooth, 5 mm long and 2.5—3 mm wide. Flowering VII; fruiting
VITI-IX.

Gravelly slopes in middle mountain zone.—Soviet Central Asia:
Pamiro-Alai (south). Endemic. Described from the Gandadarra Ravine.
Type in Leningrad.

117. C. coerulea Kult. ex Tschern. in Bot. Mat. Gerb. Inst. Bot.
Akad. Nauk UzbSSR, 15 (1959) 51.—C. subinermis Tschern. Ibid. (1959)
40.

Perennial. Stem, erect, floccose-arachnoid hairy-tomentose, up to
100 cm high, branched in upper half; branches long, secondary branches
rather short, one-headed, forming lax inflorescence. Leaves coriaceous,
gray arachnoid-hairy to tomentose on both sides; basal leaves short-

236 petiolate, with thick floccose-tomentose petioles bearing few narrow
spinose lobes, oblong or oblong-oblanceolate, prickly-lobed, with broadly
deltoid, spinose-toothed, acuminate-cuspidate, rather approximate lobes,
as a result leaf somewhat crisped; cauline leaves gradually reduced above;
lower cauline leaves lanceolate, sessile, with auricles connate with stem,
narrow deltoid-prickly-lobed, crisped; lobes deltoid, abruptly attenuate
in long and stiff spine, unequally and broadly spinose-toothed; middle
leaves oblong, otherwise like lower; upper leaves ovate, sessile,
semiamplexicaul, with connate auricles, spinose-toothed; leaves on
branches reduced, remote. Capitula globose-ovoid, 14-20 mm wide
(excluding cusps), densely arachnoid hairy-tomentose. Involucral bracts
numerous, narrow-lanceolate, acuminate into fine acicular cusp,
squarrose; outer bracts slightly decurved, up to 5 mm long; middle bracts
erect-spreading, 12-16 mm long; inner bracts erect, narrow-lanceolate.

237

233

gradually acuminate in short cusp, 22 mm long. Receptacular bristles
smooth. Corolla whitish or light pink. Achenes obovoid, glabrous,
smooth, weakly ribbed, 6 mm long and 3 mm wide. Flowering VII-—
VII; fruiting VIII.

Gravelly stony slopes in middle mountain zone.—Soviet Central
Asia: Pamiro-Alai (Gissar Range). Endemic. Described from the Varzob
River Basin, upstream along the Ziddy River from the confluence of
Ziddy and Mai-Kur. Type in Tashkent.

Note. Study of additional material of this species leads to the
conclusion that C. coerulea is more widely distributed in the Gissar
Range and C. subinermis is a synonym of it. Now we can give a more
detailed and expanded description of the species, because many of the
features were not characterized in the original description, given that
the specimens in our hands were very young.

Section 21. Lopholepis Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962).—Capitula solitary terminal on branches or
main stem; involucral bracts coriaceous, lanceolate, cristate-prickly along
margin, short-acuminate; receptacular bristles slightly scabrous; corolla
whitish or pale yellow; anther tube glabrous, purple; achenes obovate,
glabrous, smooth, roundish above.

Perennial plants with thick, woody, many-headed roots and sessile
semiamplexicaul leaves.

Type of section: C. rigida Kult.

118. C. rigida Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIlIc, VI
(1929) 26.—Ie.: Kult. Ibid., Fig. 26.

Perennial. Root woody, blackish-fibrous, many-headed; root collar
white-hairy, with remains of old leaves. Stem erect, 35 cm high, weakly
sulcate, finely arachnoid-hairy below, glandular throughout, densely
above, simple, one-headed. Leaves coriaceous, green on both sides, with
network of prominent veins, glandular; basal leaves long-petiolate, 20
cm long, with petioles somewhat thickened at base, lanceolate, attenuate
toward both ends, entire, with remote, thin and short spines along margin,
more approximate toward leaf base and continued on petiole; lower
cauline leaves like basal but with shorter petioles; other cauline leaves
sessile, semiamplexicaul, with thin spines at base. Capitula globose-
cylindrical, truncate basally, 15-20 mm wide, 25—28 mm long (excluding
corollas), and with very thin arachnoid-hairy pubescence. Involucral
bracts densely glandular, all except inner ovate to oblong-ovate, 12—22
mm long and 6—7 mm wide at base, midrib somewhat prominent, more
or less flat, cristate-prickly along entire margin, with numerous, thin,
yellow spines, acuminate into erect cusp; inner bracts membranous,

23

oo

234

narrow lanceolate, scabrous along margin, acuminate into thin cusp,
dorsally brownish. Receptacular bristles smooth or few with one or two
serrations. Corolla pale yellow; anther tube purple. Achenes obovoid,
smooth, roundish above, 6 mm long, gray. Flowering VII—VIII. (Plate
XVUI, Fig. 1).

Stony slopes.——Soviet Central Asia: Tien Shan (northern slope of
the Alai Alatau and Kirgiz Alatau). Endemic. Described from the Kirgiz
Alatau, Kosh-Archa-Sai. Type in Tashkent.

119. C. mindshelkensis B. Fedtsch. Perech. Rast. Turk. IV (1911)
261; B. Fedtsch. in Fedde, Repert, X, 164.

Perennial. Root thick, blackish-fibrous, with branched collar, covered
with numerous remains of old leaves, with numerous sterile leaf rosettes
and stems, forming thick pads. Stems 10-30 cm high, usually simple,
one-headed, less often weakly branched above and then with two or
three capitula on branch tips, white arachnoid-hairy. Leaves coriaceous,
finely gray-tomentose on both sides, more densely beneath, glandular
above with sessile glandular hairs; basal leaves attenuate toward both
ends, short-petiolate, narrow-lanceolate or lanceolate, sparsely crenate-
prickly-toothed along margin; cauline leaves strongly reduced, sessile,
semiamplexicaul, like basal. Capitula globose-ovoid, 12-15 mm wide
(excluding cusps) and 18 mm long (excluding corollas), almost glabrous
or slightly arachnoid-hairy above; involucral bracts except inner,
coriaceous, lanceolate, dorsally glandular, acuminate, erect, cristate-
prickly; inner and innermost bracts membranous, acuminate-spinescent,
irregularly toothed along margin. Receptacular bristles unequal, longest
bristles slightly scabrous above. Corolla whitish or light pink; anther
tube purple. Achenes oblong-obovoid, glabrous, smooth, roundish above,
gray, 6 mm long, 2.5 mm wide. Flowering VII—VIII; fruiting IX.

Stony gravelly slopes.—Soviet Central Asia: Tien Shan (Karatau).
Endemic. Described from Bish-Saz area. Type in Leningrad.

120. C. gomolitzkii Juz. ex Tschern. in Bot. Mat. Gerb. Inst. Bot.
Akad. Nauk UzbSSR, XVII (1962) 95.

Perennial. Root thick, woody, covered with black fibrous bark, many-
headed, with thick collar densely covered with remains of old petioles,
stems 25-50 cm high, numerous, straw-yellow to light brown, finely
striate-sulcate, glandular with sessile glandular hairs and papillae scattered
throughout. Basal leaves oblong, narrowed toward both ends, on short
and thick petioles, margins boldly sinuate-toothed, teeth acuminate-
spinescent, hard coriaceous, light green on both sides, glossy, with
strongly prominent network of veins, densely glandular-hairy with short
sessile hairs, sometimes finely white-tomentose beneath along margin
or throughout (f. hypoleuca); cauline leaves gradually reduced, sessile,

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235

semiamplexicaul, dense, otherwise like basal leaves. Capitula oblong-
cylindrical, 22 mm long (excluding corollas) and 12-15 mm wide
(excluding cusps), usually finely white- or ferruginous arachnoid-hairy
above. Involucral bracts very densely glandular dorsally, with sessile
glandular hairs, all except inner bracts narrowed from ovate base to
narrow-lanceolate, acuminate, erect with cristately arranged thin spines
along margin; inner and innermost bracts broad-linear, purple dorsally,
almost membranous, scabrous, irregularly toothed along margin,
acuminate-spinescent. Receptacular bristles unequal, longest slightly
flattened above and weakly scabrous. Corolla white, anther tube purple.
Achenes immature, oblong-obovoid, glabrous, smooth, roundish above,
7 mm long. Flowering VII-VIII.

Stony and rocky slopes.—Soviet Central Asia: Tien Shan (Karatau).
Endemic. Described from the Obaly-Dzhan Mountain. Type in Tashkent.

Section 22. Pectinatae Winkl. in Tr. Peterb. Bot. Sada, XII (1892)
263 p. p—Congestae Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2 (1865)
11 p.p.—Uncinatae Winkl. op. cit. 186 p. p.—Capitula in terminal
corymbose or lax panicles, usually aggregated at branch apices; involucral
bracts somewhat flat, cristate-toothed along margin, acuminate in straight
or uncinate cusp; receptacular bristles smooth; corolla yellow, with pale-
purple, sparsely hairy anther tube; achenes obovoid, smooth, roundish
above. Perennial monocarpic plants with large coriaceous basal leaves
and smaller stems.

Lectotype of section: C. albertii Rgl. and Schmalh.

121. C. albertii Rgl. and Schm. in Tr. Peterb. Bot. Sada, VI (1879)
315; Winkl. ibid XII, 264.

Perennial. Stem erect, strong, floccose-arachnoid-hairy, striate-
sulcate, in upper half profusely and dense cymosely branched, branches
with capitula on short peduncles aggregated at tips, forming compact
corymbose inflorescence. Leaves coriaceous, with network of prominent
veins, green, sparsely arachnoid-hairy, glabrescent, densely white-
tomentose beneath; basal and lower cauline leaves with thick petioles,
slightly broadened at base, cordate-ovate or ovate, about 20 cm long
excluding petioles, 15 cm wide, weakly crisped with two prickly
teeth; middle and upper cauline leaves gradually reduced, sessile,
semiamplexicaul, ovate-lanceolate, crisped, with two large prickly teeth,
spines at leaf base stiff, strong, covering petiole. Capitula oblong-ovoid,
with 12-15 florets, 17-19 mm long and 8-9 mm wide (excluding cusps),
glabrous. Involucral bracts except inner ovate-lanceolate to oblong-
lanceolate, appressed, cristate-spinulose on both sides in upper half, with
two to six pairs of thin spines, acuminate in weakly incurved cusp;
inner bracts linear-oblong, with somewhat membranous, usually

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236

irregularly incised appendages, roundish above, with stiff and weakly
curved spine. Receptacular bristles smooth. Corolla yellow, conspicuous,
anther tube pale purple, sparsely sericeous. Achenes obovoid, 6 mm
long, roundish above. Flowering V; fruiting VI.

Dry loamy-gravelly slopes.—Soviet Central Asia: Tien Shan
(Karatau). Endemic. Described from Boroldai. Type in Leningrad.

Note. The hybrid between C. albertii Rgl. and Schmalh. and C.
umbrosa Bge. is discussed in the note to the latter species.

122. C. pentacantha Rgl. and Schm. in Tr. Peterb. Bot. Sada, VI
(1879) 315; Winkl. ibid., XII, 189.—Arctium pentacanthum O. Ktze.
Rev. Gen. 1 (1891) 308.

Perennial. Plants mostly with one stem. Stem erect, striate-sulcate,
arachnoid-hairy, 30-60 cm high, profusely branched from base; branches
relatively short, densely leafy, in turn branched into numerous secondary
branches, bearing one to three or four capitula on each (1-2 cm long)
peduncle, inflorescence lax, paniculate. Leaves coriaceous, basal leaves
large, 25-30 cm long and 17-19 cm wide, ovate or oblong-ovate,
attenuate toward both ends, finely arachnoid-hairy above, with isolated
glandular hairs, gray-tomentose beneath, petiolate; petioles floccose-
tomentose, flat adaxially slightly convex abaxially, sometimes with
occasional narrow lobes; cauline leaves much smaller than basal,
gradually reduced toward stem apex; middle cauline leaves sessile,
elliptical or almost orbicular, narrowed toward base, finely toothed,
glabrous above and gray-tomentose beneath. Capitula 15 mm long and
6 mm wide (excluding cusps), oblong-ovoid, glabrous. Involucral bracts
numerous, usually 60, with (up to 3 mm long), uncinate apical spines;
outer bracts narrow lanceolate, 3 mm long, with one or two pairs of
very thin spines above along margin; middle bracts narrow lanceolate,
8-10 mm long, closely appressed at base, abruptly horizontal in upper
part, with two or three pairs of thin, up to 2 mm long, spines along
margin; inner bracts erect, lanceolate, 12 mm long, somewhat broadened
at apex, with membranous border and up to 3 mm long, thin, uncinate
spines. Receptacular bristles smooth. Corolla yellow; anther tube pale
purple, sparsely pubescent. Achenes obovoid, gray. Flowering VI-VII;
fruiting VII-VIII. (Plate XVII, Fig. 2).

Stony slopes in foothills ——Soviet Central Asia: Tien Shan (Mogoltau,
foothills of the Chatkal Range, adjoining the Fergana Valley). Endemic.
Described from the Maili River Basin. Type in Leningrad.

123. C. horrescens Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1,
3 (1936) 322.—C. albertii ssp. horridula Bornm. in Fedde, Repert.
XXXVI (1934) 337.—Exs.: H.F.A.M. No. 478 (sub C. albertii Rgl. and
Schmalh.).

241

237.

Perennial. Plants with one stem. Stem erect, deeply straite-sulcate,
finely arachnoid-hairy, almost glabrous, 50-60 cm high, branched from
middle; branches, in turn, repeatedly branched in relatively short one-
headed secondary branches; inflorescence dense, broadly corymbose.
Leaves coriaceous, basal leaves large, cauline leaves abruptly reduced
towards stem. apex; basal leaves petiolate, ovate or oblong-ovate, with
lamina up to 25 cm long and 15-16 cm wide, toothed, with broadly
deltoid, somewhat approximate teeth, in turn, irregularly prickly-toothed,
with prominent veins, green above, very finely and sprasely arachnoid-
hairy with glandular hairs scattered throughout, white-tomentose beneath;
upper leaves oblong, sessile, with two or three pairs of stiff basal spines,
spinose-toothed, glabrous above, loosely tomentose beneath, with
prominent midrib, terminating in stiff, 7-10 mm long, three-angled spine.
Capitula ovoid, 10 mm wide (excluding cusps) and 22 mm long, glabrous,
somewhat foveate at base. Involucral bracts numerous, more or less
carinate, with scattered fine glandular hairs; outer bracts 7 mm long,
divergent, lanceolate, acuminate into short spine; with two to four pairs
of thin stiff spines along margin; middle bracts 10 mm long, lanceolate,
basally appressed, cartilaginous-serrated, divergent in upper half, with
three to five pairs of thin stiff hairs along margin; inner bracts oblong,
12 mm long, slightly broadened above, with wide membranous border,
and short apical spine. Receptacular bristles smooth. Corolla greenish-
yellow; anther tube pale purple, sparsely pubescent in upper part. Achenes
obovoid, spotted, roundish above, weakly curved, about 5—6 mm long
and 2.5—3 mm wide. Flowering V—VI; fruiting VI-VII.

Stony slopes from foothills to middle mountain zone—Soviet Central
Asia: Tien Shan (west). Endemic. Described from the Bos-Su River
Basin, vicinity of Niazbek. Type in Tashkent.

Note. Specimens with some resemblance to C. albertii Rgl. and
Schmalh. are found in the foothills of Karatau, and possibly, are hybrids
of these two species, which come in contact in the Karatau Mountains.

124. C. pentacanthoides Juz. ex Tschern. in Bot. Mat. Gerb. Inst.
Bot. Akad. Nauk UzbSSR, XVII (1962) 102.

Perennial. Plants with one stem. Stem erect, striate-sulcate, very
finely arachnoid-hairy, almost smooth, 70-80 cm high, branched from
base, branches long, repeatedly branched in relatively short branches
with several capitula on 5-6 mm long peduncles; inflorescence lax,
paniculate. Leaves coriaceous; basal leaves petiolate with short (6—7.5
cm long) petioles thickened at base and with isolated spines, lamina
oblong-elliptical, 20-30 cm long and 10-15 cm wide, with prominent
veins, with wide lobes, lobes finely toothed, green, densely or less
often sparsely glandular, and very finely arachnoid-hairy above, white-

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238

tomentose beneath; cauline leaves much smaller than basal, abruptly
reduced toward stem apex; lower cauline leaves sessile, elliptical,
narrowed toward both ends, acuminate into stiff, short spine, with three
or four pairs of basal spines, almost glabrous and glandular above, gray-
tomentose beneath; middle and upper leaves strongly reduced, oblong,
sessile, with one or two pairs of basal spines. Capitula ovoid, 20 mm
long and 7 mm wide (excluding cusps), glabrous. Involucral bracts 40—
50; outer bracts 4-5 mm long, somewhat carinate, broad-lanceolate, with
two or three pairs of thin spines along margin; middle bracts 7-9 mm
long, closely appressed at base, entire, weakly divergent in upper half,
acuminate into stiff, up to 3 mm long, thin, weakly curved spine, with
three or four pairs of thin, up to 3 mm long spines along margin; inner
bracts oblong, acuminate-deltoid, with membranous border, and up to 2
mm long slightly curved spine. Receptacular bristles smooth. Corolla
yellow; anther tube pale purple, sparsely pubescent. Achenes narrow
oblong, 7 mm long and 1.5—2 mm wide, roundish above, dorsally slightly
curved, spotted. Flowering VI—VII; fruiting VII-—VIII.

Stony slopes in lower mountain zone.—Soviet Central Asia: Tien
Shan (west). Endemic. Described from the Kuraminskii Range, the
mountain between Kurashi-Kuduk and the Kainar River. Type in
Tashkent.

Note. Plants were collected from the region of the village of Kansai
with characters intermediate between C. pentacantha Rgl. and Schmalh.
(known from this region on the basis of collections from the Ungar-
Tyube Mountain) and C. pentacanthoides Juz.; however, the strongly
glandular surface of their leaves suggests that it would be more
appropriate to include these plants under C. pentacanthoides Juz.

Specimens with less glandular leaves and a shorter, almost straight
cusp of the involucral bracts were collected from the region of the village
of Zarkent; on the basis of these herbarium specimens, S.V. Juzepczuk
named the new species C. intersita Juz. The paucity of material from
this region and the attendant impossibility of checking the constancy of
these characters in a large collection compels us at present to interpret
the species C. pentacanthoides more broadly, leaving the solution to the
problem of segregating the new species until further observations in
nature can be made.

Section 23. Oligantha Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR,
8 (1940) 15, 565.—Congestae Bge. in Mém. Akad. Pétersb. 7 sér. 9, 2
(1865) 11 p. min. p—Capitula oblong-cylindrical, aggregated on branch
tips, readily falling, small; involucral bracts herbaceous, flat, closely
appressed, with slightly divergent short cusp, cristate-toothed; receptacular
bristles smooth; corolla whitish; achenes glabrous, smooth, roundish

243

Plate XVII.
1—Cousinia triflora Schrenk.; 2—C. pentacantha Rgl. an

d Schm.

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240

above. Perennial herb with large, sparsely glandular and coriaceous
leaves.
Type of section: C. triflora Schrenk.

125. C. triflora Schrenk in Bull. phys.-math. Akad. Pétersb. 3 (1845)
108; Bge. in Mém. Akad. Pétersb. 7 sér. 9, 2, 11; Winkl. in Tr. Peterb.
Bot. Sada, XII, 264; XIV, 211; Bornm. in Beih. bot. Centralbl. XXXIV,
2, 194; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 565.—Arctium
triflorum O. Ktze. Rev. Gen. (1891) 308.—Exs.: H.F.A.M. No. 197.

Perennial. Stem solitary, 60-120 cm high, erect, thick, striate-sulcate,
glabrous, densely leafy, branched in upper half; branches short, leafy,

~in turn branched into branchlets bearing groups of 9-15 capitula at apices;

general inflorescence corymbose-paniculate. Leaves later coriaceous;
basal leaves petiolate, petioles thick and broadened at base, oblong-
ovate, attenuate toward both ends, up to 40 cm long and 25 cm wide,
sparsely glandular, with reticulate venation above, finely arachnoid-hairy
beneath with wide crenate-toothed lobes; upper cauline leaves oblong-
ovate, sessile, short-decurrent, finely toothed. Capitula on very short
peduncles, readily falling, 15 mm long and 4 mm wide, oblong-
cylindrical, with few florets (three). Involucral bracts herbaceous, flat,
closely appressed, with slightly divergent cusp; outer and middle bracts
oblong-lanceolate, with membranous border from base, acuminate-
spinescent, with three or four pairs of horizontal, sometimes flexuous
spines on both sides; inner bracts entire, acuminate, with membranous
border from base to apex. Receptacular bristles smooth. Corolla exserted
from involucre, whitish. Achenes in capitulum usually solitary, large, 7
mm long and 4 mm wide, obovoid, glabrous, smooth, roundish above,
whitish or light gray. Flowering VI-VII; fruiting VII—VIII. (Plate XVII,
Fig. 1).

Loamy-stony slopes.—Soviet Central Asia: Tien Shan (northern and
western Tien Shan), Pamiro-Alai (Alai Range), mountainous Turkmenia
(Kopetdag).

Note. The hybrids of C. triflora and C. umbrosa are described in
the note to the latter species.

Section 24. Chrysis Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8
(1940) 515, 548.—Nudicaules Winkl. in Tr. Peterb. Bot. Sada, XII (1892)
191 p. p.——Capitula at the apices of short branches, solitary, glabrous or
weakly arachnoid-hairy; involucral bracts gradually acuminate into thin
cusps; corolla bright yellow; receptacular bristles smooth; achenes
obovoid, angular, distinctly toothed above. Perennial herbs with less
numerous and more or less poorly developed, cauline leaves and large,
usually rosulate, basal leaves.

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241
Lectotype of section: C. aurea Winkl.

126. C. aurea Winkl. in Tr. Peterb. Bot. Sada, X (1889) 476 quoad
sp. seravschan.; XII, 196 p. p.; XIV, 218; Bornm. in Beith. bot. Centralbl.
XXXIV, 2, 138 (excl. var.); Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR,
8, 548.—Arctium aureum O. Ktze. Rev. Gen. 1 (1891) 307.

Perennial. Stem erect, 30-100 cm high, striate-sulcate, almost
glabrous, branched from base, branches long, repeatedly branched further,
forming lax, broadly paniculate inflorescence. Basal leaves distinctly
rosulate; cauline leaves less numerous, much smaller than basal, strongly
reduced, and poorly developed, as a result, stem appearing almost
leafless, especially in upper part; leaves of basal rosette large, petiolate,
with petioles broadened at base, almost half as long as lamina, oblong-
ovate, undivided, finely toothed, with two or three pairs of broadly deltoid
basal lobes; lower cauline leaves petiolate, petioles winged, auricles of
wings fused with stem; upper leaves scaly; all leaves glabrous above,
green, white-tomentose beneath. Capitula small, 10 mm wide, ovoid,
very finely arachnoid-hairy. Involucral bracts oblong, closely appressed
at base, with erect or weakly divergent cusp; outer bracts smaller than
others but usually all bracts very similar to each other. Receptacular
bristles smooth. Corolla yellow. Achenes obovoid, almost three-angled,
with small teeth above, 6 mm long and 3 mm wide, slightly curved.
Flowering VI-VII; fruiting VII—VIII.

Pebbled and gravelly slopes in lower and middle mountain zone_—
Soviet Central Asia: Pamiro-Alai. Endemic. Described from the Voru
River Valley, between Santuruch and the Kshtut Pass. Type in Leningrad.

127. C. medians Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, VIII, Ser.
1, 3 (1936) 297; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, VIII, 549.

Perennial. Stems erect, up to 120 cm high, striate-sulcate, thick,
glabrous, profusely branched; branches long, in turn repeatedly branched,
forming broadly paniculate lax inflorescence. Basal leaves distinctly
rosulate; cauline leaves less numerous, much smaller than basal, poorly
developed, as a result stem appearing leafless, especially in upper part;
basal and lower cauline leaves petiolate, broad-ovate, broadly toothed;
middle and upper leaves sessile, lanceolate, prickly-toothed, reduced; all
leaves almost green, glabrous or finely arachnoid-hairy above, finely
tomentose beneath. Capitula numerous, 10—12 mm wide (excluding cusps),
ovoid, almost glabrous. Involucral bracts numerous, ovate to oblong-
ovate, abruptly narrowed into 5—7 mm long, horizontal cusp. Receptacular
bristles smooth. Corolla yellow. Achenes obovoid, angular, with scarcely
noticeable teeth above. Flowering VI-—VII; fruiting VII-VIII.

Stony screes in middle mountain zone.—Soviet Central Asia: Pamiro-

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242

Alai. Endemic. Described from the Kugitang Mountain, near the village
of Kugitang. Type in Leningrad.

128. C. refracta Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3
(1936) 297; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 549.—C.
aurea B. refracta Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916)
138.

Perennial. Stems erect, 60-120 cm high, striate-sulcate, almost
glabrous, profusely branched in upper half; branches long, in turn,
repeatedly branched, forming lax and spreading semiumbellate
inflorescence. Basal leaves distinctly rosulate; cauline leaves less
numerous, much smaller than basal, somewhat weakly developed, as a
result stem appearing almost leafless in upper part; basal leaves petiolate,
with petioles half as long as lamina, oblong-ovate or orbicular, entire,
irregularly toothed; cauline leaves strongly reduced, with narrow-winged
petioles, entire, finely toothed; uppermost leaves scaly; all leaves finely
arachnoid-hairy above, white-tomentose beneath. Capitula numerous,
globose-ovoid, 15 mm wide (excluding cusps), almost glabrous.
Involucral bracts imbricate, with up to 5 mm long thin and sharply
decurved cusp, all similar. Receptacular bristles smooth. Corolla yellow.
Achenes obovoid, almost three-angled, mildly curved, 6 mm long, 3
mm wide. Flowering VI-VII; fruiting VII-—VIII.

Outcrops of varicolored rocks in middle mountain zone.—Soviet
Central Asia: Pamiro-Alai. Endemic. Described from the Karatag River
Valley, the village of Hakimi. Type in Leningrad.

129. C. schmalhausenii Winkl. in Tr. Peterb. Bot. Sada, X (1889)
474; XII, 195; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 549.—
Arctium schmalhausenii O. Ktze. Rev. Gen. 1 (1891) 308.

Perennial. Stems erect, 120 cm high, striate-sulcate, glabrous or
somewhat glandular, branched in upper half; branches long, in turn,
repeatedly branched, forming lax paniculate inflorescence. Basal leaves
distinctly rosulate, cauline leaves less numerous, strongly reduced; as a
result, stem appearing almost leafless, especially above; basal leaves
petiolate, petioles strongly broadened at base, oblong-ovate, pinnatipartite
in upper half, and then undivided below, or pinnatilobate or pinnatipartite
throughout, and then petioles winged; cauline leaves sessile, strongly
reduced, uppermost leaves almost scaly; all leaves finely arachnoid-hairy
above, white-tomentose beneath. Capitula numerous, small, 9-10 mm
wide (excluding cusps), ovoid, finely arachnoid-hairy. Involucral bracts
oblong-ovate above with very short cusp, imbricate, all bracts similar.
Receptacular bristles smooth. Corolla yellow. Achenes oblong-obovoid,
angular. Flowering V—VII; fruiting VI-—VIII.

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243

Stony and gravelly slopes in lower mountain zone.—Soviet Central
Asia: Pamiro-Alai. Endemic. Described from the Zeravshan River Valley.
Oalyk Ravine. Type in Leningrad.

130. C. karatavica Rgl. and Schm. in Tr. Peterb. Bot. Sada, VI
(1880) 317; Winkl. ibid. XII, 194; XIV, 192; Bornm. in Beih. bot.
Centralbl. XXXIV, 2, 138; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR,
8, 549.—Arctium karatavicum O. Ktze. Rev. Gen. 1 (1891) 308.

Perennial. Stem erect, 40-150 cm high, striate-sulcate, weakly
pubescent, branched almost from base, branches long, in turn repeatedly
branched, forming broadly paniculate inflorescence. Basal leaves
distinctly rosulate; cauline leaves less numerous, much smaller than basal,
poorly developed; as a result, stem appearing almost leafless, especially
above, leaves in rosette petiolate, uppermost oblong-ovate, pinnatisect
in upper half, undivided below; inner leaves in rosette often pinnatisect
throughout, with lateral segments deeply lobed or pinnatipartite; cauline
leaves sessile, pinnatipartite or lobed; uppermost leaves reduced to scales;
all leaves very finely arachnoid-hairy above, almost smooth, white-
tomentose beneath. Capitula numerous, 7-10 mm wide, ovoid, very finely
arachnoid-hairy, almost glabrous. Involucral braccts closely appressed
at base, with falcately decurved, thin, up to 3 mm long cusp; outer
bracts small, up to 3 mm long; middle bracts oblong-ovate, 12-13 mm
long; inner bracts scarious, oblong-lanceolate, with short, divergent, thin
cusp. Receptacular bristles smooth; corolla yellow. Achenes obovoid,
angular, scarcely toothed above, 5 mm long and 2 mm wide. Flowering
VI-VII; fruiting VII—-VIII.

Stony and gravelly slopes in lower mountain zone.—Soviet Central
Asia: Tien Shan (west and center), Pamiro-Alai. Endemic. Described
from the Karatau Mountains, near Ishtyube. Type in Leningrad.

131. C. haesitabunda Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR,
8 (1940) 516, 549.

Perennial. Stems erect, 20-25 cm high, shallow-sulcate, finely
arachnoid-hairy, cymosely branched in upper half, sparsely leafy. Basal
leaves rosulate, less numerous, petiolate, oblong-lanceolate, sparsely
finely arachnoid-hairy above, white-tomentose beneath, pinnatisect, their
lateral segments pinnatilobate, prickly-toothed; cauline leaves strongly
reduced, sessile, semiamplexicaul, pinnatilobate or large-prickly-toothed.
Capitula ovoid, 10-12 mm wide (excluding cusps), glabrous. Involucral
bracts numerous, stiff, coriaceous, closely appressed at base, falcate in
upper half and acuminate-spinescent; inner bracts lanceolate, erect,
scarious with almost transparent border. Receptacular bristles smooth.
Corolla yellow. Achenes obovoid, almost four-angled. Flowering V—VI;
fruiting VI-VII. (Plate XIX, Fig. 1).

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244

Middle mountain zone.—Soviet Central Asia: Pamiro-Alai (Kaitash
Mountains, Malguzar). Endemic. Described from the Kaitash Mountains,
near the village of Karatash. Type in Leningrad.

132. C. chlorantha Kult. in Tr. Sredneaz. Gos. Univ. Ser. VI (1929)
4; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 549.—TIc.: Kult. op.
cit. Fig. 4.

Perennial. Stem erect, 30-35 cm high, striate-sulcate, partly floccose-
tomentose, sparsely branched from base, leaves crowded mainly in lower
part of stem; stem appearing almost leafless above because of less
numerous and usually poorly developed middle and upper leaves. Basal
leaves oblong-lanceolate, petiolate, pinnatipartite or pinnatisect, with
remote, oblong-lanceolate, sessile, entire or more or less distinctly toothed
lateral lobes; lower cauline leaves like basal; middle leaves one-third to
half as long as lower, sessile, semiamplexicaul, prickly-toothed; upper
and uppermost leaves scaly; all leaves finely arachnoid-hairy above,
white-tomentose beneath. Capitula ovoid, 12-17 mm wide (excluding
cusps), partly pubescent. Involucral bracts closely appressed at base,
coriaceous, ovate-oblong, herbaceous in upper half, abruptly and strongly
decurved and gradually acuminate in thin, falcate cusp; all bracts similar,
except inner oblong-lanceolate bracts, with membranous border above;
receptacular bristles smooth; corolla yellow. Achenes obovoid or almost
pyramidal, 5 mm long, three-angled, with scarcely noticeable three teeth
above, slightly curved. Flowering VI-VII; fruiting VII—VIII. (Plate
XXVII, Fig. 2).

Gravelly slopes in middle mountain zone.—Soviet Central Asia:
Pamiro-Alai (Turkestan Range). Endemic. Described from the region
between Gop and Suforidum. Type in Tashkent.

133. C. korolkovii Rgl. and Schm. in Tr. Peterb. Bot. Sada, VI
(1880) 315; Winkl. ibid. XII, 194; XIV, 192, 218; Juz. in Tr. Tadzh.
Bazy Akad. Nauk SSSR, 8, 549.—Arctium korolkovii O. Ktze. Rev.
Gen. 1 (1891) 308.

Perennial. Stem erect, 40-60 cm high, striate-sulcate, pubescent,
sparsely branched mainly in lower half, leaves crowded in lower half,
stem appearing leafless above because of few and poorly developed
middle and upper leaves. Basal leaves short-petiolate, oblong-lanceolate,
entirely pinnatisect, their lateral segments oblong, approximate, sessile,
lobed or entire; lower cauline leaves like basal, but sessile; middle leaves
one-fourth to one-third as long as lower, oblong, entirely pinnatipartite,
semiamplexicaul; upper and uppermost leaves scaly; all leaves coriaceous,
very finely arachnoid-hairy above, white-tomentose beneath. Capitula
narrowly ovoid, 6—9 mm wide (excluding cusps), glabrous. Involucral

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bracts ovate, closely appressed at base, acuminate in very short, erect
or erect-spreading cusp, all similar, except inner; inner bracts oblong,
obtuse, with membranous border. Receptacular bristles smooth; corolla
yellow. Achenes obovoid, angular, toothed above, 5 mm long. Flowering
VI-VII; fruiting VII-VIII.

Stony-clayey slopes in middle mountain zone.—Soviet Central Asia:
Pamiro-Alai (city of Nuratau), Kyzyl-Kum. Endemic. Described from
Kyzyl-Kum, between Khalaati and Adamkiriulgan. Type in Leningrad.

Section 25. Serratulopsis Tschern. in Bot. Mat. Gerb. Inst. Bot.
Akad. Nauk UzbSSR, XVII (1962) 83——Capitula solitary terminal on
long branches; involucral bracts closely appressed, with narrow
membranous border, acuminate; inner bracts with scarious, incised, purple —
apical appendage. Receptacular bristles smooth; corolla whitish. Achenes
obovoid, smooth, roundish above. Perennial plant with many-headed
woody root and coriaceous, almost undivided or crenate-lobed leaves.

Type of section: C. vavilovii Kult.

134. C. vavilovii Kult. in Sched. ad Herb. Fl. As. Med. 8 (1926)
No. 198; Byull. SAGU, 12, Suppl. 15.—Exs.: H.F.A.M. No. 198.

Perennial. Roots woody, many-headed. Stems numerous, erect,
slender, elegant, about 40 cm high, longitudinally sulcate, finely and
floccosely arachnoid-hairy, somewhat slender-branched almost through-
out; branches axillary, long, one-headed. Leaves coriaceous, with network
of prominent veins, light green and very finely arachnoid-hairy above,
finely appressed-gray tomentose beneath; basal leaves petiolate, about
15—20 cm long and 5-6 cm wide, ovate, undivided, crenate-lobed or
with two or three basal lobes; cauline leaves petiolate, ovate, undivided,
somewhat acuminate, gradually reduced; upper cauline leaves sessile,
oblong-lanceolate, entire or irregularly and finely crenate-toothed.
Capitula oblong-ovoid, glabrous, with about 20-florets, 20-22 mm long
and 8 mm wide. Involucral bracts closely appressed, about 55, ovate to
oblong-ovate, roundish above, acuminate, with narrow membranous
border, green; inner bracts with membranous appendage at apex
irregularly incised and slightly purple. Receptacular bristles smooth.
Corolla whitish; anther tube purple. Achenes obovoid, roundish above,
matty, transversely black-striate, 6 mm long and 2.5-3 mm wide.
Flowering VII; fruiting VII.

Rocky slopes in subalpine mountain zone.—Soviet Central Asia:
Tien Shan (Kirgizian Range). Endemic. Described from Uch-Bulak. Type
in Tashkent.

Section 26. Amberbopsis Tschern. in Bot. Mat. Gerb. Inst. Bot.
Akad. Nauk UzbSSR, XVII (1962) 83.—Capitula solitary terminal on

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246

long, almost leafless branches; involucral bracts somewhat flat, with
membranous border, acuminate; receptacular bristles smooth. Perennial
plant with many-headed woody root and thin coriaceous, almost without
spines, rather large leaves.

Type of section: C. grandifolia Kult.

135. C. grandifolia Kult. in Tr. Peterb. Bot. Sada, VIIIc, VI (1929)
9.—Ie.: Kult. ibid., Fig. 10.

Perennial. Root vertical, woody, many-headed. Stem erect, finely
longitudinally sulcate, finely and sparsely arachnoid-hairy, 80-100 cm
high, somewhat sparsely branched in upper half, branches axillary, long,
leafless, in turn, usually sparingly branched, one-headed. Leaves thin
coriaceous, more densely spinose, green, glabrous, and lustrous above,
appressed white-tomentose beneath; lower cauline leaves large; about
40 cm long and 25 cm wide, petiolate, oblong-ovate, almost lyrate,
with large irregularly crenate, orbicular, terminal lobe, and 7—10 pairs
broadly sessile, remote, crenate-toothed, lateral lobes reduced toward
base of lamina; middle cauline leaves few, like lower, but much smaller,
sessile, semiamplexicaul, with distinct basal lobes; upper leaves sessile,
attenuate toward base, undivided or crenate-lobed; uppermost leaves
strongly reduced. Capitula obovoid, 18—20 mm long (excluding corolla)
and 12-13 mm wide, basally truncate or weakly foveate, glabrous.
Involucral bracts about 55, closely appressed, green, with very short
cusp and irregularly incised membranous border from ovate to oblong-
ovate (outer bracts 4 x 5 mm, subsequent bracts 7 x 4 mm; middle bracts
8 x 4 mm, and inner bracts 17 x 3 mm); innermost bracts linear-
lanceolate, 17 mm long and 1.5 mm wide, short acuminate. Receptacular
bristles smooth. Corolla yellow; anther tube yellow. Achenes obovoid,
smooth, roundish above, with irregular black dots, 8 mm long and
3 mm wide. Flowering V—VI; fruiting VI.

Stony slopes in ravines.—Soviet Central Asia: Tien Shan (western
Tien Shan). Endemic. Described from the ravine of the Dau-Baby River.
Type in Tashkent.

Section 27. Stenoloma Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR,
8 (1940) 513, 547.—/nermes Winkl. in Tr. Peterb. Bot. Sada, XII (1892)
196 p. p.—Capitula solitary terminal on stem and branches, or in twos;
involucral bracts gradually acuminate into very long and short-subulate
cusp; corolla whitish, light pink or deep yellow; receptacular bristles
smooth or scabrous; achenes obovoid or obpyramidal, somewhat angular.
Perennial plants with entire leaves.

Lectotype of section: C. komarowii (O. Ktze.) Wink.

Series 1. Komarowianae Tschern. Stem wingless and, like leaves,

247

sparingly pubescent; capitula oblong-ovoid; cauline leaves sessile, short-
decurrent. Receptacular bristles smooth.

136. C. komarowii (O. Ktze:) Winkl. in Tr. Peterb. Bot. Sada, X
(1889) 470; XI, 132; XII, 199; Kult. in Fl. Turkm. VII, 178.—Arctium
komaroffii O. Ktze. in Tr. Peterb. Bot. Sada, X (1887) 197.—TIe.: Fl.
Turkm. VII, Plate 27, Fig. 1.

Perennial. Root woody, many-headed. Stem numerous, 30-40 cm
high, with remains of old petioles at base, usually simple, less often
branched, loosely arachnoid-hairy, grayish-green, later grayish-brown,
stems and branches one-headed. Leaves soft, grayish-green and arachnoid-
hairy above, finely white-tomentose beneath; basal leaves linear-
lanceolate, petiolate, attenuate toward both ends, up to 15 cm long and
1—2 mm wide, usually undivided and entire or pinnatilobate, acuminate
into short cusp; cauline leaves gradually reduced toward apex; middle
leaves up to 5 cm long and 5—12 mm wide, sessile, semiamplexicaul,
decurrent, with usually prickly-toothed auricles; upper cauline leaves
reduced. Capitula ovoid-oblong. 7-8 mm wide, arachnoid hairy-
pubescent. Involucral bracts 60-80, all except inner linear-subulate, 5—
14 mm long, with triquetrous, erect cusps; inner bracts coriaceous; linear-
lanceolate, short-spinescent, usually light pink. Receptacular bristles
smooth, slightly broadened at apex. Corolla pink; anther tube purple.
Achenes obovoid, more or less angular, 4.5-5 mm long, light gray,
with small teeth above. Flowering VI; fruiting VII.

Stony-loamy slopes in middle mountain zone.—Soviet Central Asia:
mountainous Turkmenia (Kopetdag). General distribution: Northern Iran.
Described from Kopetdag in the Ashkhabad District. Type in Leningrad.

137. C. stenophylla Kult. in Tr. Tadzh. Bazy Akad. Nauk SSSR,
Villc, VI (1929) 7; Kult. in Fl. Turkm. VII, 176.—Ic.: Fl. Turkm. VII,
Plate 27, Fig. 2.

Perennial. Root woody, many-headed. Stems numerous, 30-40 cm
high, slender, dark brown, finely arachnoid-hairy, simple or sparsely
branched. Leaves soft, withering early, linear, acuminate, undivided,
recurved, gray arachnoid-hairy on both sides; basal leaves petiolate,
attenuate toward both ends, 12 cm long and 4-5 mm wide; cauline
leaves gradually reduced, sessile, short-decurrent, slightly broadened at
base, with three to five small teeth. Capitula ovoid-oblong or more or
less ovoid, 8 mm wide (excluding cusps) and 15 mm long (excluding
florets), densely arachnoid-hairy. Involucral bracts about 60, all except
inner with subulate-triquetrous, long, almost horizontal cusp, 7-22 mm
long; inner bracts erect, coriaceous, scabrous along margin, usually
purple, linear, 16-18 mm long. Receptacular bristles broadened above,

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smooth. Corolla whitish; anther tube pink. Achenes obpyramidal, three
or four-angled, gray, toothed above, 4 mm long, and 1.5 mm wide.
Flowering VI-VII; fruiting VII—VIII.

Loamy mountain slopes.—Soviet Central Asia: mountainous Turkmenia
(Kopetdag). Endemic. Described from the vicinity of Ashkhabad. Type
in Tashkent.

Series 2. Leucanthae Tschern.—Plants very sparsely pubescent;
stems wingless; cauline leaves broadly long-decurrent. Capitula oblong-
ovoid; receptacular bristles smooth.

138. C. leucantha Bornm. and Sint. in Russk. Bot. Zhurn. 1 (1911) -
1; Kult, in Fl. Turkm. VII, 178.—C. simplex Kult. in FI. Turkm. VII
(1960) 180 in obs. (ross.).—Ie.: Bornm. and Sint. op. cit. Plate 1; Kult.
op. cit. Plate 28, Fig. 1.

Perennial. Root woody, many-headed; sterile branches creeping,
somewhat decumbent; stems several or solitary, floccose-arachnoid-hairy,
with remains of old leaves at base, sparsely branched in upper half,
with long, one-headed branches. Leaves soft, finely arachnoid-hairy
above, finely gray-tomentose beneath; basal leaves lanceolate, petiolate,
attenuate toward apex, undivided entire, 13-17 cm long and 1.5—3 cm
wide; cauline leaves gradually reduced toward stem apex, sessile, rather
long-decurrent, usually finely prickly-toothed; leaves on branches strongly
reduced. Capitula ovoid-oblong, 9-13 mm wide (excluding cusps), finely
arachnoid-hairy. Involucral bracts about 70, outer bracts broadened at
base, attenuate into short, usually divergent cusp; middle bracts broad-
lanceolate, deltoidly broadened in upper part, acuminate-spinescent with
short, erect dark cusp, margin scabrous; inner bracts membranous-
coriaceous, about 20 mm long, broadened above, gradually attenuate
into short thin cusp, scabrous along margin. Receptacular bristles slightly
broadened above, smooth. Corolla whitish or light pink; anther tube
pink. Achenes obovoid, 4-5 mm long, gray, with dark spots, somewhat
angular. Flowering VI—VII; fruiting VII.

On dry slopes.—Soviet Central Asia: mountainous Turkmenia
(Kopetdag). Endemic. Described from Kyzyl-Arvat. Type in Leningrad.

Series 3. Griseae Tschern.—Stem very narrowly and discontinuously
winged, arachnoid hairy-tomentose; cauline leaves long- and narrow-
decurrent. Capitula ovoid; receptacular bristles scabrous.

139. C. grisea Kult. in Tr. Peterb. Bot. Sada, VIIIc, VI (1929) 5;
Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 547.—C. komarowii var.
patula Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916) 145.—Ic.: Kult.
opyicity) Figai6:

249

Plate XVIII.
rigida Kult.; 2—C. grandifolia Kult.

253

—Cousinia

254

250

Perennial. Stems numerous, slender, 10-25 cm high, very narrowly
and discontinuously winged, arachnoid hairy-tomentose, glabrescent, and
then lustrous, pinkish, leafy, moderately branched above; branches
slender, one-headed. Leaves soft, grayish-green above, white-tomentose
beneath, entire; basal leaves linear-oblong, thin-petiolate, acuminate,
lower and middle cauline leaves sessile, linear, narrowly long-decurrent,
gradually narrowed toward apex in thin and short spine, entire or
sometimes with two or three pairs of small teeth at base; upper cauline
leaves small, sessile, almost nondecurrent or short-decurrent. Capitula
ovoid, 8 mm wide (excluding cusps), pubescent. Outer and middle
involucral bracts green, appressed at base, linear-lanceolate, gradually
attenuate into falcate, subulate cusp; inner bracts erect, linear, finely
cuspidate. Receptacular bristles broadened above, scabrous. Corolla
whitish (?); anther tube pink. Achenes oblong, scarcely angular, light
gray with dark spots, 4 mm long and 2 mm wide. Flowering VI-VII;
fruiting VII—VIII.

Loamy-gravelly slopes in middle mountain zone.—Soviet Central
Asia: Pamiro-Alai (southwestern spurs of the Gissar Range). Endemic.
Described from the Yakkabag District near Tutakat. Type in Tashkent.

Series 4. Alaicae Tschern.—Stems narrow-winged and like leaves
sparingly pubescent; cauline leaves sessile, decurrent. Capitula ovoid;
receptacular bristles smooth.

140. C. alaica Juz. ex Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 90.

Perennial. Stems numerous, erect, slender, pubescent, 15—25 cm high
narrow-winged throughout, wings 1-2 mm wide, leafy, branched in upper
half; branches slender, long, with two capitula each at tips on relatively
short peduncles, or stems simple, with two terminal capitula. Leaves
soft, long-decurrent, entire, grayish green above due to fine pubescence,
gray-tomentose beneath; basal leaves thin-petiolate, linear, acuminate-
finely spinescent; cauline leaves gradually reduced toward stem apex,
sessile, decurrent, linear, acuminate-finely spinescent, sometimes with
two subulate teeth at base. Capitula ovoid, 7 mm wide (excluding cusps),
pubescent. Involucral bracts, except inner, imbricate, appressed at base,
acuminate in fine, erect cusp; inner bracts linear, broadened above,
orbicular, with thin membranous border and thin spine. Receptacular
bristles slightly broadened above, smooth. Corolla apparently white.
Achenes weakly angular, gray, toothed above, 3—3.5 mm long. Flowering
VI-VII; fruiting VIJ-VII.

Gravelly slopes in middle mountain zone.—Soviet Central Asia:
Pamiro-Alai (Alai and Trans-Alai ranges). Endemic. Described from the

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251

Alai Range, vicinity of Shahimardan, along Peshkaut Gorge. Type in
Tashkent.

141. C. fallax Winkl. in Tr. Peterb. Bot. Sada, X (1889) 471; XII,
200; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 547.—C. fallax var.
armata Winkl. ex O. and B. Fedtsch. Perech. Rast. Turk. IV (1911)
243.

Perennial. Many-branched plants. Stems with remains of petioles of
previous year’s leaves at base, 12-35 cm high, erect or somewhat
ascending, finely white arachnoid-hairy, winged throughout with wings
2—2.5 mm wide, leafy, usually branched above; branches short, one-
headed. Leaves soft, entire, less often with sparse small teeth, grayish-
green and finely arachnoid-hairy above, finely grayish tomentose beneath;
basal leaves thin-petiolate, narrow-lanceolate, attenuate toward both ends,
acuminate-finely cuspidate; cauline leaves reduced, sessile, decurrent into
stem wings, narrow-lanceolate, attenuate above into short cusp. Capitula
ovoid, 6—9 mm wide (excluding cusps), finely arachnoid-hairy. Involucral
bracts except inner appressed at base, oblong-ovate, attenuate-acute with
falcate cusp; inner bracts broadened above, scarious, orbicular-ovate,
acuminate-finely cuspidate. Receptacular bristles slightly broadened
above, smooth. Corolla deep yellow, less often turning red on drying.
Achenes weakly angular, obovoid, 3 mm long, light gray. Flowering
VII; fruiting VIII-IX. (Plate XIX, Fig. 2).

Stony-gravelly slopes in the upper part of middle mountain zone.—
Soviet Central Asia: Pamiro-Alai (southern slopes of the Gissar Range,
Baldzhuan). Endemic. Described from the vicinity of the village of
Kalkot and the Sanglak Mountains. Type in Leningrad.

Series 5. Baranovianae Tschern.—Stems wingless and like leaves
densely pubescent; cauline leaves sessile. Capitula globose; receptacular
bristles smooth.

142. C. baranovii Juz. ex Tschern. in Bot. Mat. Gerb. Inst. Bot.
Akad. Nauk UzbSSR, XVII (1962) 92.

Perennial. Root slender, woody, many-headed, their collars covered
with scaly remains of old leaves, bearing numerous sterile rosettes and
floriferous stems. Stems erect, white-tomentose, simple, one-headed,
leafy. Leaves finely gray-tomentose above, white-tomentose beneath,
entire; leaves of basal rosette narrow-lanceolate, narrowed toward both
ends, short-petiolate, 11-12 cm long, 7-8 mm wide, soft; cauline leaves
gradually reduced, sessile, short-acuminate, sometimes with three to five
pairs of thin spines at base. Capitulum globose, 12-13 mm wide
(excluding cusps), rusty-tomentose; involucral bracts carinate to narrow-
lanceolate, gradually acuminate, falcately decurved; middle and inner

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Pe

bracts dorsally dark, purple; innermost bracts erect, spinescent.
Receptacular bristles smooth. Corolla whitish (?); anther tube light pink.
Flowering VII.

On mountain slopes.—Soviet Central Asia: Tien Shan (Talass
Alatau). Endemic. Described from the ravine of Kara-Berek River Valley.
Type in Tashkent.

Section 28. Kuekenthalia Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR,
8 (1940) 513, 548.—Capitula solitary terminal on branches, more or
less numerous on long peduncles; involucral bracts gradually attenuate-
cuspidate; corolla pale yellow or purple; receptacular bristles scabrous;
achenes glabrous, smooth, roundish above. Biennial plants with narrow-
winged stems and sessile or decurrent leaves.

Lectotype of section: C. kuekenthalii Bornm.

Series 1. Neglectae Juz.—Stems strongly branched, with a significantly
large number (10-60) of capitula; leaves weakly spinose.

143. C. kuekenthalii Bornm. in Beih. bot. Centralbl. XXXIV, 2
(1916) 146; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 548.

Perennial. Stems erect, 20-50 cm high, narrow-winged, with wings
3-4 mm wide, sparsely arachnoid-hairy, sparsely branched in upper
half. Leaves decurrent, narrow lanceolate, glaucous-green and sparsely
arachnoid-hairy above, white-tomentose beneath; basal leaves petiolate,
with narrow-winged petioles, pinnatisect; lower cauline leaves like basal
but sessile; middle and upper leaves narrow-lanceolate, prickly-toothed;
uppermost leaves reduced, spinescent. Capitula 15-25, on relatively short
and wingless peduncles, oblong-ovoid, 8 mm wide (excluding cusps),
finely arachnoid-hairy. Involucral bracts erect-spreading, strong, narrow-
lanceolate, carinate, with thin, 3-5 mm long cusp; inner and innermost
bracts coriaceous, linear, acuminate-spinescent. Receptacular bristles
white below, smooth, broadened above, scabrous. Corolla pink; anther
tube purple. Achenes obovoid, glabrous, smooth, glossy, roundish above,
light gray with dark spots, 3 mm long. Flowering VII—VIII; fruiting
Vil.

Outcrops of red clay or red sandstone in middle mountain zone.—
Soviet Central Asia: Pamiro-Alai. Endemic. Described from the Takhta-
Karacha Pass, near the village of Amankutan. Type in Leningrad.

144. C. neglecta Juz. in Bot. Mat. Gerb. Glavn. Bot. Sada, IV
(1923) 78; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 548.—C.
komarowii auct. non Winkl. in Tr. Peterb. Bot. Sada, XIV (1897) 219.

Biennial. Stems erect, 30-50 cm high, winged throughout, wings in
many rows and 1-3 mm wide, weakly pubescent, smooth at extreme
tip, branched almost from base, with long, slender, repreatedly branched

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253

twigs, forming broad inflorescence, leaves long-decurrent, mostly
recurved, greenish and arachnoid hairy-tomentose above, white or gray-
tomentose beneath; basal leaves withering before anthesis; lower cauline
leaves oblong-lanceolate, pinnatipartite or pinnatisect, with lanceolate,
entire, short-acuminate segments; upper leaves undivided, with large and
remote teeth; uppermost leaves reduced. Capitula numerous (25-65),
ovoid, 6-9 mm wide (excluding cusps), pubescent. Involucral bracts 75,
herbaceous, closely appressed at base, ovate, gradually acuminate into
subulate, thin, erect spine, almost as long as appressed part; inner bracts
coriaceous, lanceolate, acuminate-spinescent. Receptacular bristles whitish
below, smooth, thickened toward apex, light brown, scabrous. Corolla
straw-yellow; anther tube purple. Achenes obovoid, almost truncate
above, weakly longitudinally ribbed. Flowering VII—VIII; fruiting
VITI-IX.

Gravelly slopes in middle mountain zone.—Soviet Central Asia:
Pamiro-Alai (Zeravshan Range). Endemic. Described from Mussabazari.
Type in Leningrad.

145. C. dissectifolia Kult. in Tr. Turk. Nauchn. Obshch. 1 (1923)
111; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 548.—Iec.: Kult. op.
cit. Plate (1).—Exs.: H.F.A.M. No. 480.

Biennial. Stem erect, 30-50 cm high, with narrow, 3—4 mm wide
wings, finely floccose-arachnoid-hairy, lax corymbosely branched in
upper half. Leaves decurrent, gray-tomentose on both sides, more densely
beneath; basal leaves petiolate, with narrow-winged petioles, pinnatisect,
with numerous (up to 10) lateral segments; lower cauline leaves like
basal but sessile; middle and upper leaves lanceolate, entire or sparsely
prickly-toothed; uppermost leaves reduced. Capitula 15-30, ovoid, 10
mm wide (excluding cusps), finely arachnoid-hairy. Involucral bracts
stiff, carinate, acuminate in divergent, 3-4 mm long cusp; inner and
innermost bracts linear, coriaceous, with erect acuminate-spinescent cusp.
Receptacular bristles whitish below, smooth, broadened in upper half,
scabrous. Corolla bright yellow; anther tube purple. Achenes obovoid,
glabrous, smooth, roundish above, 4 mm long. Flowering V—VI; fruiting
VI-VII.

On clayey slopes.—Soviet Central Asia: Pamiro-Alai (Samarkand
Mountain). Endemic. Described from Zerabulak. Type in Tashkent.

146. C. decurrentifolia Juz. ex Tschern. in Bot. Mat. Gerb. Inst.
Bot. Akad. Nauk UzbSSR, XVII (1962) 53.

Biennial. Stem erect, 25-70 cm high with narrow, 3-4 mm wide
wings, finely arachnoid-hairy, profusely cymosely branched in upper
half; branches long. Leaves decurrent, sparsely above and densely gray-

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254

tomentose beneath; basal leaves petiolate, with narrow-winged petioles,
lanceolate, almost entire, sometimes with one or two pairs of baSal teeth;
lower cauline leaves like basal, but sessile; middle and upper leaves
linear-lanceolate, entire; uppermost leaves reduced. Capitula 10—25(60),
on long peduncles, 6 mm wide (excluding cusps), ovoid, arachnoid-
hairy. Involucral bracts stiff, carinate, with short, up to 2 mm long,
arcuate cusp; inner bracts linear, with straight, erect cusp. Receptacular
bristles broadened upward, scabrous. Corolla bright yellow; anther tube
purple. Achenes obovoid, glabrous, smooth, roundish above, 3 mm long.
Flowering VI—VII; fruiting VII—VIII. (Plate XX, Fig. 2).

Loamy-gravelly and stony slopes in middle mountain zone.—Soviet
Central Asia: Pamiro-Alai. Endemic. Described from the vicinity of the
village of Tash-Kurgan. Type in Tashkent.

Series 2. Ulotomae Juz.—Stem with few branches; leaves almost
unarmed. Capitula three to seven.

147. C. ulotoma Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916)
148; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 548.

Biennial. Stem erect, 15—20 cm high, densely arachnoid hairy-
gray tomentose and narrow-winged from base, or with few branches
from middle; branches long, one-headed. Leaves soft, grayish-green,
densely arachnoid-hairy on both sides, pinnatisect; lower leaves often
bipinnatisect, with narrow-lanceolate segments, usually weakly crisped,
early withering; all leaves except lower short-petiolate, sessile, decurrent
into wings; cauline leaves gradually reduced toward stem apex; leaves
on branches strongly reduced. Capitula broadly cylindrical, 9 mm wide
(excluding cusps), floccose-arachnoid hairy. Involucral bracts about 40;
outer bracts appressed at base, ovate, attenuate into short and recurved
cusp; middle bracts oblong, appressed at base, gradually narrowed into
almost arcuate cusp; inner bracts yellowish, membranous, linear-
lanceolate (about 2 mm wide), 15-17 mm long, gradually acuminate in
thin short cusp. Receptacular bristles more or less broadened above,
scabrous. Corolla pale yellow; anther tube pink. Achenes obovoid, 4
mm long and 1.5 mm wide, glabrous, smooth, roundish above, light
brown with dark spots. Flowering VII; fruiting VII. (Plate XX, Fig. 1).

Soviet Central Asia: Pamiro-Alai (Zaravshan Range). Endemic.
Described from the vicinity of the villages of Madm and Kum. Type in
Leningrad.

Section 29. Lasiandrae Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2
(1865) 40.—Capitula solitary terminal; involucral bracts appressed,
attenuate at divergence and broadened into grooved, subulate ciliate-

255

259 toothed cusp; receptacular bristles smooth; corolla yellow; anther tube
yellow, velutinous due to flexuous, thin, and arachnoid-hairy pubescence;
achenes narrow-obpyramidal, weakly ribbed, roundish above. Perennial
(may be monocarpic) plant with woody many-headed roots, white-
tomentose caudices and more or less cordate basal cauline leaves.

Lectotype of section: C. lasiandra Bge.

148. C. lasiosiphon Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1,
3 (1936) 311; Kult. in Fl. Turkm. VII, 170.

Perennial (Monocarpic ?). Roots woody, caudices white-tomentose
with remains of old petioles. Stem erect, 30-35 cm high, arachnoid
hairy-tomentose, sulcate, straw-yellow, branched almost from base or
middle, branches long, usually repeatedly branched further, one-headed.
Leaves weakly floccose-arachnoid hairy, green, and slightly lustrous
above, densely gray-tomentose beneath; basal leaves short-petiolate, with
flattened petiole, usually arachnoid hairy-tomentose below; lamina 11-—
14 cm long and 5—7 cm wide, ovate or obovate, somewhat attenuate
toward both ends, with large and irregularly erose-toothed, somewhat
obtuse, abruptly acuminate-spinescent; cauline leaves reduced toward
apex; lower cauline leaves broad-ovate, middle almost orbicular; all
sessile, deeply cordate, crenate-prickly-toothed, apex roundish, with short
and thin spine; upper leaves strongly reduced. Capitula 7-14 on each
plant, 2—2.5 cm wide (excluding cusps), ovoid or broadly ovoid, truncate
at base or weakly roundish, arachnoid-hairy. Involucral bracts about 80;
outer and middle bracts lanceolate, appressed at base, slightly narrowed
at divergence and broadened into grooved cusp with thin erect spines
along margin, acuminate into hard, stiff and subulate spine; inner bracts
thin-coriaceous, linear, scabrous dorsally, somewhat flat, gradually
acuminate in long and thin spine. Receptacular bristles smooth, white.
Corolla yellow; anther tube yellow, velutinous due to flexuous, thin,
arachnoid-hairy pubescence. Achenes narrow-obpyramidal, acuminate
towards base, 3.5-5 mm long, 1—1.5 mm wide, somewhat ribbed, with
narrow fringe along margin, roundish above, matty, finely matted, light
brown with dark spots. Flowering VII; fruiting VIII. (Plate XXI,
Fig. 2).

Stony slopes.—Soviet Central Asia: mountainous Turkmenia
(Badkhyz). Endemic. Described from the vicinity of Pul-i-Khatum. Type
in Leningrad.

Section 30. Cynaroideae Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2

(1865) 47.—Macrocephalae Bge. Ibid., 41.—Appendiculatae Winkl. in

Tr. Peterb. Bot. Sada, XII (1892) 268.—Constrictae Winkl. Ibid., 264

260 p. min. p.—Capitula solitary terminal on stem on branches; involucral

256

bracts narrowed in middle and broadened above into coriaceous, broadly
lanceolate, weakly carinate or almost sagittate, short-acuminate, divergent
appendages; corolla purple, whitish, or yellow; receptacular bristles
scabrous, less often smooth; achenes obpyramidal, angular, narrow-
winged along angles, toothed above. Perennial monocarpic or biennial
plants, usually with long-decurrent cauline leaves; stems and branches
almost leafless in upper part.
Lectotype of section: C. cynaroides (MB.) C.A.M.

149. C. onopordioides Ldb. in Echwald, Casp. Cauc. (1831-1833)
40; DC. Prodr. VI, 556; Ldb. Fl. Ross. II, 680; Bge. in Mém. Acad.
Pétersb. 7 sér. 92, 54; Boiss. FI. or. III, 507; Winkl. in Tr. Peterb. Bot.
Sada, XI, 137; XII, 277; XIV, 235; Takht. in Tr. Arm. Fil. Akad. Nauk
SSSR, Biol. Ser. 2, 191; Kult. in Fl. Trukm. VII, 172.—C. karelini
Less. in Bull. Soc. Nat. Mosc. XII (1839) 159.—C. albicaulis Boiss.
and Buhse in Nouv. Mém. Soc. Nat. Mosc. XII (1860) 128; Bge. op.
cit. 2, 53; Boiss. Fl. or. III, 507; Winkl. in Tr. Peterb. Bot. Sada, XII,
278; Kult. in Fl. Turkm. VII, 174.—Onopordon polyacanthum Eichwald,
op. cit. (1831-1833) 3, nomen.—Arctium polyacanthum, A. albicaule
and A. karelini O. Ktze. Rev. Gen. 1 (1891) 307 and 308.—Ie.: Eichwald,
op. cit. tab. 37; Fl. Turkm. VII, Plate 25; Boiss. and Buhse, op. cit. tab.
a

Perennial (apparently monocarpic). Stem erect, densely arachnoid
hairy-tomentose at base, with remains of old petioles; 80-100 cm high,
whitish, glossy, glabrous or arachnoid-hairy, sulcate, broad-winged due
to wide long-decurrent leaves, branched, branches spreading, up to 50
cm long, one-headed. Leaves coriaceous, grayish-green on both sides;
finely arachnoid-hairy or glabrous, with prominent veins; basal leaves
petiolate, broad-lanceolate, 18-30 cm long, 6—8 cm wide, pinnatilobate,
with acuminate-spinescent, prickly-toothed lobes; cauline leaves sessile,
broadly long-decurrent, gradually reduced toward stem apex, oblong-
lanceolate, deeply crenate-toothed, with acuminate-spinescent, prickly-
toothed teeth, cuspidate, slightly divergent from stem. Capitula solitary,
about 40 mm wide (excluding cusps), arachnoid-hairy, ovoid-globose,
with many florets. Involucral bracts numerous (about 100); outer bracts
coriaceous, green, 35 mm long, 10 mm wide, carinate dorsally, broad-
lanceolate, with three or four pairs of thin erect spines along margin,
cuspidate, decurved; middle bracts about 50 mm long, appressed at base,
8-9 mm wide, hard-coriaceous, yellowish, glabrous, attenuate in middle,
then broadened into broad-lanceolate, coriaceous, green, somewhat
carinate, divergent appendage, acuminate into stiff cusp toward apex and
with two or three pairs of stiff erect spines along margin; inner bracts 60 —
mm long, 4 mm wide, linear-lanceolate, glabrous, glossy, gradually and

257

Plate XIX.

1—Cousinia haesitabunda Juz.; 2—C. fallax Wink\.

261

263

258

long acuminate, scabrous along margin, erect. Receptacular bristles
scabrous. Corolla pink (?); anther tube pink. Achenes obpyramidal, 5—
7.5 mm long, 2-4 mm wide, ribbed, with corona above. Flowering V—
VI; fruiting VII.

Stony, gravelly slopes of lower mountain zone and in the foothills —
Caucasus: Eastern Transcaucasia; Talysh; Soviet Central Asia: Aralo-
Caspian (Mangyshlak), Kara-Kum; mountainous Turkmenia (Kopetdag,
Bolshie Balkhany). General distribution: Iran, Pakistan, Afghanistan.
Described from the vicinity of Tyu-Karagan (Mangyshlak). Type in
Leningrad.

150. C. cynaroides (MB.) C.A.M. in Verz. (1831) 230; DC Prodr.
VI, 556; Ldb. Fl. Ross. II, 680; Bge. in Mém. Acad. Pétersb. 7 sér. 9,
2 50; Boiss. Fl. or. III, 506; Winkl. in Tr. Peterb. Bot. Sada, XI, 137;
XII, 276; XIV, 215; Grossh. Fl. Kavk. IV, 168; Takht. in Tr. Arm. Fil.
Akad. Nauk, Biol. Ser. 2, 191; Kult. in Fl. Turkm. VII, 174; Isaev in
Fl. Azerb. VIII, 363.—Carthamus cynaroides MB. in Beschr. Land.
Casp. (1800) 195; MB. FI. taur.-cauc. II, 285——Onobroma cynaroides
Spreng. Syst. veg. ed. 3 (1826) 392; C.A.M. Verz. (1831) 67.—Arctium
cynaroides O. Ktze. in Rev. Gen. 1 (1891) 307.—Iec.: Fl. Azerb. VIII,
Plate 37.

Perennial. Stem with numerous remains of old leaves at base,
tomentose, erect, 30-40 cm high, usually white arachnoid-hairy, densely
leafy, discontinuously winged, with decurrent leaves, usually branched;
branches one-headed. Leaves coriaceous, usually arachnoid-hairy on both
sides, more dense beneath, grayish green; basal leaves petiolate,
lanceolate-oblong, pinnatilobate, with acuminate-spinescent, approximate
prickly-toothed lobes; cauline leaves sessile, broadly long-decurrent,
gradually reduced toward stem apex, ovate, prickly-toothed, acuminate-
cuspidate, slightly divergent; uppermost leaves reaching capitula. Capitula
20 mm wide (excluding cusps), ovoid-globose, arachnoid-hairy. Involucral
bracts numerous; outer bracts 18 mm long, green, coriaceous, about 10
mm wide at base, broad-lanceolate, with two or three pairs of spinescent
teeth along margin, acuminate into stiff, decurved cusp; middle bracts
appressed at base, stiff-coriaceous, glabrous, attenuate in middle and
broadened into lanceolate, coriaceous, green, cuneate, 13 mm long and
up to 5 mm wide, divergent appendage with one or two pairs of spines
along margin, acuminate-cuspidate, subsequent bracts similar but with
longer base. and slightly shorter appendage; inner bracts 26 mm long
and 4 mm wide, linear-lanceolate, gradually acuminate into long, thin
deflexed cusp, scabrous dorsally and along margin. Receptacular bristles
unequal, longest scabrous. Corolla yellow; anther tube pink. Achenes
angular, narrow-winged along edges, 44.5 mm long and 1.5 mm wide,

264

259

gray, obpyramidal, with fine teeth above. Flowering V—VI; fruiting
VIL.

Dry slopes.—Caucasus: Eastern Transcaucasia, Talysh; Soviet
Central Asia: mountainous Turkmenia. General distribution: Iran (north).
Described from Talysh. Type in Leningrad.

151. C. macrocephala C.A.M. Verz. (1831) 231; DC. Prodr. VI,
556; Ldb. FI. Ross. II, 681; Bge. in Mém. Acad. Pétersb. 7, sér. 9, 2,
41; Boiss. Fl. or. III, 503; Winkl. in Tr. Peterb. Bot. Sada, XII, 271;
Grossh. Fl. Kavk. IV, 168; Takht. in Tr. Arm. Fil. Akad. Nauk SSSR,
Biol. Ser. 2, 190; Fl. Azerb. VIII, 370.—C. macrocephala B. spinulosa
Lomak. ex Lipsky; Fl. Kavk. (1899) 355; Grossh. Fl. Kavk. IV, 168.—
Onobroma macrocephalum C.A.M. in Verz. (1831) 67.—Arctium
macrocephalum O. Ktze. Rev. Gen. 1 (1891) 308.

Biennial (or monocarpic). Stem erect, tomentose at base, with
remains of old petioles, 30-40 cm high, arachnoid-hairy, longitudinally
sulcate, discontinuously winged due to decurrent leaves, usually weakly
branched from base; branches densely leafy, one-headed. Leaves
coriaceous, grayish-green, arachnoid-hairy on both sides, more densely
beneath, lower side with strongly raised veins; basal leaves petiolate,
with short, somewhat flat petioles, oblong, pinnatilobate, with unequal
lanceolate, acuminate-spinescent lobes; cauline leaves gradually reduced,
sessile, broadly long-decurrent, oblong-lanceolate to ovate, broadly
crenate-prickly-toothed attenuate-cuspidate. Capitula 30-35 mm wide
(excluding cusps), somewhat arachnoid-hairy, flat-globose. Involucral
bracts numerous (about 80); outer bracts broad-lanceolate, herbaceous,
coriaceous, carinate, cuspidate, usually recurved, 20-40 mm long, smooth
along margin or less often with thin short spines; middle bracts 30-45
mm long, appressed at base, 6 mm wide, attenuate in middle, broadened
above into broad-lanceolate, coriaceous, carinate appendages, latter
smooth, acuminate into stiff short cusp, smooth along margin; inner
bracts 35 mm long, narrow-lanceolate, gradually acuminate. Receptacular
bristles unequal, longest somewhat scabrous. Corolla yellow; anther tube
purple. Achenes obovoid, 5 mm long, indistinctly longitudinally ribbed,
truncate above. Flowering VI-VII; fruiting VIII-IX.

Dry slopes in middle mountain zone.—Caucasus: Eastern and
southern Transcaucasia, Talysh. General distribution: Iran (north).
Described from Talysh. Type in Leningrad.

152. C. lyrata Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2 (1865) 53;
Boiss. Fl. or. III, 509; Winkl. in Tr. Peterb. Bot. Sada, XII, 278; XIV,
235; Kult. in Fl. Turkm. VU, 170.—C. caesia Winkl. op. cit. (1897)
238; Kult. op. cit. VII, 171—Arctium lyratum O. Ktze. Rev. Gen. 1
(1891) 308.

—N

260

Biennial, or perennial. Plants branched from base. Stem erect, weakly
branched in upper half, arachnoid-hairy; branches one-headed. Leaves
green and finely arachnoid-hairy above, gray-tomentose beneath; basal
leaves 10-16 cm long, petiolate, lyrate, with large, roundish, somewhat
cordate, and finely prickly-toothed terminal lobe and broad-lanceolate,
sessile lateral lobes; cauline leaves 5—7 cm long, sessile, broadly short-
decurrent on stem, oval-ovate, divergent, obtuse, with short spine,
irregularly prickly-toothed, gradually reduced toward stem apex;
uppermost leaves 1.5—2 cm long. Capitula globose, 2—2.5 cm wide
(excluding cusps, 4-6 cm with cusps) and 4—4.5 cm long (with florets),
arachnoid-hairy, somewhat foveate at base. Involucral bracts about 65,
all except inner with thick coriaceous, broad-lanceolae or almost rhombic
appendages, 8-12 mm broad at base, usually with prominent network of
veins, gradually acuminate into stiff grooved cusp with two or three
spinescent teeth along margin; involucral bracts themselves coriaceous,
ovate, closely appressed; inner bracts erect, almost linear, often pink,
gradually acuminate, short-spinescent. Corolla pink, with yellow anther
tube. Receptacular bristles scabrous. Achenes obpyramidal, dark, four-
angled, 5 mm long, with three or four small teeth above. Flowering VI—
VII; fruiting VII—-VIII. (Plate XXI, Fig. 1).

Stony-gravelly slopes.—Soviet Central Asia: mountainous Turkmenia
(Kopetdag, Badkhyz). General distribution: Northern Iran. Described from
Iran. Type in Leningrad.

153. C. purpurea C.A.M. ex DC. Prodr. VI (1837) 555; Ldb. FI.
Ross. II, 680; Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2, 51; Boiss. FI. or.
III, 511; Winkl. in Tr. Peterb. Bot. Sada, XII, 277; Grossh. Fl. Kavk.
IV, 167; Takht. in Tr. Arm. Fil. Akad. Nauk SSSR, Biol. Ser. 2, 191;
Takht. and Fedtsch. Fl. Erev. 318; Fl. Azerb. VIII, 372.—Arctium
purpureum O. Ktze. Rev. Gen. 1 (1891) 308.—Exs.: Pl. or. exs. No.
Pia

Biennial (or monocarpic). Stem erect, with scaly remains of old
petioles at base, 15-30 cm high, finely arachnoid-hairy, longitudinally
sulcate, discontinuously winged with long-decurrent leaves, branched
almost from base; branches leafy, one-headed. Leaves coriaceous, finely
arachnoid-hairy above, finely grayish-tomentose beneath; basal leaves
petiolate, broad-lanceolate, finely crenate-spinose-toothed; cauline leaves
gradually reduced upward, sessile, broadly long-decurrent, oblong-
lanceolate, crenate and finely spinose-toothed. Capitula globose or
globose-ovoid, very finely arachnoid-hairy, 15—17(25) cm wide
(excluding cusps). Involucral bracts numerous, all except inner closely
appressed at base, constricted in middle, and broadened in small,
lanceolate, dorsally carinate horizontal or arcuately decurved appendages

266

261

with two pairs of thin spines along margin; inner bracts lanceolate,
gradually acuminate, thin cuspidate, erect. Receptacular bristles unequal,
weakly scabrous above. Corolla purple; anther tube creamish. Achenes
obpyramidal, almost four-angled, with four small teeth above, 5 mm
long. Flowering VII—VIII; fruiting VIII-IX.

Dry slopes in lower and middle mountain zone, often as weeds.—
Caucasus: Southern Transcaucasia. General distribution: Northern Iran.
Described from the area between towns of Nakhichevan and Kok. Type
in Leningrad.

154. C. freynii Bornm. and Sint. in Russk. Bot. Zhurn. 1 (1911) 4;
Kult. in Fl. Turkm. VII, 204.—Ic.: Bornm. and Sint. op. cit., Plate III.

Perennial (monocarpic ?). Root vertical, woody blackish-fibrous; root
collar thickened. Stem 100—150 cm high, basally covered with numerous
woody remains of old leaves, tomentose, erect, glabrous above, long
paniculately branched; branches glabrous, long, often bent, with solitary
terminal capitula, usually capitula also axillary; stem and branches leafy
mainly in lower half, almost leafless above. Leaves coriaceous, appressed
white-tomentose on both sides; basal leaves petiolate, narrow oblong-
lanceolate, 15-20 cm long, 4 cm wide, pinnatipartite, sinuate with
approximate lobes, lobes 12-15 pairs, deltoid, acuminate-spinescent,
spinose-toothed; cauline leaves like basal but sessile, smaller; uppermost
leaves 1—-1.5 cm long, lanceolate, acuminate-spinescent, divergent.
Capitula flat-cylindrical, 20-25 mm wide (excluding cusps), finely
arachnoid-hairy, somewhat foveate at base. Involucral bracts 50-60, all
except inner basally appressed, slightly narrowed in middle and then
broadened into coriaceous, green, somewhat grooved, deltoid-triangular,
acuminate-spinescent appendage, smooth along margin, 7-10-12 mm
wide at base and 12—20 mm long; inner bracts membranous, straw-
yellow, lanceolate, scabrous along margin, gradually and short thin
acuminate; receptacular bristles smooth. Corolla light purple. Achenes
5—6 mm long, 3 mm wide, indistinctly ribbed; slightly compressed, with
small corona. Flowering VI-—VII; fruiting VII.

Mountain slopes.—Soviet Central Asia: mountainous Turkmenia
(Kopetdag). General distribution: Northern Iran. Described from Sulyukli.
Type in Czechoslovakia.

155. C. beckeri Trautv. in Tr. Peterb. Bot. Sada, IX (1886) 457;
XII, 268; XIV, 212; Kult. in Fl. Turkm. VII, 172.—TIc.: Fl. Turkm. VII,
Plate 26.

Perennial (monocarpic). Root woody, root collar thick, covered with
remains of old leaves. Stem erect, 20-40 cm high, white-tomentose,
simple or sparsely branched; branches one-headed, stem and branches

267

262

densely leafy,’ appearing discontinuously winged with broadly long-
excurrent leaves. Leaves coriaceous, finely arachnoid-hairy above, white-
tomentose beneath; basal leaves oblong, 7-10 cm long, lyrate, lateral
lobes triangular-ovate, decurrent along midrib, acuminate-spinescent,
prickly-toothed; cauline leaves strongly reduced, 2—2.5 cm long, 13-20
mm wide, almost obovate, acuminate-spinescent, prickly-toothed, sessile,
broadly long-decurrent over stem. Capitula globoe, densely arachnoid-
hairy, 20-22 mm wide (excluding cusps; with cusps 55 mm); 35-40
mm long (with florets). Involucral bracts numerous, constricted in middle,
lanceolate and closely appressed at base, with foliaceous, recurved,
carinate, acuminate-spinescent, basally sagittate appendage in upper half,
with 2 opposite teeth; inner bracts with shorter appendage, with shorter
obtuse teeth; innermost bracts finely coriaceous, linear, scabrous along
margin, short acuminate-spinescent. Receptacular bristles almost smooth,
with scarcely noticeable indentation. Corolla whitish. Achenes obovoid,
four-angled, with narrow and thick wings along edges, gray, 5 mm long,
3 mm wide. Flowering V—VI; fruiting VI-—VII.

Mountain slopes.—Soviet Central Asia: mountainous Turkmenia
(Kopetdag). Endemic. Described from Kyzyl-Arvat. Type in Leningrad.

Section 31. Kopetdagia Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 84.—Odontocarpae Winkl. in Tr. Peterb.
Bot. Sada, XII (1892) 253 p. min. p.—Capitula solitary terminal on
stem or branches; involucral bracts lanoeolate, carinate, gradually
attenuate into short cusp; corolla pink or pale-yellow; receptacular bristles
scabrous; achenes four-angled, narrow-winged along edges. Monocarpic
perennial or biennial plants with decurrent cauline leaves.

Type of section: C. smirnowii Trautv.

156. C. smirnowii Trautv. in Tr. Peterb. Bot. Sada, VIII (1883)
488; Winkl. ibid. XI, 136; XII, 259; Kult. in Fl. Turkm. VII, 191—C.
smirnowii var. armata Winkl. op. cit. XI (1890) 136; XII, 259.—C.
smirnowii var. globosa Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIIIc, VII
(1929) 5.—C. russeola Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1,
3 (1936) 319; Kult. in Fl. Turkm. VII, 190.—Arctium smirnowii O.
Ktze. Rev. Gen. 1 (1891) 308.—Ice.: Fl. Turkm. VII, Plate 30, Fig. 1.—
Exs.: H.F.A.M. No. 49.

Perennial (monocarpic). Root woody, root collar with remains of
old leaves. Stem erect, white-tomentose, striate-sulcate, leafy, cymosely

‘branched in upper half. Leaves soft, grayish-green, gray arachnoid-hairy

on both sides; basal leaves in many-leaved rosette, 8 cm long, 2.5 cm
wide, petiolate, with 2 cm long petioles; oblong-lanceolate, pinnatisect,
with deltoid-lanceolate, acuminate-spinescent segments or bipinnatipartite

268

(263

with prickly-toothed lobes; cauline leaves gradually reduced, sessile,
usually decurrent, and then stem appearing discontinuously winged; lower
cauline leaves usually like basal; middle leaves pinnatilobate, with
narrow-triangular or almost lanceolate acuminate-spinescent lateral lobes,
broadly decurrent; upper cauline leaves ovate, crenate-spinose-toothed,
long-decurrent, narrowed into stiff, divergent, short cusp; uppermost
leaves strongly reduced. Capitula solitary terminal on branches, arachnoid
hairy-tomentose, with white or rusty tomentum, 20-28 mm wide
(excluding cusps). Involucral bracts about 70, all except inner lanceolate,
dorsally carinate, appressed at base, deflected almost from middle in
grooved tip; outer bracts almost horizontal or slightly decurved, 15-20
mm long, middle bracts divergent, 22-28 mm long; inner bracts
membranous, erect, narrow-lanceolate, short and thin acuminate, scabrous
along margin, usually pink dorsally, 28-30 mm long; receptacular bristles
scabrous; corolla pink. Achenes 5 mm long, 2.5—3 mm wide, four-angled,
obovoid or narrow-winged along angles, smooth, usually with four teeth
above, dark gray. Flowering VI-VII; fruiting VII—VIII.

In the zone of alpine steppes, montane xerophytes and juniper
forests.—Soviet Central Asia: mountainous Turkmenia (Kopetdag).
General distribution: Iran. Described from the vicinity of Akhal-Teke.
Type in Leningrad.

Note. The plants from Chapandag, described by S.V. Juzepczuk
under the name C. russeola, differ somewhat from the type of C.
semenowii Trautv. by larger, rusty-tomentose capitula and thicker
involucral bracts. At present, we have extensive material of this species
from the town of Chapandag, from which it can be judged that C.
russeola does not deserve recognition; only one species grows in
Kopetdag, which varies somewhat in its characters.

157. C. botschantzevii Juz. ex Tschern. in Bot. Mat. Gerb. Inst.
Bot. Akad. Nauk UzbSSR, XVII (1962) 93.

Biennial. Stem erect, 20-40 cm high, shallow-sulcate, arachnoid-
hairy, discontinuously winged, branched in upper half; branches one-
headed; stem wings 2-4 mm wide, with sparse small teeth. Leaves
grayish-green and scatteredly arachnoid-hairy above, finely gray-
tomentose beneath; basal leaves short-petiolate, lanceolate, pinnatisect
into unequal, narrow-lanceolate, remote lateral lobes; lower cauline leaves
like basal but sessile, decurrent in stem wings; middle leaves oblong,
pinnatipartite, decurrent in stem wings; upper and uppermost leaves short-
lanceolate, with small teeth, short decurrent. Capitula globose, with many
florets, 18 mm wide (excluding cusps), arachnoid-hairy. Involucral bracts
green, almost coriaceous, all squarrose, gradually acuminate into very
short, thin spine; inner and innermost bracts with scarious, lanceolate,

269

264

short acuminate appendages. Receptacular bristles scabrous. Corolla
yellow, exserted. Achenes obovoid, glabrous, glossy, roundish above,
grayish with dark spots. Flowering V; fruiting VI. (Plate XXIII, Fig. 1).

Stony-gravelly slopes in middle mountain zone.—Soviet Central
Asia: Pamiro-Alai (Nuratau Mountains). Endemic. Described from
Nuratau. Type in Tashkent.

Section 32. Helianthae Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2
(1865) 39; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 553.—Capitula
solitary terminal on stem and branches; involucral bracts acuminate in
long, stiff, three-angled cusp; corolla pale yellow; receptacular bristles
smooth. Achenes obovoid, flat-compressed, broad-winged along edges.
Biennial herbs with wingless stem and coriaceous, sessile cauline leaves.

Type of section: C. heliantha Bge.

158. C. spryginii Kult. in Pochv. i Bot.-Geogr. Issl. Bass. mr. Amu-
i Syr-Dari, 2 (1916) 78; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8,
553.—Ie.: Kult. op. cit. No. 24.

Biennial. Stem 20—40 cm high, erect, arachnoid-hairy, striate-sulcate,
branched from base; branches unicapitulate, leafy. Leaves finely
arachnoid-hairy above, gray-tomentose beneath, prickly-toothed; basal
leaves short-petiolate (petioles broad, winged), oblong-obovate, crenate-
lobate, lobes short-acute; lower cauline leaves like basal but sessile;
middle and upper cauline leaves sessile, oblong-obovate, roundish above
with short and thin spine, crenate-toothed, half to two-thirds as long as
basal leaves. Capitula globose, large, 25—35 mm wide (excluding cusps),
densely arachnoid-hairy. Involucral bracts acuminate. into long, stiff,
three-angled spine; outer bracts slightly decurved, 25 mm long, stiff,
carinate; middle bracts squarrose, up to 40 mm long, subulate, carinate;
subsequent bracts somewhat flat, gradually acuminate into spine; inner
bracts coriaceous, linear-lanceolate, acuminate, not exceeding middle
bracts. Receptacular bristles smooth. Corolla pale yellow. Achenes
obovoid, flat, margins with broad wings (one-third or one-fourth of
achene width), not connivent at apex, grayish-green with dark dots, 4.5—
5 mm long and 3 mm wide. Flowering V—VII; fruiting VII—VIII.

On varicolored rock strata in the foothills—Soviet Central Asia:
Pamiro-Alai (southwest). Endemic. Described from the vicinity of Baisun.
Type in Tashkent.

Section 33. Dichotomae Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2
(1865) 8 p. max. p.—Squarrosae Boiss. in Fl. or. III (1875) 460 p. min.
p.—Constrictae Winkl. in Tr. Peterb. Bot. Sada, XII (1892) 264 p. p—
Series Dichotomae Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8 (1940)

270

265

545.—Plants dichotomously branched. Capitula solitary or aggregated
at stem or branch tips; involucral bracts gradually attenuate into thin,
squarrose or almost horizontal, less often almost falcate cusp; corolla
pink, whitish or pale yellow; receptacular bristles smooth or scabrous,
achenes longitudinally ribbed, narrow-winged along ribs, toothed above.
Annual plants with soft, sessile, almost undivided or prickly-toothed
leaves.
Lectotype of section: C. dichotoma Bge.

159. C. dichotoma Bge. in Delect. Sem. Hort. Dorpat. (1845) 3;
Bge. in Mém. Acad. Pétersb. 7 sér. 9,-2, 9; Boiss. Fl. or. HI, 478;
Winkl. in Tr. Peterb. Bot. Sada, XII, 267 p. p.; XIV, 231; Juz. in Tr.
Tadzh. Bazy Akad. Nauk SSSR, 8, 545; Kult. in Fl. Turkm., VII, 166.

Annual. Stem erect, 10-25 cm high, slender, with isolated many-
celled hairs, leafy, dichotomously branched right from base; branches
one-headed. Leaves green on both sides, upper surface with isolated
many-celled hairs scattered throughout, lower surface glandular with
many-celled hairs, sessile, semiamplexicaul, ovate, spinose-toothed.
Capitula 5-6 mm wide (excluding cusps), arachnoid-hairy. Involucral
bracts less numerous, lanceolate, gradually acuminate into thin, erect
cusp; inner and middle bracts green, sparsely glandular, 5-8 mm long,
inner bracts membranous above, short-acuminate and erect; receptacular
bristles white, smooth; corolla pink, eglandular. Achenes obovoid, with
longitudinal narrow-winged ribs, toothed above. Flowering V—VI; fruiting
VI.

Sands in foothill deserts—Soviet Central Asia: Kyzyl-Kum, Amu-
Darya. Endemic. Described from Kyzyl-Kum. Type in Paris; cotype in
Leningrad.

160. C. erectispina Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 94.

Annual. Stem dichotomously branched right from base, 15-30 cm
high, erect, with isolated many-celled hairs and weakly arachnoid-hairy
pubescence, leafy; branches somewhat ascending, long, in turn branched,
one-headed. Leaves green on both sides, almost glabrous above with
isolated glandular and many-celled hairs, glandular beneath with very
thin arachnoid-hairy pubescence and many-celled hairs on midrib, sessile,
semiamplexicaul, lanceolate or oblong-ovate, prickly-toothed. Capitula
ovoid, 7-8 mm wide (excluding cusps), finely arachnoid-hairy. Involucral
bracts sparsely glandular; outer and middle bracts lanceolate, 10-15 mm
long, gradually acuminate in stiff, horizontal or erect, 9-12 mm long
cusp; inner bracts membranous above, short-acuminate; receptacular
bristles white, smooth; corolla pink. Achenes obovoid, longitudinally

272

266

ribbed, with narrow discontinuous wings along nibs, pitted-rugose, spotted,
3-3.5 mm long. Flowering V; fruiting VI.

On sandy takyrs, old fields, saline areas with sandy soil.—sSoviet
Central Asia: Syr-Darya, Kyzyl-Kum, Kara-Kum. Endemic. Described
from Ziadin. Type in Tashkent.

161. C. polytimetica Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 103.

Annual. Stem dichotomously and divaricately profusely branched
from base, with many-celled, flat, white hairs and arachnoid-hairy
pubescence. Leaves coriaceous, green on both sides, glandular above,
glandular and finely arachnoid-hairy beneath; basal leaves short-petiolate,
oblong, crenate-spinose-toothed; cauline leaves sessile, semiamplexicaul,
crenate-toothed, teeth terminating into stiff erect spine. Capitula ovoid,
9-10 mm wide (excluding cusps), finely arachnoid-hairy. Involucral
bracts green, acuminate into stiff, yellow, subulately three-angled spine,
all except inner narrow-lanceolate, dorsally carinate, glandular, erect-
spreading, 12-15 mm long; inner bracts membranous, scabrous along
margin, short and thin acuminate, about 10 mm long; receptacular bristles
smooth, white; corolla pink. Achenes obovoid, longitudinally ribbed,
narrow-winged along ribs, toothed above, 3 mm long. Flowering VI;
fruiting VII.

In the foothills and high deserts—Soviet Central Asia: Pamiro-Alai
(along the Zeravshan River in the foothills of Nuratau). Endemic.
Described from the foothills of the Nuratau Range, Tutak-Sai. Type in
Leningrad.

162. C. patentispina Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 101.—Exs.: GRF No. 2777 (sub. C.
bungeana Rgl. and Schm.).

Annual. Stem erect, 20-30 cm high, white, with isolated glandular
and many-celled hairs, leafy, dichotomously branched from base;
branches one-headed. Leaves green on both sides, glabrous above with
occasional many-celled hairs, glandular beneath with very thin arachnoid-
hairy pubescence and many-celled hairs along midrib; basal leaves
apparently petiolate; cauline leaves sessile, amplexicaul, oblong-
lanceolate, prickly-toothed along margin. Capitula 7-8 mm wide
(excluding cusps), finely arachnoid-hairy. Involucral bracts green, very
sparsely glandular; outer and middle bracts lanceolate, 8-11 mm long,
acuminate in somewhat decurved stiff cusp; inner bracts membranous
above, erect, short-acuminate; receptacular bristles white, smooth; corolla
pink. Achenes obovoid, longitudinally ribbed, with very narrow
discontinuous wings, pitted-rugose. Flowering V; fruiting VI.

267

Plate XX.
1—Cousinia ulotoma Bornm.; 2—C. decurrentifolia Juz.

271

273

268

On takyrs.—Soviet Central Asia: Syr-Darya, Kyzyl-Kum, Kara-Kum.
Endemic. Described from Mubarek. Type in Tashkent.

163. C. sylvicola Bge. in Delect. Sem. Hort. Dorpat. (1845) 3;
Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2, 9; Juz. in Tr. Tadzh. Bazy
Akad. Nauk SSSR, VIII, 545.—C. dichotoma auct. non Bge.: Winkl. in
Tr. Peterb. Bot. Sada, XII (1892) 263 p. p.; Bornm. in Beih. bot.
Centralbl. XXXIV, 2, 194.

Annual. Stem erect, 7-10 cm high, slender, with isolated many-
celled hairs, leafy, branched above; branches short, one-headed. Leaves
finely coriaceous, green and glabrous above; grayish-green and finely
arachnoid-hairy beneath; basal leaves petiolate, oblong-lanceolate,
crenate-toothed, teeth acuminate into thin and short (1—-1.5 mm long)
spines; cauline leaves sessile, somewhat amplexicaul, lanceolate, short
prickly-toothed, gradually reduced toward stem apex. Capitula 5-6 mm
wide (excluding cusps), finely arachnoid-hairy. Involucral bracts green,
somewhat flat; outer and middle bracts 8-9 mm long; lanceolate, with
falcately decurved cusp, inner bracts lanceolate, membranous above,
short-acuminate, erect; receptacular bristles smooth, white; corolla
creamish (?). Achenes three-angled, pitted, with teeth above.

On sands.—Soviet Central Asia: Amu-Darya, Kyzyl-Kum (?).
Endemic. Described from the area between the Kuvan and Yandarya
rivers. Type in Paris.

Note. A little known species; additional material is necessary in
order to compare it with all members of the section.

164. C. bungeana Rgl. and Schmalh. in Tr. Peterb. Bot. Sada, VI
(1880) 318; Winkl. ibid. XII, 267 p. p.; Juz. in Tr. Tadzh. Bazy Akad.
Nauk SSSR, 8, 545.—Arctium bungeanum O. Ktze. Rev. Gen. 1 (1891)
307.

Annual. Stem spreadingly branched from base, 25—30 cm high, white,
with numerous thick hairs and arachnoid-hairy pubescence, often
glandular below capitulum, leafy; branches one-headed. Leaves thin-
coriaceous, green, very finely and sparsely arachnoid-hairy, glandular
above, grayish-green and finely arachnoid hairy-tomentose beneath; basal
leaves usually short-petiolate, lanceolate; cauline leaves ovate or
sometimes orbicular, sessile, auriculate, prickly crenate-toothed. Capitula
13-14 mm wide (excluding cusps), finely arachnoid-hairy. Involucra]
bracts green, densely glandular; outer and middle bracts lanceolate,
squarrose, dorsally carinate, wide, up to 2 mm wide at base; inner bracts
linear, membranous at apex, short-acuminate; receptacular bristles
smooth, white; corolla creamish or pale yellow, eglandular. Achenes
obovoid, 3 mm long, almost three-angled, longitudinally mbbed, with

274

269

narrow discontinuous wings and small pits. Flowering V—VI; fruiting
VI.

On sandy, loamy-gravelly substrates—Soviet Central Asia: Syr-
Darya, Tien Shan (western foothills). Endemic. Described from the
vicinity of Leninabad (Khodzhent). Type in Leningrad.

Note. The species varies greatly in the glandular cover of the leaves
and involucral bracts; this character is very important in this group of
species and deserves further study, in the first instance in the populations.

165. C. pusilla Winkl. in Tr. Peterb. Bot. Sada, X (1889) 468; XII,
197; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, VIII, 545.—Arctium
pusillum O. Ktze. Rev. Gen. 1 (1891) 308.

Annual. Stem erect, 8—15(25) cm high, white, arachnoid-hairy leafy,
sparsely branched in upper half; branches short, one-headed. Leaf
coriaceous, green and glabrous above, sparsely grayish-tomentose
beneath, broad-lanceolate, sessile (except lower short petiolate), with
small teeth, teeth acuminate into thin, short, 1-2 mm long spine. Capitula
broadly ovoid, 7-9 mm wide (excluding cusps), arachnoid-hairy.
Involucral bracts green; outer and middle bracts lanceolate, 7-10 mm
long, without glands, acuminate into horizontal thin cusp; inner bracts
membranous, short-acuminate, usually purple; receptacular bristles white,
smooth; corolla pink, eglandular. Achenes obovoid, 3 mm long, almost
three-angled, longitudinally nbbed, with narrow discontinuous wings and
small spots, pitted-rugose. Flowering V; fruiting V—VII.

Saline sandy loams, in the foothills—Soviet Central Asia: Pamiro-
Alai (south). Endemic. Described from the vicinity of Dzhalikul on the
left bank of the Vakhsh River. Type in Leningrad.

166. C. orthacantha Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 98.

Annual. Stem divaricately branched from base, 15-20 cm high,
white, sparsely glandular, with thick many-celled hairs and arachnoid-
hairy pubescence, leafy; branches somewhat ascending, leafy, in turn
profusely branched with numerous short branchlets. Leaves coriaceous,
grayish-green, sparsely sessile glandular-hairy, and finely arachnoid-hairy
above, sparsely white-tomentose beneath with prominent veins, large-
toothed, teeth acuminate into stiff, 3-5 mm long spine; basal leaves
oblong-lanceolate, short-petiolate; cauline leaves ovate, sessile, weakly
amplexicaul. Capitula broadly ovoid, 10-12 mm wide (excluding cusps),
arachnoid-hairy. Involucral bracts green; outer and middle bracts
divergent, 11-14 mm long, narrow-lanceolate, carinate dorsally,
glandular, acuminate in short cusp; inner bracts lanceolate, 13 mm
long, membranous, white, short acuminate; receptacular bristles smooth;

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270

corolla creamish (?) or light pink. Achenes obovoid, 3—3.5 mm long,
longitudinally ribbed with very narrow undivided wings along ribs.
Flowering V; fruiting VI.

On stable sands.—Soviet Central Asia: Pamiro-Alai (south).
Endemic. Described from the Kafirnigan River Valley near Tshkan-Tepe.
Type in Tashkent.

167. C. tedshenica Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 105.

Annual. Stem divaricately branched almost from base, 10—20 cm
high, whitish, indistinctly striate-sulcate with many-celled, somewhat
flat, flexuous hairs. Leaves thin-coriaceous, green above with isolated
many-celled hairs scattered throughout, finely arachnoid-hairy beneath
and also with many-celled hairs; basal leaves oblong, short-petiolate,
crenate-toothed, teeth acuminate in thin spine; cauline leaves sessile,
semiamplexicaul, reduced toward stem and branch apices. Capitula broad
cylindrical, 7-10 mm wide (excluding cusps), very finely arachnoid-
hairy, usually foveate at base. Involucral bracts numerous, green, all
except inner ovate, closely appressed, free in middle, very sharply
decurved, acuminate in short cusp; inner bracts membranous, broad-
lanceolate, weakly scabrous along margin, 10-12 mm long, with short
thin cusp, 2—2.5 mm wide in upper broadened part; receptacular bristles
unequal, longest slightly scabrous. Corolla creamish (?). Achenes
obovoid, three-angled, 4 mm long, narrow-winged along edges, pitted-
rugose. Flowering IV, fruiting V.

Stable sands and outcrops of bedrocks in deserts——Soviet Central
Asia: mountainous Turkmenia (Badkhyz), Kara-Kum. Endemic. Described
from the vicinity of Lake Er-Oilan-Duz. Type in Tashkent.

168. C. murgabica Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 97.—C. pygmaea auct. non Winkl.; Kult.
in Fl. Turkm. VII (1960) 164.—C. bungeana auct. non Winkl.: Kult.
ibid. 168.

Annual. Stem 20—25 cm high, weakly and repeatedly branched from
base, whitish, cylindrical, with many-celled hairs. Leaves thin-coriaceous,
green above, sparsely glandular, finely arachnoid-hairy beneath, very
slightly reduced toward stem and branch apices; cauline leaves sessile,
semiamplexicaul, crenate-toothed, with teeth acuminate into thin erect
spine. Capitula globose-pyramidal, 9-11 mm, wide (excluding cusps),
arachnoid-hairy. Involucral bracts numerous, all except inner ovate to
oblong-ovate, free in middle, narrow-lanceolate, about 10 mm long,
gradually acuminate into thin subulate, horizontal cusp; inner bracts
membranous, narrow-lanceolate, 1—1.5 mm wide, scabrous along margin,

276

271

acuminate into thin, short, erect cusp; receptacular bristles unequal,
weakly scabrous; corolla whitish or pale pink (?). Achenes obovoid,
angular, narrow-winged, toothed, 3—3.5 mm long. Flowering V~ fruiting
V.

On sands.—Soviet Central Asia: Kara-Kum. Endemic. Described
from the sands near the village of Sary-Yazy. Type in Leningrad.

169. C. angusticeps Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser.
1, 3 (1936) 323; Kult. in Fl. Turkm. VII, 167.

Annual. Stem erect, somewhat arachnoid-hairy, glossy. Leaves
sessile, ovate-oblong, almost undivided, spinescent, erose-toothed, green
on both sides, somewhat arachnoid-hairy-flocculose beneath. Capitula
12-13 mm long (with florets) and 4 mm wide (excluding cusps), ovoid-
oblong, roundish below, aggregated at stem apices. Involucral bracts
(about 30) pale green, somewhat arachnoid-hairy; outer bracts 2 mm
long and 0.5 mm wide at base; middle bracts appressed at base,
lanceolate, 5 mm long and 1 mm wide, attenuate into thin 1.5 mm long
divergent spine at apex; inner bracts 11 mm long and 1 mm wide,
linear, membranous, glossy, gradually acuminate, toothed, whitish;
receptacular bristles weakly scabrous; corolla yellowish; anther tube pink.
Achenes 4 mm long and 1.5 mm wide, oblong-pyramidal, pitted-rugose,
toothed above. Flowering IV—V; fruiting V. (Plate X, Fig. 1).

Sandy and clayey deserts.—Soviet Central Asia: Kara-Kum,
mountainous Turkmenia (Kopetdag). Endemic. Described from Kara-
Kum between the village of Annenkovo and the Kara-Kuyu well. Type
in Leningrad.

170. C. eugenii Kult. in Bot. Mat. Gerb. Glavn. Bot. Sada, V (1924)
141; Kult. in Fl. Turkm. VI, 164.

Annual. Stem dichotomously branched from base, forming
hemispherical shrub, 6-10 cm high, whitish, flocculose-arachnoid-hairy,
leafy. Leaves coriaceous, sessile, attenuate toward both ends, broad-
lanceolate, acuminate into thin spine, crenate-toothed, teeth acuminate
into 6-10 mm long, thin spine, green and finely arachnoid-hairy above,
glaucous and finely tomentose beneath. Capitula small, 7 mm wide
(excluding cusps), broadly ovoid, finely and appressed pubescent.
Involucral bracts numerous, all equal, narrow lanceolate, gradually
acuminate in long, weakly uncinate, thin spine, deflexed only at tip;
outer bracts 5—6 mm, middle up to 10 mm, inner up to 12 mm long,
purple-tipped. Corolla purple. Receptacular bristles smooth. Achenes
narrow, obpyramidal, almost four-angled, 3.5 mm long and 1 mm wide,
narrow-winged, rugose-pitted, toothed above. Flowering V; fruiting VI—
VI.

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272

On semistable sands.—Soviet Central Asia: Kara-Kum, Amu-Darya,
Kyzyl-Kum, mountainous Turkmenia (Badkhyz). Endemic. Described
from region between Karshi and Kagan. Type in Tashkent.

Section 34. Xiphiolepides Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2
(1865) 38 p. p—Serratuloideae Bge. in Boiss. Fl. or. III (1875) 459 p.
p.—Constrictae Winkl. in Tr. Peterb. Bot. Sada, XII (1892) 264 p. p—
Capitula solitary terminal on stem and branches; involucral bracts
numerous, carinate, gradually acuminate; corolla whitish; receptacular
bristles smooth; achenes obovoid, without pappus.. Biennial plants with
many rows of wings on stem, and cauline leaves decurrent into wings.

Lectotype of section: C. xiphiolepis Boiss.

171. C. hystrix C.A.M. in Ind. sem. hort. Petrop. 2 (1835) 34; DC.
Prodr. VI, 554; Ldb. Fl. Ross. II, 677; Bge. in Mém. Acad. Pétersb. 7
sér. 9, 2, 38; Winkl. in Tr. Peterb. Bot. Sada, XII, 267; Boiss. FI. or.
Ill, 466; Bornm. in Vestn. Tifl. Bot. Sada, 30, 19, 24; Grossh. Fl. Kavk.,
IV, 167; Takht. in Tr. Arm. Fil. Akad. Nauk SSSR, Biol. Ser. 2, 177;
Fl. Azerb. VIII, 364.—Carthamus pterocaulos C.A.M. Verz. (1831) 67.—
Arctium hystrix O. Ktze. Rev. Gen. 1 (1891) 308.

Annual. Stem erect, white arachnoid-hairy, winged throughout, wings
in many rows, crenate-prickly-toothed. Leaves coriaceous, gray
arachnoid-hairy on both sides, more densely beneath; basal leaves
petiolate, oblong-lanceolate, pinnate-prickly lobed, with narrow triangular,
spinescent lobes; cauline leaves gradually reduced upward, sessile,
decurrent in wings, like basal but usually with more approximate lobes,
not in same plane, as a result leaf margin appearing spinose. Capitula
globose-pyramidal, 25 mm wide (excluding cusps), finely arachnoid-
hairy. Involucral bracts numerous, erect or outermost bracts almost
horizontal, narrow-lanceolate, weakly carinate, gradually attenuate in
short cusp, 20-30 mm long; inner bracts membranous above, gradually
acuminate into subacute cusp, dorsally scabrous in middle; receptacular
bristles smooth. Corolla whitish; anther tube pink. Achenes obovoid, 4—
4.5 mm long, with scarcely noticeable longitudinal stripes and short
corona above, without pappus. Flowering VII—VIII; fruiting VIIJ-—IX.
(Plate XXII, Fig. 2).

Gravelly dry mountain slopes in middle mountain zone.—Caucasus:
Eastern Transcaucasia, Talysh. General distribution: Northwestern Iran.
Described from Talysh. Type in Leningrad.

Section 35. Pugioniferae Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2
(1865) 30 p. p.—Calcitrapae Boiss. Fl. or. HI (1875) 460 p. p.—
Xiphaeanthae Winkl. in Tr. Peterb. Bot. Sada, XII (1892) 232 p. p.— ©

213

Macropterae Takht. in Tr. Arm. Fil. Akad. Nauk SSSR, Biol. Ser. 2
(1937) 189.—Capitula solitary terminal on stem and branches; involucral
bracts less numerous, closely appressed at base, ovate, abruptly attenuate
in stiff carinate cusp; corolla purple; receptacular bristles smooth; achenes
obovoid, longitudinally ribbed. Biennial plants with discontinuously
winged stems and long-decurrent cauline leaves.

Lectotype of section: C. pugionifera Spach.

172. C. macroptera C.A.M. in DC. Prodr. VI (1837) 555; Ldb. FI.
Ross. II, 679; Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2, 32; Boiss. FI. or.
Ill, 492; Winkl. in Tr. Peterb. Bot. Sada, XII, 234; Grossh. Fl. Kavk.
IV, 165; Takht. in Tr. Arm. Fil. Akad. Nauk SSSR, Biol. Ser. 2, 190;
Takht. in Fl. Erev. 318; Fl. Gruzii, VIII, 443; Fl. Azerb., VIII, 370.—C.
obovata Boiss. Fl. or. III (1875) 491.—C. macroptera var. obovata
Winkl. in Tr. Peterb. Bot. Sada, XII (1892) 235.—Arctium macropterum
O. Ktze. Rev. Gen. 1 (1891) 308.—A. obovatum O. Ktze. Ibid—Cousinia
carlinoides auct. non DC.: Boiss. and Buhse in Nouv. Mém. Soc. Nat.
Mosc. XII (1860) 126; Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2, 32
(quoad Boiss. and Buhse).—Ic.: Fl. Azerb., VIII, Plate 36.

Biennial. Stem erect, 15-40 cm high, white arachnoid-hairy,
spreadingly branched from middle, forming hemispherical inflorescence;
stem and branches densely leafy, discontinuously winged with long and
broadly excurrent leaves. Leaves coriaceous, grayish green and finely
arachnoid-hairy above, white-tomentose beneath, with network of
prominent veins; basal leaves short-petiolate, obovate-orbicular or ovate,
crenate-prickly-toothed or shallow lobate; cauline leaves gradually
reduced, sessile, broadly long-decurrent, lanceolate or oblong-lanceolate,
finely prickly-toothed, acuminate-spinescent. Capitula globose-ovoid, 8—
9 mm wide (excluding cusps), about 20 mm long, corolla erect, finely
arachnoid-hairy basally truncate. Involucral bracts less numerous, 25-
30, abruptly attenuate towards apex into divergent or almost horizontal,
short, carinate cusp; inner bracts lanceolate, purple in middle, transparent-
scarious above, orbicular, erect; receptacular bristles smooth. Corolla
purple; anther tube purple. Achenes narrow-obovoid, longitudinally
ribbed, transversely rugose, with small toothed above, about 4 mm long,
dark. Flowering V—VI; fruiting VI-—VII.

Dry stony slopes in lower mountain zone.—Caucasus: Southern
Transcaucasia. General distribution: Asia Minor (Turkey), Iran. Described
from Nakhichevan. Type in Leningrad.

Section 36. Microcarpae Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2
(1865) 43.—Alpinae Bge. Ibid. (1865) 45 p. p.—Xiphiolepides Bge. Ibid.
(1865) 38 p. p.—Psilacanthae Boiss. Fl. or. III (1875) 461.—

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274

Serratuloideae Boiss. Ibid. (1875) 459 p. p.—Lampocarpae Winkl. in
Tr. Peterb. Bot. Sada, XII (1892) 242.—Odontocarpae Winkl. Ibid.
(1892) 253 p. p.—Homalochatae Winkl. Ibid. (1892) 200 p. p.—
Microcarpa Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8 (1940) 514,
553.—Capitula solitary terminal on stem or branches; involucral bracts
either almost linear, acuminate in thin stiff cusps, or almost lanceolate,
acuminate in short, stiff, triangular cusp; corolla purple, whitish, or
yellow; receptacular bristles scabrous, less often smooth; achenes
glabrous, smooth, less often almost three-angled. Biennial, less often
perennial or monocarpic herbs, with wingless or winged stems and sessile,
less often short-decurrent cauline leaves.
Lectotype of section: C. microcarpa Boiss.

Subsection 1. Coronophora Juz.—Involucral bracts numerous,
gradually acuminate in thin stiff cusp; inner bracts with membranous
appendages, forming “collar” or “corona”; achenes glabrous, smooth;
receptacular bristles scabrous. Biennial or perennial plants with wingless
stem.

173. C. radians Bge. in Delect. Sem. Hort. Dorpat. (1845) 3; Bge.
in Mém. Acad. Pétersb. 7 sér. 9, 2, 44; Boiss. Fl. or. III, 495; Winkl. in
Tr. Peterb. Bot. Sada, XII, 253; XIV, 207; Bornm. in Beih. bot. Centralbl.
XXXIV, 2, 172; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR 8, 553.—C.
stenolepis Rgl. and Schmalh. in Izv. Obshch. Lyubit Estestv. Antrop. 1
Etn. 34, 2 (1882) 49; Winkl. in Tr. Peterb. Bot. Sada, XII, 252; XIV,
207.—C. interrupta Winkl. Ibid. XII (1892) 229.—C. coronata auct.
non Franch.: Winkl. Ibid. XIV (1897) 229 p. p.—Arctium radians O.
Ktze. Rev. Gen. | (1891) 307.—A. stenolepis O. Ktze. Ibid. (1891)
308.—Exs.: GRF No. 1423 (sub. C. stenolepide Rgl. and Schmalh.).

Perennial or biennial. Stems 30-60 cm high, erect, partly arachnoid-
hairy, glabrescent, weakly branched, usually from middle; branches one-
headed, leafy. Leaves coriaceous, relatively finely spinose-toothed, green
and glabrous above, white-tomentose beneath; basal leaves short-petiolate,
lyrate, with ovately acuminate terminal lobe and ovate lateral lobes or
pinnatisect; lower cauline leaves like basal but sessile; middle leaves
oblong-lobate, sessile, auriculate; upper leaves oblong, acuminate-
spinescent with short spine, one-third to half as long as middle leaves,
sessile, auriculate; uppermost leaves strongly reduced. Capitula ovoid-
globose, 15-20 mm wide (excluding cusps), weakly arachnoid-hairy.
Involucral bracts numerous, stiff, 1-1.2 mm wide at base, with subulate
cusp, as wide as thick, green, erect-spreading; outermost bracts 5S—15
mm long and outer 18-20 mm long, both not exceeding middle bracts;
middle bracts coriacous, carinate, 20-22 mm long; inner bracts with

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membranous, oblong-lanceolate, usually pale purple appendages, forming
corona, exceeding adjacent middle bracts by 20 mm; receptacular bristles
scabrous. Corolla pink, not exserted from collar. Achenes oblong-obovoid,
grayish, indistinctly three-angled. Flowering VI—VII; fruiting VII—VIII.

Clayey and gravelly slopes in middle mountain zone.—Soviet Central
Asia: Pamiro-Alai. General distribution: Iran. Described from the
Zeravshan Basin, near Samarkand. Type in Paris.

174. C. coronata Franch. in Ann. Sc. Nat. Bot. 6 sér. 16 (1883)
321; Winkl. in Tr. Peterb. Bot. Sada, XIV, 229 p. p—C. campylacantha
Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3 (1936) 312; Juz. in
Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 554.—C. radians B. paradoxa
Bornm. in Beih. bot. Centralbl. XXXIV.—2 (1916) 178.—Arctium
coronatum O. Ktze. Rev. Gen. | (1891) 307.—Ie.: Franch. op. cit. tab.
18.

Perennial or biennial. Stem 25—50 cm high, erect, sparsely arachnoid-
hairy, striate-sulcate, weakly branched from middle or base; branches
one-headed, leafy. Leaves coriaceous, finely spinose-toothed, green and
glabrous above, finely gray-tomentose beneath; basal leaves short-
petiolate, lyrate, with ovately acuminate terminal lobe or pinnatisect;
lower cauline leaves like basal but sessile; middle leaves lobed, lobes
acuminate-spinescent; upper leaves oblong, shorter than middle;
uppermost leaves reduced. Capitula ovoid-globose, 18-22 mm wide
(excluding cusps), weakly arachnoid-hairy. Involucral bracts stiff,
carinate, with short cusp, green, wider than in previous species, 2 mm
wide at base, erect, weakly arcuately decurved; outermost bracts half as
long as outer; outer bracts 15 mm long; middle bracts coriaceous,
carinate, slightly flat above, with short cusp; inner bracts with
membranous broadly lanceolate appendages, 5-6 mm wide, forming
corona, exceeding middle bracts by 15-20 mm; receptacular bristles
scabrous. Corolla pink, not exserted from collar. Achenes obovoid, light
brown. Flowering VI-VII; fruiting VII-VIII.

Loamy-gravelly and stony slopes in middle mountain zone.—Soviet
Central Asia: Pamiro-Alai. Endemic. Described from the vicinity of
Unrmitan. Type in Paris.

Note. C. radians Bge. and C. coronata Franch. are quite distinct in
the extreme forms, but in the contact regions of their ranges transitional
forms are produced probably of hybrid origin, which are very difficult
to distinguish. We see such a picture, for example, in the vicinity of the
village of Urgut from where numerous specimens are represented mainly
by transitional forms. We see such abundance of transitional forms also
in the Tupalang River Valley.

Numerous hybrids of C. coronata and C. microcarpa Boiss.,

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described under the name C. atripurpurea Juz. (cf. note to C. microcarpa
Boiss.), are known.

175. C. pulchra Winkl. in Tr. Peterb. Bot. Sada, X (1889) 478;
XII, 252; XIV, 207; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8,
554.—C. subtilis Juz. Ibid. (1940) 559, quoad typum.—C.
pseudostenolepis Rech. f. and Edelbg. in Dan. Biol. Skr. 8, 2 (1955)
147, fig. 99 and 103E.—Arctium pulchrum O. Ktze. Rev. Gen. 1 (1891)
308.—Ic.: Rech. f. l.c.

Perennial or biennial. Stem 25-50 cm high, erect, partly arachnoid-
hairy, glabrescent, branched from base or middle; branches one-headed,
leafy. Leaves finely spinose-toothed, gray-tomentose beneath, green and
glabrous above; basal leaves petiolate, lyrate, with ovately acuminate
terminal lobe; lower cauline leaves like basal, sessile or short-petiolate
with narrow-winged petioles; middle cauline leaves sessile, lyrate, with
one or two pairs of lateral lobes or oblong, sessile, semiamplexicaul;
upper leaves oblong, acuminate; uppermost leaves strongly reduced.
Capitula ovoid-oblong, 8-12 mm wide (excluding cusps), somewhat
arachnoid-hairy. Involucral bracts stiff, carinate, with subulate cusp, 1—
1.5 mm wide at base, green, somewhat horizontal; outer bracts as long
as or longer than capitulum diameter (considering only divergent part
without appressed base); middle bracts coriaceous, with short cusp, shorter
than outer bracts, 1-2 mm wide at base; inner bracts with membranous,
purple, lanceolate, up to 1 cm long and 2 mm wide appendage forming
a collar and two times as long as adjacent middle bracts; receptacular
bristles scabrous above. Corolla pink, slightly exserted. Achenes obovoid,
almost three-angled, spotted, about 5—5.5 mm long. Flowering VI-VII;
fruiting VII—VIII.

Stony slopes on the outcrops of red sandstones in middle mountain
zone.—Soviet Central Asia: Pamiro-Alai (Babatag, Gazimailik, Gardani-
Ushti, Khodzha-Kozian, Koi-Pioz-Tau, Sarsaryak, mountain slopes along
Pyandzh). General distribution: Afghanistan. Described from the
Gazimailik Mountains above Khoshbulak. Type in Leningrad.

Note. The species is quite polymorphic. Plants with involucral bracts
shorter than in typical C. pulchra are found in Babatag. We find plants
with longer leaves and capitula in the eastern part of the range in southern
Tadzhikistan. It is possible that such variability can be explained by
hybridization between closely related species, or perhaps that these are
small geographical races, which quite likely will be considered as separate
narrowly localized species after further study of extensive material.

While studying the variability in C. pulchra, we came to the
conclusion that it is difficult to distinguish the type of C. subtilis from
C. pulchra, in any case from its eastern members, and we refer this

A |

| Wi iy Wy yy

SZ
Fie

281 Plate XXI.

1—Cousinia lyrata Bge.; 2—C. lasiosiphon Juz.

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278

species to the synonyms of C. pulchra. Cf. also the note to C. alpina
Bge.

176. C. alpestris Bornm. ex Juz. in Tr. Tadzh. Bazy Akad. Nauk
SSSR, 8 (1940) 554.—C. radians y. alpestris Bornm. in Beith. bot.
Centralbl. XXXIV, 2 (1916) 178.

Biennial or perennial. Stem 35-60 cm high, erect, arachnoid-hairy,
glabrescent, cymosely branched in upper part or almost from base;
branches one-headed, leafy. Leaves green and very finely arachnoid-
hairy above, gray-tomentose beneath; basal leaves petiolate, almost
bipinnatisect into spinose-toothed narrow-lanceolate lateral segments;
lower. cauline leaves like basal, other cauline leaves sessile, oblong,
spinose-lobate, gradually reduced toward stem apex. Capitula ovoid-
globose, very finely arachnoid hairy, 12-14 mm wide (excluding cusps);
involucral bracts numerous, stiff, carinate, with subulate cusp; outer bracts
decurved, almost half as long as middle; middle bracts almost horizontal,
almost as long as or somewhat longer than capitulum diameter;
subsequent bracts smaller, flat, 10-12 mm long, short-acuminate, erect;
inner bracts with membranous, broad-lanceolate, 10-12 mm long, 2-3
mm wide appendages forming corona; receptacular bristles apically
broadened, scabrous. Corolla pale pink or whitish. Achenes obovoid,
glabrous, smooth, glossy, 3.5—4 mm long. Flowering VIJ—VIII.

Stony-gravelly slopes in upper mountain zone.—Soviet Central Asia:
Pamiro-Alai. Endemic. Described from the Yagnob River Basin near
the village of Novobad. Type in Leningrad.

Note. Hybrids of C. alpestris Bornm. and C. submutica Franch.,
described as C. heteromorpha Bornm. (B.B.C. 34, 2 (1916) 145), are
known from Novobad (Yagnob River Basin). These plants have almost
lyrate basal and lower cauline leaves, ovoid-globose capitula with a
somewhat foveate base arid involucral bracts with a more or less arcuately
decurved, short, thin cusp, the inner bracts with membranous appendages
usually as long as the corolla.

177. C. mulgediifolia Bornm. in Beih. bot. Centralbl. XXXIV, 2
(1916) 139; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 554.

Biennial or perennial. Stem 45—100 cm high, erect, almost glabrous,
usually branched from base, leafy; branches in turn branched, branchlets
one-headed. Leavés coriaceous, glabrous and green above with network
of prominent veins, white-tomentose beneath, sparsely very finely
spinose-toothed or almost entire; basal leaves oblong-ovate, attenuate
toward both ends, petiolate, petioles almost as long as lamina, or less
often almost lyrate, with large terminal lobe and one or two pairs of
small lateral lobes; lower cauline leaves like basal or almost sessile,

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279.

semiamplexicaul; middle leaves oblong, sessile, semiamplexicaul, with
wide auricles; upper leaves gradually reduced, almost ovate, sessile,
semiamplexicaul, acuminate. Capitula ovoid-globose, 12-15 mm wide
(excluding cusps), very finely arachnoid-hairy. Involucral bracts
numerous, with subulate, squarrose cusp, about 1 mm wide at base;
outer and middle bracts not exceeding capitulum diameter, 10-12 mm
long; inner bracts with membranous, lanceolate, usually pale purple
appendages, forming corona, exceeding adjacent involucral bracts by 8—
10 mm; receptacular bristles scabrous. Corolla pale purple or whitish,
slightly exserted from corona. Achenes oblong-obovoid, glabrous, smooth,
about 3.5 mm long, light gray. Flowering VII—VIII; fruiting VIII.

Stony slopes in middle mountain zone—Soviet Central Asia: Pamiro-
Alai (south). Endemic. Described from the vicinity of Charga (former
Shuroabad District). Type in Leningrad.

178. C. leptocampyla Bornm. in Beih. bot. Centralbl. XXXIV, 2
(1916) 179; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 554.

Biennial or perennial. Stem erect, 70-80 cm high, very finely
arachnoid-hairy, sparsely branched from base; branches in turn branched
into slender, long, leafy, one-headed branchlets. Leaves coriaceous, finely
arachnoid-hairy above, white-tomentose beneath, finely spinose-toothed;
middle leaves sessile, crenate-lobed, with large, about 4 cm long, terminal
lobe, usually finely spinose-toothed and with three or four pairs of almost
entire, acuminate-spinescent, 1-1.5 cm long lateral lobes; upper leaves
oblong-ovate, semiamplexicaul, with wide auricles, finely spinose-
toothed, acuminate; leaves on branches small, sessile, spinose-toothed.
Capitula numerous, globose, 10-13 mm wide (excluding cusps), finely
arachnoid-hairy. Involucral bracts numerous; outer and middle bracts
linear-subulate, rather weak, almost arcuately decurved, with somewhat
short cusps, 3—12 mm long; inner bracts membranous, flat, with whitish,
lanceolate appendages above, exceeding adjacent bracts by 8-10 mm,
forming corona; receptacular bristles scabrous. Corolla pale-yellow or
almost white. Achenes obovoid, glabrous, smooth, about 3 mm long.
Flowering VII-VII; fruiting VIII.

Stony-gravelly slopes in middle mountain zone.—Soviet Central
Asia: Pamiro-Alai (south). Endemic. Described from the vicinity of
Faizabad (central Tadzhikistan). Type in Leningrad.

Note. A little known species that, except for the type specimen, no
one has collected. It is morphologically very unique and stands out
prominently among the closely related species.

Subsection 2. Carduncellus Juz.—lInvolucral bracts relatively less
numerous, gradually acuminate in cusp, inner bracts with or without

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“crown” or “corona”; achenes glabrous, smooth or almost three-angled;
receptacular bristles scabrous or smooth. Biennial or perennial plants
with wingless stems.

Note. The division of the subsection into series will be possible
only after monographic treatment of the genus.

179. C. alpina Bge. in Delect. Sem. Hort. Dorpat. (1845) 3; Bge.
in Mém. Acad. Pétersb. 7 sér. 9, 2, 46; Boiss. FI. or. III, 495;-Winkl. in
Tr. Peterb. Bot. Sada, XII, 249; XIV, 229.—C. acicularis Franch. in
Ann. Sc. Nat. Bot. 6 sér. 16 (1883) 319; Winkl. op. cit. XII, 252; Bornm.
in Beih. bot. Centralbl. XXXIV, 2, 170.—C. lyratifolia Winkl. op. cit.
X (1887) 90; XII, 249; XIV, 229; Bornm. op. cit. 170.—C. subtilis Juz.
in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8 (1940) 559, quoad specimina
examinata.—Arctium alpinum O. Ktze. Rev. Gen. 1 (1891) 307.—A.
aciculare O. Ktze. Ibid.

Biennial. Stem erect, 30-50 cm high, striate-sulcate, finely arachnoid-

“hairy, leafy almost from base, cymosely branched almost from middle;

branches long, leafy, one-headed. Leaves coriaceous, greenish-gray and
finely arachnoid-hairy above, finely gray-tomentose beneath; basal leaves
short-petiolate, oblong, pinnatipartite into triangular, uncinate, lateral
lobes acuminate in stiff cusp, or lyrate with large ovate, terminal lobe;
cauline leaves gradually reduced, sessile, short-decurrent; lower cauline
leaves like basal; other cauline leaves lanceolate, acuminate in stiff cusp,
spinose-toothed. Capitula ovoid-globose, 10-15 mm wide (excluding
cusps), white or rusty-tomentose. Involucral bracts about 80; outer bracts
narrow-subulate-triquetrous, decurved, 8-12 mm long; middle bracts
appressed at base, gradually attenuate in stiff erect, carinate, triquetrous-
subulate, 20-35 mm long, less often 10-12 mm long cusp; inner bracts
thin-coriaceous, lanceolate, somewhat attenuate above, usually brownish
dorsally, 18-22 mm long; receptacular bristles scabrous. Corolla purple;
anther tube pink. Achenes obovoid, 4 mm long, glabrous, smooth,
roundish above. Flowering VIIJ—VIII; fruiting VIII-IX.

Loamy-stony slopes in upper part of the middle or upper mountain
zone.—Soviet Central Asia: Pamiro-Alai (Turkestan, Gissar, Zeravshan,
Peter the Great ranges, and mountains of southern Tadzhikistan).
Endemic. Described from the Karatau Range (Lehmann). Type in Paris.

Note. The species is extremely variable in capitulum size and length
of involucral bracts. Apparently, it hybridizes easily with C. submutica
Franch. and other species of section Microcarpae, which possibly explains
the variety of forms.

In describing C. subtilis Juz., S.V. Juzepczuk, as is evident from
the specimens labeled by him and the characters listed in the key for
identification, had in mind the plants without a corona and with white-

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tomentose capitula, which cannot be considered a separate species, given
the variation of C. alpina Bge. However, as the type for C. subtilis,
Juzepczuk selected a plant that undoubtedly belongs to the group of
species with a corona. Cf. also the note to C. pulchra Winkl.

180. C. carduncelloidea Rgl. and Schmalh. in Izv. Obshch. Lyubit
Estestv. Antrop. Etn. 34, 2 (1882) 50; Winkl. in Tr. Peterb. Bot. Sada,
XI, 244; XIV, 206; Bornm. in Beth. bot. Centralbl. XXXIV, 2, 160;
Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 556.—Arctium
carduncelloideum O. Ktze. Rev. Gen. 1 (1891) 307.

Biennial. Usually many-stemmed plants; stems erect, arachnoid-hairy,
striate-sulcate, 15-30 cm high, branched; branches densely leafy, one-
headed. Leaves coriaceous, green and finely arachnoid-hairy above, finely
gray-tomentose beneath; basal leaves petiolate with petioles broadened
at base, unevenly pinnatipartite into triangular, acuminate-spinescent
lobes, with stiff spine, large-spinose-toothed; cauline leaves gradually
reduced, sessile, semiamplexicaul, short-decurrent, lanceolate, lobed or
large spinose-toothed. Capitula ovoid-globose, 17 mm wide (excluding
cusps), arachnoid-hairy. Involucral bracts about 60, narrow-lanceolate,
gradually and long-acuminate, carinate; outermost bracts slightly
decurved, about 10 mm long; subsequent bracts almost horizontal, 23—
25 mm long; middle bracts 15—18 mm long, erect; inner bracts scarious,
erect, somewhat flat, 20-22 mm long, scabrous above along margin,
acuminate; corolla pink; receptacular bristles scabrous. Achenes about 4
mm long, glabrous, smooth. Flowering VI—VII; fruiting VII—VIII.

Stony gravelly slopes.—Soviet Central Asia: Pamiro-Alai (Alai
Range). Endemic. Described from the Isfairam Pass in “Kokania.” Type
in Tashkent; cotype in Leningrad.

Note. Plants very similar to C. carduncelloidea are found in Shugnan,
but whether they belong to this species or are some sort of hybrids
between C. rava Winkl. and C. buphtalmoides Rgl. that resemble C.
carduncelloidea, in fact can be decided only after studying the plants in
nature.

181. C. outichaschensis Franch. in Ann. Sc. Nat. Bot. 6 sér. 16
(1883) 322; Winkl. in Tr. Peterb. Bot. Sada, XII, 256; XIV, 208; Juz.
in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 566.—C. commixta Kult. in
Tr. Sredneaz. Gos. Univ. Ser. VIIIc, VI (1929) 18; Juz. op. cit. 566.—
C. glaucifolia Kult. op. cit. (1929) 19; Juz. op. cit. 566.—C.
buphthalmoides auct. non Rgl.: Winkl. in Tr. Peterb. Bot. Sada, XII
(1892) 258 p. p.; XIV, 230; Bornm. in Beih. bot. Centralbl. XXXIV, 2,
185.—C. carduncelloidea auct. non Rgl. and Schm.; Winkl. op. cit.
XIV (1897) 227.—Arctium outichaschense O. Ktze. Rev. Gen. 1 (1891)
308.—Ie.: Kult. op. cit. Figs. 18, 19.—Exs.: H.F.A.M. No. 481.

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282

Biennial. Stem 15-35 cm high, striate-sulcate, arachnoid-hairy,
branched from base or middle; branches one-headed, leafy. Leaves
greenish and finely arachnoid-hairy above, gray-tomentose beneath; basal
leaves petiolate, lanceolate, pinnatipartite into unequal, triangular-
lanceolate, acuminate-spinescent, spinose-toothed lobes, alternating with
smaller lobes, reduced to spines or lyrate, with oblong-orbicular terminal
lobe, acuminate-spinescent at base, slightly cuneately decurrent on
petiole, finely spinose-toothed, 25-30 mm long, with lateral lobes about
12-15 mm long, broadly sessile, ovate-orbicular, acuminate-spinescent;
lower cauline leaves like basal; middle leaves sessile, semiamplexicaul,
with small auricles, ovate-oblong, large spinose-toothed, acuminate into
stiff, slightly divergent spine; upper leaves thin, reduced, oblong-
lanceolate, spinose-toothed, acuminate-spinescent; uppermost leaves
strongly reduced. Capitula ovoid-globose, 15—20 mm wide (excluding
cusps), loosely arachnoid-hairy. Involucral bracts about 75; outer bracts
stiff, subulate-triquetrous, about 15-25 mm long, gradually acuminate
into decurved cusp; subsequent bracts longer, 25-28-30 mm long, almost
horizontal; middle bracts somewhat flat, 18-20 mm long, scarcely
carinate dorsally, mucronate; inner bracts coriaceous, 21—25 mm long,
flat, scabrous along margin, very short-acuminate into thin spine, often
pale purple above; innermost bracts linear, mucronate, whitish; corolla
pink, exserted from involucre; receptacular bristles scabrous. Achenes
obovoid; glabrous; smooth, roundish above, about 4—5 mm long.
Flowering VI-VII; fruiting VII—VIII.

Limestone and gypsum deposits, stony and gravelly slopes in middle
and upper mountain zones.—Soviet Central Asia: Pamiro-Alai (Kugitang,
Gissar, Zeravshan, Turkestan and Alai ranges). Endemic. Described from
the Zeravshan Valley (Outikasch). Type in Paris.

Note. A highly polymorphic species. At one time attempts were
made to separate several species on the basis of the shape of the
basal leaves. However, the study of extensive material shows that
plants with different kinds of basal leaves are found in the same
population, and there is no basis to consider such plants as separate
species.

Hybrids of C. outichaschensis Franch. and C. submutica Franch.,
described as C. iskanderi Bornm. (cf. the note to C. submutica Franch.)
are known. Hybrids of C. outichaschensis Franch. and C. pulchella Bge.
have also been described, as C. heterogenetos Bornm. (B.B.C. 34, 2
(1916) 163), from the Dzhidzhikrud River Valley near the village of
Zimarl (C. heterogenetos a. pulchelliceps Bornm. ibid. 164 and B.
majoriceps Bornm. ibid. 165). This is a plant with thin and rather long
decurrent leaves, arachnoid-hairy capitula, and numerous, thin, squarrose
involucral bracts.

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283

182. C. stellaris Bornm. in Beith. bot. Centralbl. XXXIV, 2 (1916) 161;
Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 559.—C. carduncelloidea
B. sonchifolia Bornm. op. cit. (1916) 160.—C. sonchifolia Bornm. ex
Juz. op. cit. (1940) 556.—C. rotundifolia auct. non Winkl.: Bornm. op. cit.
vie

Biennial. Stem 30-35 cm high, arachnoid-hairy, glossy, striate-
sulcate, branched at base or throughout; branches one-headed, leafy.
Leaves greenish, finely arachnoid-hairy above, densely tomentose
beneath; basal leaves petiolate, lyrate, with almost orbicular, 5—8.5—11
cm long, sometimes more or less cordate, spinose-toothed terminal lobe,
lateral lobes 1.5—2 cm long, thin, often three pairs, finely spinose-toothed,
broadly sessile, terminating in small, stiff spine; cauline leaves sessile,
auriculate, ovate-orbicular, spinose-toothed, terminating in small, stiff
spine; upper cauline leaves strongly reduced. Capitula ovoid-globose,
(1.5)2—2.5 cm wide (excluding cusps), arachnoid-hairy. Involucral bracts
about 60; outer bracts stiff, subulate-triquetrous, gradually acuminate in
long spine, decurved, about 25 cm long; middle bracts like outer but
longer, 30-32 mm long and horizontal; subsequent bracts coriaceous,
somewhat flat, slightly carinate, weakly attenuate in middle, scabrous
along margin, terminating in small stiff spine, 17-19 mm long; inner
bracts membranous, with broad ovate, about 5—6 mm wide whitish
appendage along margin, acuminate; innermost bracts linear-lanceolate,
scabrous along margin; corolla pink, exserted from involucre; receptacular
bristles scabrous. Achenes obovoid, glabrous, smooth, glossy, roundish
above, blackish, longitudinally whitish-striate, about 5 mm long.
Flowering V—VI-VII; fruiting VII—VIII.

In the foothills and lower mountain zone.—Soviet Central Asia:
Pamiro-Alai (foothills of the Alai Range). Endemic. Described from the
Alai Range (Margelan District, Shiman). Type in Leningrad.

183. C. buphtalmoides Rgl. in Bull. Soc. Nat. Mosc. XL, 3 (1867)
155; Boiss. Fl. or. III, 501; Winkl. in Tr. Peterb. Bot. Sada, XII, 258
quoad sp. afganica.—Arctium buphthalmoides O. Ktze. Rev. Gen. 1
(1891) 307.

Biennial. Stem erect, 15—45 cm high, striate-sulcate, arachnoid-hairy,
usually branched throughout or at base, leafy; branches one-headed.
Leaves coriaceous, finely arachnoid-hairy above, gray-tomentose beneath;
basal leaves petiolate, lyrate, with large, ovate, almost cordate, and finely
spinose-toothed terminal lobe, lateral lobes small, three to five pairs,
lanceolate, finely spinose-toothed or less often almost bipinnatilobate;
cauline leaves lanceolate-oblong, sessile, short-decurrent, pinnatilobate,
lobes acuminate-spinescent, spinose-toothed gradually reduced toward

| apex. Capitula ovoid, 13-15 mm wide (excluding cusps), finely

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284

arachnoid-hairy. Involucral bracts about 50; outermost bracts lanceolate,
attenuate into narrow-lanceolate cusp, usually decurved, 8—15(23) mm
long, with small thickening on inner side at transition of closely appressed
part with narrowed cusp; subsequent bracts similar but almost horizontal
or erect-spreading, about (17)30-32 mm long, strongly carinate dorsally;
middle bracts more flat, lanceolate, erect, short-acuminate, 12-15 mm
long; inner bracts lanceolate, flat, erect, acuminate, usually dark colored
dorsally in upper part, about 20 mm long; corolla purple; receptacular
bristles scabrous. Achenes obovoid, about 4 mm long; young achenes
with conspicuous ribs, terminating in teeth above, mature achenes
glabrous, smooth, with inconspicuous ribs. Flowering VI—VII; fruiting
VIII.

Gravelly slopes in middle mountain zone—Soviet Central Asia:
Pamiro-Alai (western Pamir). General distribution: Afghanistan.
Described from Afghanistan.

Note. Plants with lyrate basal leaves predominate, but plants with
leaves nearly bipinnate occur in the same population. Many different
forms are found within the range of C. buphthalmoides, whose taxonomic
status is difficult to determine at present. It is possible, that these are
hybrids of C. rava Winkl. and C. buphthalmoides among them and it is
probable that separate species also occur, but this can be decided only
after studying them in nature. Probably, these also include the already
described C. hilariae Kult. and C. darwasica Winkl. The plants from
Shugnan, which resemble C. carduncelloidea, evidently, also belong to
such forms.

184. C. hilariae Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIIIc, VI
(1929) 22; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 556.—Ic.:
Kult. op. cit. Fig. 21.

Biennial. Stem erect, about 70 cm high; branches one-headed,
arachnoid-hairy, striate-sulcate, leafy. Leaves green, arachnoid-hairy
above, gray-tomentose beneath; basal leaves ovate-oblong, short-petiolate,
with broadened petioles, finely spinose-toothed, lyrate, crenate-
pinnatipartite in lower part, terminal lobe ovate-oblong; lower cauline
leaves like basal but sessile, semiamplexicaul; middle and upper leaves
strongly reduced, oblong-ovate, sessile, semiamplexicaul, with roundish
auricles, attenuate-spinescent, spinose-toothed. Capitula ovoid-globose,
about 20 mm wide (excluding cusps), arachnoid-hairy. Involucral bracts’
40-50, all except inner broad-lanceolate, strongly carinate; outer bracts
12-20 mm long, usually decurved; middle bracts about 30 mm long,
divergent, grooved in upper half; subsequent bracts more flat, mucronate,
about 20 mm long; inner bracts membranous; lanceolate, yellowish,
scabrous along margin, acuminate, receptacular bristles scabrous. Corolla

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285

pale-pink, prominent. Achenes obovoid, roundish above, glabrous,
smooth, about 4 mm long. Flowering VI-—VII; fruiting VIJ—VIII.

In middle mountain zone.—Soviet Central Asia: Pamiro-Alai
(Shugnan). Endemic. Described from Pyandzh River Valley, near Ravak.
Type in Tashkent.

Note. A little studied species, known only from one locality.

185. C. knorringiae Bornm. in Beith. bot. Centralbl. XXXIV, 2
(1916) 201.

Biennial. Stem erect, 20-30 cm high, finely arachnoid-hairy, almost
glabrous in upper half, with scattered, flexuous flat hairs throughout,
branched almost from base, leafy; branches one-headed. Leaves
coriaceous, glabrous and green above, finely gray-tomentose beneath;
basal leaves short-petiolate, 25 cm long, oblong, lyrate, with roundish,
somewhat cordate and finely spinose-toothed terminal lobe and unequal,
almost ovate, slightly decurved, somewhat attenuate at both ends and
spinose-toothed lateral lobes; lower cauline leaves like basal but sessile
and smaller; other cauline leaves gradually reduced toward stem apex,
broad ovate or orbicular, cordate, sessile with roundish auricles, crenate-
spinose-toothed, stiff cuspidate. Capitula oblong-cylindrical, 40 mm long
(with florets) and 15—20 mm wide (excluding cusps), glabrous. Involucral
bracts less numerous (about 25); outer bracts lanceolate, gradually
acuminate, short spinescent, falcately decurved, 5 mm wide at base,
about 10-12 mm long; middle bracts somewhat flat, broad-lanceolate,
10-14 mm wide at base, 27-30 mm long, gradually acuminate-spinescent,
closely appressed at base, divergent in middle, weakly furrowed at
divergence; subsequent bracts flat, broad-lanceolate, roundish above, with
short spine, erect, 8-10 mm wide, 27 mm long; inner bracts membranous,
broad-lanceolate, usually light purple, scabrous along margin, short-
acuminate 32-35 mm long, 3—5 mm wide; receptacular bristles scabrous.
Corolla light purple; anther tube light creamish. Achenes obovoid, 4—
4.5 mm long, glabrous, smooth, roundish above. Flowering VII—VIII;
fruiting VIII-IX.

Stony-gravelly slopes.—Soviet Central Asia: Tien Shan (Uzun-
Akhmat Range). Endemic. Described from the Ungar-Tyube and Shatak-
Tape mountains. Type in Leningrad.

186. C. grigoriewii Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1,
3 (1936) 313; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 556.

Biennial. Stem erect, 30-35 cm high, arachnoid hairy-tomentose,
flexuously and repeatedly branched from base. Leaves finely arachnoid-
hairy and grayish-green above, white-tomentose beneath; basal leaves
oblong, short-petiolate, almost lyrate, lateral lobes oblong-triangular,
acuminate-spinescent, irregularly spinose-toothed, terminal lobe oblong-

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286

ovate, sometimes almost cordate, finely spinose-toothed; cauline leaves
gradually reduced; lower cauline leaves sessile, ovate, semiamplexicaul;
middle and upper leaves orbicular, sparsely spinose-toothed, somewhat
cordate, auriculate, usually adnate; uppermost leaves strongly reduced.
Capitula almost globose, 12-20 mm wide (excluding cusps), gray
arachnoid-hairy, with many florets. Involucral bracts lanceolate; outer
and middle bracts acuminate stiff spinescent, short dorsally carinate,
weakly grooved’ in divergent part, arcuately decurved, 10-35 mm long;
subsequent bracts coriaceous, somewhat flat, broad-lanceolate, somewhat
acuminate, terminating in thin and short erect spine, 17-19 mm long,
inner bracts membranous, broadened above, dark, with thin short spine,
scabrous along margin, 20-25 mm long; réceptacular bristles scabrous.
Corolla exserted, dark purple; anther tube glabrous, yellowish when dry.
Achenes obovoid, glabrous, smooth, light gray with dark spots, about
4.5 mm long. Flowering VI; fruiting VII.

- Stony, loamy and gypsic slopes in middle mountain zone.—Soviet
Central Asia: Pamiro-Alai (south). Endemic. Described from the top of
the Sarsaryak Range, above the village of Dagana. Type in Leningrad.

187. C. rava Winkl. in Tr. Peterb. Bot. Sada, XIV (1897) 227; Juz.
in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 556.

Perennial. Root woody, many-headed. Stem erect, arachnoid-hairy,
25—40 cm high, branched from base; branches long, one-headed. Leaves
coriaceous, bright green, finely arachnoid-hairy above, finely gray-
tomentose beneath; basal leaves short-petiolate, oblong to narrow-
lanceolate, pinnatipartite or crenate-lobate, segments or lobes broadly
triangular, acuminate-spinescent, finely spinose-toothed; lower cauline
leaves like basal but sessile, semiamplexicaul; middle and upper leaves
lanceolate, gradually reduced, acuminate-spinescent, spinose-toothed,
sessile, irregularly short-decurrent. Capitula cylindrical, about 15—17 mm
wide (excluding cusps), arachnoid-hairy. Involucral bracts about 60; outer
bracts lanceolate, acuminate-spinescent, carinate, almost horizontal, 15—
17 mm long; middle bracts erect, somewhat flat, lanceolate, usually
somewhat dark colored dorsally, acuminate-spinescent, 12-13 mm long;
inner bracts scarious, lanceolate, mucronate, flat, whitish, erect, 15—17
mm long. Corolla purple, 16 mm long; receptacular bristles scabrous.
Achenes obovoid, glabrous, smooth, about 4 mm long. Flowering VII—
VII; fruiting VII-IX.

Stony-gravelly slopes in upper mountain zone.—Soviet Central Asia:
Pamiro-Alai (Pamir, Shugnan). Endemic of Lake Yashil-Kul. Type in
Leningrad.

188. C. glabriseta Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIIIc, VI
(1929) 7; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 557.

287

Plate XXII.

1—Cousinia armena Takht.; 2—C. hystrix C.A.M.

291

288

Biennial. Stem 10-30 cm high, arachnoid-hairy, striate-sulcate, leafy,
branched from base; branches leafy, one-headed. Leaves coriaceous,
finely arachnoid-hairy above, finely white-tomentose beneath, spinose-
toothed; basal leaves short-petiolate, lyrate, with large ovate, about 7
cm long and 5 cm wide, terminal lobe slightly cuneate at base and
lanceolate, 2—2.5 cm long, 1—-1.2 cm wide lateral lobes; lower cauline
leaves like basal but sessile; middle and upper leaves gradually reduced,
orbicular, sessile, semiamplexicaul, with large roundish auricles and short
apical spine; uppermost leaves strongly reduced. Capitula ovoid-globose,
arachnoid-hairy, 20-22 mm wide (excluding cusps). Involucral bracts
about 50, squarrose; outer bracts stiff, carinate dorsally, horizontal;
middle bracts slightly reduced, weakly carinate dorsally; inner and
innermost bracts coriaceous, greenish, somewhat acuminate. Corolla pink;
receptacular bristles smooth. Achenes obovoid, about 4 mm long,
glabrous, smooth. Flowering V—VI; fruiting VII.

Gravelly slopes in upper mountain zone.—Soviet Central Asia: —
Pamiro-Alai (Kugitang). Endemic. Described from the Kugitang and
Kyzylalma mountains. Type in Tashkent.

189. C. triceps Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIlIc, VI
(1929) 8; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR 8, 557.

Biennial. Stem 15-22 cm high, usually developing three branches
from base or middle, arachnoid-hairy, striate-sulcate, leafy; branches
one-headed. Leaves coriaceous, finely arachnoid-hairy above, white-

293 tomentose beneath; basal leaves lyrate, short-petiolate with roundish,
finely spinose-toothed, acuminate, weakly cordate terminal lobe, lateral
lobes broadly sessile, finely spinose-toothed, alternating with smaller
lobes; lower cauline leaves like basal but sessile; middle leaves oblong-
orbicular, acuminate, finely spinose-toothed; uppermost leaves very small,
almost reduced to spines. Capitula ovoid-globose, arachnoid-hairy, 10—
15(20) mm wide (excluding cusps). Involucral bracts about 50, squarrose;
outer bracts stiff, dorsally carinate, terminating in three-angled spine,
subsequent bracts like outer, horizontal; middle bracts broad linear,
carinate, short-acuminate; inner bracts coriaceous, linear, green,
acuminate; innermost bracts with narrow membranous frill along margin.
Corolla yellow; receptacular bristles smooth. Achenes obovoid, glabrous,
smooth, about 4 mm long. Flowering VI-—VII; fruiting VIJ—VIII.

Gravelly slopes in upper mountain zone.—Soviet Central Asia:
Pamiro-Alai (Kugitang). Endemic. Described from the Kugitang
Mountains of Kyzylalma. Type in Tashkent.

190. C. fetissowii Winkl. in Tr. Peterb. Bot. Sada, XII (1892) 247.
Biennial. Stem 20-40 cm high, striate-sulcate, arachnoid-hairy,
usually sparsely branched almost from base; branches one-headed, leafy.

294

289

Leaves greenish and finely sparsely arachnoid-hairy above, finely gray-
tomentose beneath; basal leaves short-petiolate, lanceolate, large spinose-
toothed, lower cauline leaves like basal; other cauline leaves sessile,
semiamplexicaul, spinose-toothed, roundish at base gradually reduced
upward. Capitula 20-25 mm wide (excluding cusps), arachnoid-hairy,
broad pyramidal, weakly foveate at base. Involucral bracts numerous,
narrow-lanceolate, acuminate-spinescent; outer and middle bracts
arcuately or completely decurved, 15—20 mm long; subsequent bracts
squarrose, 10-17 mm long; inner bracts lanceolate, short-acuminate,
usually dark-colored throughout, about 20 mm long; receptacular bristles
scabrous. Corolla purple. Achenes obovoid, glabrous, smooth, about 4—
5 mm long. Flowering VI-VII; fruiting VII—VIII. (Plate XXIII, Fig. 2).

Gravelly slopes in middle mountain zone.—Soviet Central Asia: Tien
Shan. Endemic. Described from the headwaters of the Uzun-Akhmat
(Terek-Bel Ravine). Type in Leningrad.

Note. A highly polymorphic species, its morphological and geo-
graphical separation from C. outichaschensis Franch. is not clear at
present, as the specimens of the latter species from the upper Zeravshan
are very similar to C. fetissowii Winkl. in habit, smaller capitula,
narrower and shorter involucral bracts and more or less strongly decurved
outer involucral bracts. The plants in the Talass Alatau are sturdier,
with larger capitula; plants in the region of the Fergana Range are
intermediate between C. fetissowii Winkl. and C. schischkinii Juz. in
their characters.

191. C. schischkinii Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser.
Leedu(1936), 3.16.

Biennial. Plants very similar to C. fetissowii Winkl., differing from
it by stronger stem; narrower, cuneate (and not roundish), basally always
deeply pinnatilobate cauline leaves; one and one-half times as large
capitula; shorter involucral bracts, of which outer squarrose (and not
arcuately decurved), inner erect, squarrose, pale throughout (and not
greenish-black in lower part). Flowering VII; fruiting VIII.

Stony slopes in upper mountain zone.—Soviet Central Asia: Tien
Shan (Fergana Range). Endemic. Described from the Airtash Pass. Type
in Leningrad.

192. C. speciosa Winkl. in Tr. Peterb. Bot. Sada, X (1889) 579;
XII, 257; XIV, 230; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8,
557.—C. batalini auct.; Winkl. op. cit. (1897) 299 quoad sp. alaicam.

Biennial. (Monocarpic.) Stem usually branched from base, shrubby,
30-50 cm high; branches arachnoid-hairy, striate-sulcate, leafy, usually
simple, one-headed, less often sparingly branched. Leaves coriaceous,

295

290

green and very finely arachnoid-hairy above, finely gray-tomentose
beneath; basal leaves oblong, petiolate, pinnatilobate, with spinose-
toothed, broadly triangular, acuminate-spinescent lateral lobes; lower
cauline leaves like basal but with shorter petiole; middle and upper
leaves gradually reduced upward, broad-lanceolate, attenuate toward both
ends, sessile, spinose-toothed, attenuate into short stiff spine, somewhat
erectopatent. Capitula broad cylindrical, 20-25 mm wide (excluding
cusps), finely-arachnoid-hairy, truncate at base or weakly foveate.
Involucral bracts about 70, coriaceous; outer and middle bracts erect-
spreading, lanceolate, carinate dorsally, gradually acuminate into short
stiff spine, 15-35 mm long, about 5-6 mm wide; inner bracts
membranous, whitish, lanceolate, scabrous along margin, gradually
acuminate, about 25-30 mm long, erect. Corolla pinkish; receptacular
bristles scabrous. Achenes obovoid, about 4—5 mm long, glabrous,
smooth. Flowering VII—VIII; fruiting VIII-IX. .
Stony slopes and screes in middle mountain zone.—Soviet Central
Asia: Pamiro-Alai (Alai Range). Endemic. Described from the Alai
Range (Archaty). Type in Leningrad.

193. C. batalinii Winkl. in Tr. Peterb. Bot. Sada, XII (1892) 248;
Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 557 (quoad typum).—C.
batalinii a. latisecta Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916)
167.

Biennial. (Or monocarpic?) Stem erect, 40-60 cm high, leafy,
arachnoid hairy-pubescent, spreadingly branched from base or middle;
branches long, one-headed. Leaves coriaceous, green and finely
arachnoid-hairy above, gray-tomentose beneath with prominent veins;
basal leaves oblong, petiolate, with petioles broadened at base,
pinnatilobate, with broadly triangular, usually large spinose-toothed,
acuminate in stiff spine lateral lobes; lower cauline leaves like basal
but sessile; middle and upper leaves gradually reduced, sessile, almost
semiamplexicaul with roundish auricles, ovate or oblong-ovate, spinose-
lobed or large spinose-toothed, attenuate into stiff, arcuate cusp. Capitula
broadly pyramidal, 20-25 mm wide (excluding cusps), arachnoid-hairy.
Involucral bracts about 50; outer and middle bracts almost horizontal,
lanceolate, long acuminate, carinate dorsally, 30-32 mm long; inner
bracts flat, membranous, lanceolate or linear-lanceolate, somewhat
scabrous along margin, whitish above, about 30 mm long. Corolla purple;
receptacular bristles scabrous. Achenes obovoid, glabrous, smooth, about
5 mm long. Flowering VI-VII; fruiting VII—VIII.

Stony-gravelly slopes in middle and lower parts of the upper
mountain zone.—Soviet Central Asia: Pamiro-Alai. Endemic. Described
from the Vakhsh River Valley, from Tavildar. Type in Leningrad.

296

291

194. C. calva Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3
(1936) 317; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 557.—C.
batalinii B. tenuisecta Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916)
168.

Biennial. Stem erect, striate-sulcate, 50-70 cm high, pubescent in
lower half with thick, flexuous, flat hairs, arachnoid-hairy above, sparsely
branched from middle; branches one-headed, long. Leaves coriaceous,
green above, glabrous beneath or glabrous and with flat, thick, flexuous
hairs only along midrib, or finely arachnoid-hairy; basal leaves almost
pinnatisect in narrow-lanceolate, spinose, numerous (12-15 pairs) lateral
segments, petiolate, with petioles broadened at base, densely pubescent
throughout with thick, flexuous, flat hairs; lower cauline leaves like
basal; middle and upper leaves sessile, semiamplexicaul, gradually
reduced toward stem apex, pinnatipartite into strongly spinose-toothed
lobes; uppermost leaves strongly reduced. Capitula broadly cylindrical,
about 25 mm wide (excluding cusps), white arachnoid-hairy; involucral
bracts about 70; outer and middle bracts narrow lanceolate, squarrose,
gradually acuminate-cuspidate, dorsally carinate, 20-25 mm long; inner
bracts lanceolate, membranous, whitish above, acute, scabrous along
margin, erect, about 30 mm long. Corolla purple; receptacular bristles
scabrous. Achenes obovoid, glabrous, smooth, about 6 mm long.
Flowering V—VI; fruiting VI-VII.

Exposed red clay from middle to lower border of the upper mountain
zone.—Soviet Central Asia: Pamiro-Alai (spurs of the Khozretisho
Range). Endemic. Described from the city of Imam-Askara. Type in
Leningrad.

195. C. oxytoma Rech. f. in Dan. Biol. Skr. 8, 2 (1955) 135.—C.
batalinii 8. virescens Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916)
168.—Ic.: Rech. f. op. cit. 139, fig. 93, 94G.

Biennial. Stem erect, about 30 cm high, striate-sulcate, arachnoid-
hairy, leafy, branched from base; branches one-headed. Leaves
coriaceous, green and finely arachnoid-hairy above, densely and finely
gray arachnoid-hairy beneath with prominently raised veins; basal leaves
petiolate, with petioles broadened at base, often dark colored, almost
bipinnatisect, lateral segments narrow lanceolate, acuminate-spinescent,
second order segments unequal, acuminate-spinescent, spinose-toothed;
lower cauline leaves like basal, but sessile; middle and upper leaves
gradually reduced toward stem apex, pinnatipartite into lanceolate,
acuminate-spinescent, spinose-toothed, sessile lobes with small auricles.
Capitula broadly cylindrical, 20-22 mm wide (excluding cusps),
arachnoid-hairy. Involucral bracts about 70; outer and middle bracts
divergent, sometimes outer bracts decurved, 20-22 mm long, lanceolate,

297

292

acuminate-spinescent; inner bracts lanceolate, whitish, scabrous along
margin, acuminate-short spinescent, erect, 20-22 mm long. Corolla pink;
receptacular bristles scabrous. Achenes obovoid, glabrous, smooth,
immature achenes about 4 mm long. Flowering VII.

Stony mountain slopes.—Soviet Central Asia: Pamiro-Alai (Peter
the Great Range). General distribution: Afghanistan. Described from
Afghanistan. Type in Vienna.

196. C. ovezinnikovii Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 99.

Biennial (or monocarpic?). Stem erect, arachnoid-hairy, striate-
sulcate, 50-60 cm high, usually branched from middle or almost from
base; branches long, simple, sparsely leafy, one-headed. Leaves
coriaceous, grayish-green and finely arachnoid-hairy above, finely gray-
tomentose beneath; basal leaves petiolate, with short and broadened ~
petioles, narrow-lanceolate, 2(3) cm wide and 12-13 cm long,
pinnatilobate, lobes broadly triangular, acuminate-spinescent, spinose-
toothed, approximate; lower cauline leaves like basal but somewhat
smaller and sessile; leaves on branches oblong, spinose-toothed, sessile,
short-decurrent, acuminate in short cusp. Capitula flat-globose, about 20
mm wide (excluding cusps), somewhat rusty-tomentose. Involucral bracts
about 80; outer and middle bracts lanceolate, gradually attenuate into
cusp, horizontal or weakly decurved, 15—20(23) mm long; subsequent
bracts broad lanceolate, mucronate, 10-12 mm long; inner bracts
membranous above, with small cusp, whitish, 22 mm long. Corolla pale
pink; anther tube purple; receptacular bristles scabrous. Achenes obovoid,
glabrous, smooth, dark-spotted, about 5 mm long. Flowering VII; fruiting
VIII. (Plate XXIV, Fig. 2.)

Stony screes in upper mountain zone.—Soviet Central Asia: Pamiro-
Alai (Peter the Great Range). Endemic. Described from the Garmo
Glacier. Type in Dushanbe.

197. C. peduncularis Juz. ex Tschern. in Bot. Mat. Gerb. Inst. Bot.
Akad. Nauk UzbSSR, XVII (1962) 101.

Biennial (or monocarpic?). Stem profusely branched almost from
base into long one-headed branches, almost leafless in upper half,
flocculose-arachnoid hairy, moderately leafy below, about 80 cm high.
Leaves coriaceous, green, and glabrous above, white-tomentose beneath;
basal leaves long-petiolate (with petioles 20-35 cm long), petioles almost
as long as lamina, oblong-ovate, undivided or almost lyrate, with large,
broadly ovate terminal lobe and three or four pairs of broadly sessile
oblong lateral lobes; cauline leaves few, strongly reduced, ovate, spinose-
lobate or large spinose-toothed, sessile, acuminate in stiff, divergent spine;

298

293

uppermost leaves almost reduced to spines. Capitula broadly pyramidal,
20-35 mm wide (excluding cusps), white arachnoid-hairy, similar to
capitula in C. batalinii. Corolla pale pink; receptacular bristles scabrous.
Achenes obovoid, glabrous, smooth, about 7 mm long. Flowering V—
VI; fruiting VI. (Plate XXV, Fig. 2.)

On red sandstones.—Soviet Central Asia: Pamiro Alai (south).
Endemic. Described from the Kyzyl-Su River Basin in the Karatau
Mountains. Type in Leningrad.

198. C. gulezensis Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIlIIc, VI
(1929) 16; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 557.—C.
corymbosa f. ramosissima Wink. in Tr. Peterb. Bot. Sada, XIV (1897)
219.—Ie.: Kult. op. cit. Fig. 16.

Biennial. Stem erect, 25-50 cm high, striate-sulcate, finely arachnoid-
hairy, usually branched from base; branches one-headed. Leaves grayish-
green and finely arachnoid-hairy above, gray-tomentose beneath; basal
leaves petiolate; lower cauline leaves petiolate, with petioles almost as
long as lamina, or subsessile, attenuate toward base, lanceolate,
pinnatilobate, lateral lobes long acuminate-spinescent, spinose-toothed;
middle leaves sessile, broad-lanceolate, attenuate toward both ends,
spinose-toothed, spinescent; upper leaves ovate, acuminate-spinescent
spinose-toothed, sessile, with roundish auricles. Capitula ovoid-oblong,
about 20 mm wide (excluding cusps), finely arachnoid-hairy. Involucral
bracts about 80; outer bracts narrow lanceolate, gradually acuminate-
short cuspidate, weakly carinate dorsally, arcuately decurved, 8-10 mm
long; subsequent bracts more stiff, lanceolate, mucronate, arcuately
decurved, 15—22 mm long; middle bracts oblong-lanceolate, somewhat
erect-spreading, more or less flat, 18—22 mm long; inner bracts
membranous in upper part, erect, forming “corona,” usually whitish or
very pale purple, lanceolate, gradually acuminate in short thin cusp;
scabrous along margin, 30-35 mm long; innermost bracts almost linear,
about 30 mm long; receptacular bristles scabrous. Corolla purple, usually
not exserted from corona or almost equaling. Achenes obovoid, glabrous,
smooth, roundish above, about 4.5 mm long. Flowering VI; fruiting
VU-VI.

Middle mountain zone.—Soviet Central Asia: Pamiro-Alai (Alai
Range). Endemic. Described from the Alai Range (between Karabulak
and Gulche1). Type in Leningrad.

199. C. schtschurowskiana Rg]. and Schmalh. in Izv. Obshch.
Lyubit Estestv. Antrop. i Etn. 34, 2 (1882) 50 p. p. quoad sp. e Dshisman;
emend. Kult. and Byull. Inst. Pochv. i Geobot. Sredneaz. Gos. Univ. 1,
111; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 557.—Arctium
stshurowskianum O. Ktze. Rev. Gen. 1 (1891) 308.

299

294

Biennial. Stem erect, 30-40 cm high, densely arachnoid-hairy,
branched from middle; branches long, leafy, one-headed. Leaves green,
arachnoid-hairy above, gray-tomentose beneath, short spinose-toothed,
acuminate in triquetrous spine; basal leaves broad petiolate, lyrate, with
large, ovate, cordate, short spinose-toothed terminal lobes, and three or
four pairs of broadly sessile, short spinose-toothed lateral lobes; cauline
leaves gradually reduced upward; lower cauline leaves oblong, lobate, |
acuminate-spinescent, short spinose-toothed, sessile; middle and upper
leaves sessile, semiamplexicaul, auriculate, oblong-ovate, acuminate-
spinescent, spinose-toothed; uppermost leaves strongly reduced. Capitula
with many florets, 35-40 mm wide (excluding cusps), broadly cylindrical,
flocculose-tomentose. Involucral bracts numerous, carinate dorsally,
grooved in upper half; outer bracts 20 cm long and 5 mm wide, broad-
lanceolate, acuminate-short spinescent; middle. bracts lanceolate, 25—30
mm long and 4 mm wide at base, acuminate-stiff spinescent, subsequent
bracts thin, coriaceous, somewhat flat, lanceolate, 25-30 mm long and
3.5 mm wide, acuminate-thin spinescent; inner bracts about 45-50 mm
long, 2.5 mm wide at base, with membranous appendages forming
corona, exceeding adjacent middle bracts by 25-30 mm; receptacular
bristles scabrous. Corolla not exserted from corona, pink. Achenes
obovoid, roundish above, compressed dorsally, light gray, about 5 mm
long. Flowering V—VI; fruiting VI—VII.

Gravelly and stony slopes in middle mountain zone.—-Soviet Central
Asia: Pamiro-Alai (Nuratau, Aktau, Khabduntau mountains). Endemic.
Described from Nuratau Mountains, Dzhizman Ravine. Type in Tashkent;
cotype in Leningrad.

200. C. pseudodshisakensis Tschern. and Vved. in Bot. Mat. Gerb.
Inst. Bot. Akad. Nauk UzbSSR, 16 (1961) 59.—C. dschisakensis Kult.
in Byull. Inst. Pochv. Geobot. Sredneaz. Gos. Univ. 1 (1925) 115 p. p.
quoad spec. e Pschigar.; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8,
558 p. p.—C. stshurowskiana auct. non Rgl. and Schmalh.; Winkl. in
Tr. Peterb. Bot. Sada, XII (1892) 253 quoad sp. e Warsaminor.

Biennial. Stem erect, 20-50 cm high, somewhat arachnoid-hairy,
striate-sulcate, leafy, branched almost from base; branches long, leafy,
one-headed. Leaves green, finely arachnoid-hairy above, appressed white-
tomentose beneath; basal leaves broad-petiolate, lyrate, with oval, cuneate
at base, and short spinose-toothed terminal lobe, and three or four pairs
of sessile, ovate, acuminate-spinescent, short spinose-toothed lateral lobes;
lower cauline leaves sessile, oval-oblong, crenate-lobate; middle and
upper leaves sessile, short-decurrent, semiamplexicaul, roundish, strongly
divergent, short spinose-toothed, spinescent; outermost leaves reduced.
Capitula 25-30 mm wide (excluding cusps), arachnoid-hairy, broadly

300

295

cylindrical, constricted below corona, with many florets. Involucral bracts
numerous, carinate-grooved with subulate-triquetrous cusp; outer and
middle bracts squarrose, almost horizontal, narrow-lanceolate, acuminate;
outer bracts 20 mm and middle 25 mm long; subsequent bracts somewhat
flat, lanceolate, 21-22 mm long, 3.5 mm wide at base, acuminate-thin
spinescent; inner bracts linear-lanceolate, 35 mm long, with broad-
lanceolate, membranous, acuminate-thin spinescent appendages forming
“corona”, exceeding adjacent middle bracts by 15-20 mm; innermost
bracts membranous, linear, 35 mm long and 1 mm wide; receptacular
bristles scabrous. Achenes obovoid, roundish above, glabrous, smooth,.
about 4 mm long. Flowering V—VI-VII; fruiting VII—VIII. (Plate XXIV,
Fig:,,tli.)

Gravelly and stony slopes, on rocks in middle mountain zone.—
Soviet Central Asia: Pamiro-Alai (Turkestan and Malguzar ranges).
Endemic. Described from Guralash. Type in Tashkent.

201. C. dshisakensis Kult. in Byull. Inst. Pochv. Geobot. Sredneaz.
Gos. Univ. 1 (1925) 115 quoad spec. e Dshisak; Juz. in Tr. Tadzh.
Bazy Akad. Nauk SSSR, 8, 558 p. p.

Biennial. Stem 20-35 cm high, flocculose-arachnoid hairy, striate-
sulcate, branched from middle; branches one-headed, approximate,
forming compact bush. Leaves green, arachnoid-hairy above, gray-
tomentose beneath, large spinose-toothed, acuminate in stiff spine; basal
leaves lyrate, broad-petiolate, with large, ovate, almost undivided terminal
lobe, lateral lobes remote, oblong, spinescent, three or four pairs; lower
cauline leaves oblong, lobed, spinescent; middle and upper leaves oblong-
ovate, sessile, weakly decurrent; uppermost leaves sessile, reduced.
Capitula with many florets, flocculose-tomentose, ovoid-globose,
constricted below corona, 20(25) mm wide (excluding cusps). Involucral
bracts numerous, grooved-carinate; outer bracts lanceolate, acuminate
into short decurved spine, 17 mm long and 2 mm wide at base; middle
bracts like outer but squarrose, about 20 mm long and almost 3 mm
wide at base; subsequent bracts somewhat flat, erect, mucronate, 22
mm long, 3 mm wide at base; inner bracts 35 mm long, with
membranous lanceolate appendages at apex, about 2.5 mm wide, forming
corona; innermost bracts linear, scarious, about 35 mm long; receptacular.
bristles scarious. Corolla slightly exserted from corona, pink. Achenes
obovoid, almost three-angled, roundish at apex, light gray. Flowering
V; fruiting VI.

Gravelly slopes in middle mountain zone.—Soviet Central Asia:
Pamiro-Alai. Endemic. Described from the vicinity of Dzhizak. Type in
Tashkent.

302

296

202. C. princeps Franch. in Ann. Sc. Nat. Bot. 6 sér. 16 (1883)
324; Winkl. in Tr. Peterb. Bot. Sada, XII, 258; XIV, 230; Juz. in Tr.
Tadzh. Bazy Akad. Nauk SSSR, 8, 558.—C. stshurowskiana auct. non
Rgl. and Schmalh.: Winkl. op. cit. XII, 253, quoad sp. e Iskanderkul —
C. schtschurowskiana y. subadpressa Bornm. in Beih. bot. Centralbl.
XXXIV, 2 (1916) 181.—Arctium princeps O. Ktze. Rev. Gen. 1 (1891)
308.

Biennial? Stem erect, 30-50 cm high, branched almost from base;
branches long, erect, leafy, arachnoid-hairy, one-headed. Leaves
coriaceous, green and finely arachnoid-hairy above, gray-tomentose
beneath; basal leaves petiolate, petioles thick and flattened, sometimes
armed along margin, lyrate, usually with four or five pairs of oblong,
short-spinescent and spinose-toothed lateral lobes and broad ovate, short
spinose-toothed, acuminate-spinescent, sometimes cordate terminal lobe;
lower cauline leaves usually like basal but with broader petiole, or
pinnatipartite; other cauline leaves gradually reduced toward apex, sessile,
broad-ovate, short acuminate into cusp, weakly crenate-spinose-toothed,
semiamplexicaul, auriculate, auricles adnate, short-decurrent. Capitula
globose, 22-27 mm wide (excluding cusps), finely arachnoid-hairy.
Involucral bracts numerous, somewhat flat, all erect; outer bracts
lanceolate, somewhat erect or sometimes almost horizontal, 15-25 mm
long; all others except inner 22-28 mm long; inner bracts membranous
in upper half, broad-lanceolate, gradually acuminate, usually pale purple,
about 40 mm long, forming 25 mm long “corona;” receptacular bristles
scabrous. Corolla purple, not exserted from “corona.” Achenes narrowly
obovoid, glabrous, smooth, lustrous, roundish above, about 6 mm long.
Flowering VII—VIII; fruiting VIII.

Stony slopes and outcrops of varicolored rocks in upper mountain
zone.—Soviet Central Asia: Pamiro-Alai (Gissar and Zeravshan ranges).
Endemic. Described from the Anzob Pass in the Yagnob River Valley.
Type in Paris.

203. C. laniceps Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3
(1936) 316; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 558.—C.
schtschurowskiana «a. laniceps Bornm. in Beth. bot. Centralbl. XXXIV,
2 (1916) 181.

Biennial? Stem erect, 20-25 cm high, branched almost from base,
with robust one-headed branches. Leaves on both sides arachnoid-hairy,
grayish-green, more densely beneath; basal leaves petiolate, pinnatisect
or pinnatipartite, lateral segments broadly ovate, acuminate-spinescent,
irregularly spinose-toothed; lower cauline leaves like basal but slightly
smaller and with shorter petioles; all other leaves gradually reduced,
sessile, orbicular or oblong-ovate, short-decurrent with adnate auricles,

297

SAN

int

;

S so

,

iG

ul
1 Pa — ree

Plate XXIII.
1—Cousinia botschanzevii Juz.; 2—C. fetissowii Winkl.

301

303

298

spinose-toothed, with short and stiff cusp at apex. Capitula 25-35 mm
wide (excluding cusps), densely white-tomentose, subglobose, slightly
constricted below “corona.” Involucral bracts lanceolate, carinate, outer
and middle bracts arcuately decurved, 17-22 mm long; subsequent bracts
somewhat flat, erect, short-acuminate, 20 mm long, inner bracts
membranous above, broadly lanceolate, about 40 mm long, purple, erect,
forming 15-20 mm long “corona,” obtuse above, scabrous along margin;
receptacular bristles scabrous. Corolla purple, not exserted from “corona.”
Achenes obovoid, glabrous, smooth, roundish above, about 5 mm long,
dark gray. Flowering V—VI; fruiting VI-—VII.

Gypsic stony slopes in foothills—Soviet Central Asia: Pamiro-Alai
(foothills of the Alai Range). Endemic. Described from Kokterek. Type
in Leningrad.

204. C. tianschanica Kult. in Byull. Inst. Pochv. Geobot. Sredneaz.
Gos. Univ. 1 (1925) 113 p. p., excl. sp. mogoltavica.

Biennial. Stem erect, 22-25 cm high, striate-sulcate, sparsely
arachnoid-hairy, leafy, branched from base; branches one-headed. Leaves
light green, finely arachnoid hairy above, arachnoid-hairy beneath, large
spinose-toothed; basal and lower cauline leaves lanceolate, short-petiolate,
pinnatipartite or pinnatisect, lateral segments acuminate-spinescent;
middle leaves oblong, sessile, undivided or lobate, acuminate-spinescent;
upper leaves sessile, semiamplexicaul, acuminate-spinescent. Capitula
ovoid, constricted below corona, 18-25 mm wide (excluding cusps),
weakly arachnoid-hairy, with many florets. Involucral bracts numerous,
stiff, carinate; outer bracts lanceolate, about 15 mm long and 3 mm
wide at base, gradually acuminate in falcately decurved cusp; middle
bracts like outer but divergent, about 20 mm long and 4 mm wide at
base; subsequent bracts somewhat flat, broadly lanceolate, about 20 mm
long and 5 mm wide at base, short-acuminate; inner bracts thin
coriaceous, about 30 mm long, with membranous apical appendages,
gradually acuminate in short thin spine, forming “corona;” innermost
bracts membranous, about 35 mm long, linear, short-acuminate;
receptacular bristles scabrous. Corolla pink, slightly exserted. Achenes
oblong-ovoid, almost three-angled, smooth, lustrous, roundish above,
gray, dark-spotted, about 5 mm long. Flowering VI-VII; fruiting VII—
VIII.

Stony-gravelly slope near the lower boundary of middle mountain
zone.—Soviet Central Asia: Tien Shan (western). Endemic. Described
from the Akbash-Tau Mountains, Badam River (village of Dorofeevka).
Type in Tashkent.

Note. A type is not designated in the description of the species, but
on the specimen with the label: “Syr-Darya Region, Chimkent Area,

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299

Akbash-Tau Mountains, Badam River (village of Dorofeevka). Zone of
steppe rocky xerophytes. Stony slope, 1921 VIII 10. Abolin, Popov, No.
8047”; “Typus” is written in Kultiasow’s hand. The plant cited should
be considered the lectotype of the species.

205. C. mogoltavica Tschern. and Vved. in Bot. Mat. Gerb. Inst.
Bot. Akad. Nauk UzbSSR, 16 (1961) 55.—C. tianschanica Kult. in Byull.
Inst. Pochv. Geobot. Sredneaz. Gos. Univ. 1 (1925) 113, quoad spec.
mogoltavica.—Exs.: H.F.A.M. No. 196 (sub. C. tianschanica).

Biennial. Stem erect, 25—45 cm high, arachnoid-hairy, striate-sulcate,
leafy, branched from base; branches one-headed. Leaves somewhat
pubescent, tomentose beneath; basal leaves broad-petiolate, lyrate,
terminal lobes ovate, weakly cordate, spinose-toothed, lateral lobes
remote, oblong, acuminate-spinescent, five or six pairs; lower and middle
cauline leaves crenate-lobate, sessile, short-decurrent, acuminate-
spinescent; upper leaves reduced, ovate, sessile, auriculate, large spinose-
toothed; uppermost leaves strongly reduced. Capitula with many florets,
flat-globose, somewhat constricted below corona, 20—25(30) mm wide
(excluding cusps), weakly arachnoid-hairy. Involucral bracts numerous,
grooved-carinate; outer bracts undivided, about 15—17 mm long, gradually
acuminate into spine, decurved; middle bracts like outer but erect-
spreading, more acuminate up to 25 mm long; subsequent bracts narrow-
lanceolate, somewhat flat, 20-22 mm long; inner bracts 40 mm long,
with long, broad-lanceolate, membranous appendages above, forming
corona; innermost bracts membranous, linear, about 40 mm long;
receptacular bristles scabrous. Corolla pink, not exserted from “corona.”
Achenes obovoid, glabrous, smooth, about 4 mm long. Flowering V—
VI; fruiting VI-VII.

Gravelly slopes in middle mountain zone.—Soviet Central Asia: Tien
Shan (Mogoltau). Described from the Mogoltau Range, Spa town on
the Uch-Bakh side. Endemic. Type in Tashkent.

206. C. butkovii Tschern. and Vved. in Bot. Mat. Gerb. Inst. Bot.
Akad. Nauk UzbSSR, 16 (1916) 54.

Biennial. Stems erect, 35-40 cm high, striate-sulcate, tomentose,
leafy, branched from base; branches one-headed. Leaves light green,
very finely arachnoid-hairy above, grayish arachnoid-hairy beneath,
spinose-toothed, acuminate-spinescent; basal and lower cauline leaves
short-petiolate, lanceolate, pinnatipartite, terminal lobe not exceeding
laterals, lateral lobes oblong, remote, alternating with smaller lobes;
middle cauline leaves oblong lobed; upper leaves orbicular, semi-
amplexicaul, auriculate, sometimes attenuate toward apex, acuminate-
spinescent, decurved; uppermost leaves strongly reduced or reduced to
spines. Capitula flat-globose, (20)25-35 mm wide (excluding cusps),

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300

densely arachnoid-hairy, with many florets. Involucral bracts numerous,
scabrous, grooved-carinate, outer bracts falcately decurved, lanceolate,
20-22 mm long, acuminate-spinescent; middle bracts like outer, but
divergent, 32-35 mm long; subsequent bracts somewhat flat, broadly
lanceolate, 20-25 mm long, short-acuminate; inner bracts 25-30 mm
long, with about 6 mm wide roundish, membranous appendage; innermost
bracts linear, membranous, 30-35 mm long and about 2 mm wide;
receptacular bristles scabrous. Corolla pink, not exserted from “corona.”
Achenes oblong-ovoid, roundish above, about 4.5 mm long. Flowering
V-VI-VII; fruiting VII—-VIII.

Gravelly slopes and screes in lower mountain zone.—Soviet Central
Asia: Pamiro-Alai (Samarkand Mountains, Zeravshan Range). Endemic.
Described from the Samarkand Mountains, vicinity of the village of
Sazagan. Type in Tashkent.

207. C. praestans Tschern. and Vved. in Bot. Mat. Gerb. Inst. Bot.
Akad. Nauk UzbSSR, 16 (1961) 57.

Biennial. Stem erect, up to 1 m high, flocculose-tomentose, striate-
sulcate, leafy, spreadingly branched from base; branches long, leafy,
one-headed. Leaves relatively soft, weakly spinose, finely arachnoid-
hairy above, gray-tomentose beneath; lower cauline leaves sessile,
semiamplexicaul, short-decurrent, broad-oblong, pinnatilobate, lateral
lobes broadly triangular, acuminate, short-spinescent, irregularly short
spinose-toothed; middle leaves sessile, short-decurrent, ovate, lobed, lobes
broadly triangular, acute, short spinose-toothed; upper leaves sessile,
short-decurrent, almost orbicular, acute, spinose-toothed, approximate.
Capitula globose, about 30-35 mm wide (excluding cusps), somewhat
truncate at base, densely arachnoid-hairy. Involucral bracts scabrous,
carinate-grooved; outer bracts decurved, lanceolate, with stiff, about 25
mm long cusp; middle bracts divergent, lanceolate, with long and stiff
cusp, about 45 mm long, up to 5 mm wide at base; subsequent bracts
somewhat flat, erect, short-acuminate, about 25 mm long; inner bracts
thin-coriaceous, lanceolate, about 30 mm long; apically broadened into
membranous, 5—7 mm wide, usually purple, spinescent appendages
forming short “corona;” innermost bracts membranous, linear, acuminate;
receptacular bristles scabrous. Corolla scarcely exserted from “corona,”
pink. Achenes obovoid, smooth, lustrous, roundish above, gray with dark
spots, about 5 mm long. Flowering V—VI; fruiting VI.

Rocky cliffs in middle mountain zone.—Soviet Central Asia: Pamiro-
Alai (Gissar Range). Endemic. Described from the Ayakchi River Valley
in the Gissar Range. Type in Tashkent.

208. C. strobilocephala Tschern. and Vved. in Bot. Mat. Gerb.
Inst. Bot. Akad. Nauk UzbSSR, 16 (1961) 62.

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301

Biennial. Stem erect, 30-50 cm high, striate-sulcate, flocculose-
arachnoid hairy, leafy, branched from base; branches long, one-headed.
Leaves grayish arachnoid-hairy on both sides; basal leaves petiolate,
petioles broadened at base, oblong-lanceolate, pinnatisect, lateral lobes
sessile, short-triangular, large spinose-toothed, approximate; lower cauline
leaves like basal; middle leaves sessile, short-decurrent, lanceolate,
pinnatipartite into triangular, spinose-toothed lateral lobes; upper leaves
sessile, auriculate, oblong-ovate or ovate, acuminate-spinescent with small
spinescent lobes. Capitula globose, large, about 30 mm wide (excluding
cusps), densely arachnoid-hairy. Involucral bracts about 100, relatively
weak, coriaceous, somewhat flat; outer and middle bracts narrow-
lanceolate, acuminate-short spinescent, closely appressed at base,
arcuately decurved from middle, 10-15 mm long, 3 mm wide at base;
subsequent bracts erect, narrow lanceolate, acuminate-spinescent, about
25 mm long and 2.5 mm wide; inner bracts lanceolate, with large up to
15 mm long and 5 mm wide, membranous, acuminate-spinescent, apical
appendages forming “corona;” innermost bracts membranous, linear, 27
mm long; receptacular bristles scabrous, gradually broadened toward
apex. Corolla purple, usually not exserted. Achenes obovoid, weakly
compressed, smooth, lustrous, roundish above, grayish with dark spots,
about 6 mm long. Flowering V—VI; fruiting VI—VII.

Stony-gravelly slopes in the upper part of middle mountain zone.—
Soviet Central Asia: Tien Shan (western). Endemic. Described from the
Angren River Basin, near the village of Baksuk. Type in Tashkent.

209. C. rosea Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIIIc, VI
(1929) 17, cum tab.; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8,
558.—C. schtschurowskiana B. polylepis Bornm. in Beih. bot. Centralbl.
XXXIV, 2 (1916) 181.—TIe.: Kult. op. cit. Plate 17—Exs.: H.F.A.M.
No. 486.

Biennial. Stem 15—35(45) cm high, striate-sulcate, branched from
base, less often from middle; branches one-headed, leafy. Leaves green,
with prominent veins, glabrous above, gray-tomentose beneath, spinose-
toothed; basal leaves broad-petiolate, somewhat lyrate, with large ovate-
orbicular terminal lobe and small, broadly sessile, with one or two pairs
of lateral lobes; lower cauline leaves like basal but short acuminate;
middle and upper leaves sessile, ovate-orbicular, with or without auricles,
large spinose-toothed, short acuminate in stiff spine; uppermost leaves
sometimes strongly reduced. Capitula flat-globose, arachnoid-hairy, 25—
30-40 mm wide (excluding cusps). Involucral bracts numerous, more
than 100, with long falcate cusp; outer and subsequent bracts carinate,
subulate-triquetrous; middle bracts like outer but erect-spreading; inner
bracts membranous, about 40 mm long, lanceolate, acuminate, purple,

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302

forming “corona,” exceeding adjacent bracts by 20-25 mm; receptacular
bristles scabrous. Corolla purple, not exserted from corona. Achenes
obovoid, glabrous, smooth, about 6 mm long. Flowering V—VI-VII;
fruiting VII-VIII.

Outcrops of varicolored rocks from the foothills to middle mountain
zone.—Soviet Central Asia: Pamiro-Alai (southwestern spurs of the Gissar
Range). Endemic. Described from the vicinity of Yakkabag, Bovashada.
Type in Tashkent.

210. C. rhodantha Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIIIc, VI
(1929) 22; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 558.

Biennial. Stem 22-25 cm high, striate-sulcate, paritally arachnoid-
hairy, branched from middle; branches one-headed leafy. Leaves light
green, with prominent veins, glabrous above, fine tomentose beneath,
large spinose-toothed; basal leaves petiolate, lanceolate, pinnatisect, with
spinescent apical lobe, as large as lateral lobes; lower cauline leaves
like basal; middle leaves pinnatipartite, sessile, reduced; upper leaves
lobed, sessile, uppermost strongly reduced. Capitula ovoid-globose,
somewhat compressed near “corona,” arachnoid-hairy, 25-30 mm wide
(excluding cusps). Involucral bracts numerous (more than 100), carinate,
weak; outer bracts narrow lanceolate, 15 mm long, gradually attenuate
into thin, decurved cusp; middle bracts like outer but falcately decurved,
25-27 mm long, about 2 mm wide at base; subsequent bracts more flat,
lanceolate, acuminate, 15 mm long, erect; inner bracts about 40 mm
long, lanceolate, membranous above, purple, forming “corona,” exceeding
adjacent bracts by 15—20 mm; receptacular bristles scabrous. Corolla
pink, not exserted. Achenes obovoid, roundish above, glabrous, smooth,
about 4 mm long. Flowering VI; fruiting VII.

Clayey slopes in foothills——Soviet Central Asia: panel Alai
(southwestern spurs of the Gissar Range). Endemic. Described from the
vicinity of Baisun. Type in Tashkent.

Note. A poorly studied species, known only on the basis of type
specimens.

211. C. rotundifolia Winkl. in Tr. Peterb. Bot. Sada, X (1889)
579; XII, 252; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 558—C.
rubrarenosa Kult. in Sched. Herb. Fl. As. Med. 10 (1928) Nos. 486
nom. nud.; Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIIIc, VI (1929)
18.—C. kultiasowii Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3
(1936) 317; Juz. op. cit. 558.

Biennial. Stem 30-50 cm high, striate-sulcate, branched from base,
less often from middle; branches one-headed. Leaves dark green, with
prominent veins, glabrous above, gray-tomentose beneath, short spinose-

303

- toothed; basal leaves long-petiolate, lyrate with large, rotund, cordate
' terminal lobe, and one or occasionally two pairs of almost rudimentary

308

lateral lobes; lower cauline leaves either lyrate, with small terminal lobe
and one or two pairs of remote, acuminate-spinescent lateral lobes, or
somewhat lobed, only at base, upper part of leaf rotund, undivided,
short spinose-toothed; middle leaves broadly ovate or obovate, sessile,
almost cordate, weakly amplexicaul, auriculate, acuminate-spinescent;
upper cauline leaves reduced, sessile, round above, spinescent, uppermost
often reduced to spiny scales. Capitula flat-ovoid, constricted- below
“corona,” arachnoid-hairy, 20-25 mm wide (excluding cusps). Involucral
bracts relatively less numerous (about 60), with squarrose (decurved in
outer bracts) short and broad cusp; outer bracts stiff, carinate, subulate-
triquetrous; middle bracts like outer but two times as large and only
somewhat decurved; subsequent bracts somewhat flat, erect, acuminate-
spinescent; inner bracts membranous, lanceolate, acuminate, weakly
carinate dorsally, purple, forming “corona,” almost as long as florets;
receptacular bristles scabrous. Corolla pale pink or white. Achenes
obovoid, dark gray with light brown dots, glabrous, smooth. Flowering
V-VI-VII; fruiting VII-—VIII.

Clayey slopes with outcrops of gypsum in varicolored lower
mountains.—Soviet Central Asia: Pamiro-Alai (south). Endemic.
Described from the Tashbulak Mountains near Sangtud. Type in
Leningrad.

212. C. ferruginea Kult. in Bot. Mat. Gerb. Glavn. Bot. Sada, V,
8-9 (1924) 142; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 559.—
C. truncata Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3 (1936)
318; Juz. op. cit. 558.

Biennial. Stem erect, 20-40 cm high, usually branched from base
or middle, leafy, arachnoid-hairy, glabrescent; branches one-headed.
Leaves almost coriaceous, green, arachnoid-hairy on both sides, more
densely beneath; basal leaves petiolate, lanceolate, pinnatipartite in
triangular, acuminate-spinescent lateral lobes; lower cauline leaves like
basal but sessile; middle leaves oblong-ovate or ovate, gradually reduced
toward stem apex, sessile, very short-decurrent, auriculate, large spinose-
toothed, attenuate, cuspidate, almost horizontal; uppermost leaves strongly
reduced. Capitula ovoid-globose, 15-30 mm wide (excluding cusps),
densely rusty-tomentose, sharply truncate or even foveate at base.
Involucral bracts numerous, about 100, all except inner narrow-lanceolate,
short-acuminate, arcuately decurved; inner bracts membranous, whitish
above, lanceolate, scabrous along margin, erect, about 20 mm long.
Corolla purple; receptacular bristles scabrous. Achenes obovoid, smooth,
about 4 mm long. Flowering VII—VIII; fruiting VIII-IX.

309

310

304

Gravelly slopes in middle mountain zone.—Soviet Central Asia:
Pamiro-Alai (eastern part of the Zeravshan and Gissar ranges, Peter the
Great Range). Endemic. Described from Peter the Great Range from the
Gardani-Kaftar Pass. Type in Leningrad.

213. C. dimoana Kult. in Pochv. i Bot.-Geogr. Iss]. Bass. rr Amu-
i Syr-Dari, 2 (1916) 79; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8,
558.—Ie.: Kult. op. cit. Plate 25.

Biennial. Stem 30 cm high, erect, striate-sulcate, whitish, lustrous,
branched from base; branches one-headed, leafy. Leaves coriaceous,
glabrous above, finely arachnoid-hairy beneath with prominent veins,
terminating in stiff spines; basal leaves obovate, short-petiolate, large
spinose-toothed; lower cauline leaves like basal; middle and upper cauline
leaves oval, sessile, broadly short-decurrent, spinose-toothed; uppermost
leaves reduced. Capitula globose, 20-25 mm wide (excluding cusps),
somewhat arachnoid-hairy. Involucral bracts numerous, weak, narrow-
lanceolate, acuminate-spinescent, carinate, falcately recurved; inner bracts
linear, long-acuminate, about 30 mm long, forming somewhat short (10—
15 mm long) corona above capitulum. Corolla yellow; receptacular
bristles scabrous. Achenes obovoid, glabrous, smooth, about 5 mm long.
Flowering V; fruiting V.

Outcrops of varicolored rocks.—Soviet Central Asia: Pamiro-Alai
(Kugitang). Endemic. Described from Kugitang (village of Khodzhafilata).
Type in Tashkent.

214. C. darwasica Winkl. in Tr. Peterb. Bot. Sada, XII (1892) 249;
XIV, 207; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 557.

Biennial? Stem erect, striate-sulcate, arachnoid-hairy, leafy, branched
from base, 30-35 high. Leaves greenish-gray or greenish above, finely
flocculose-arachnoid hairy, gray-tomentose beneath; basal and lower
cauline leaves petiolate, petioles broadened at base, oblong-lanceolate,
almost pinnatisect, with remote, large spinose-toothed, broadly lanceolate
lateral lobes, large, 20-30 cm long; other cauline leaves strongly reduced,
lanceolate, sessile, semiamplexicaul, large spinose-toothed, acuminate-
spinescent, 2-5 cm long; uppermost leaves strongly reduced. Capitula
globose, 12-13 mm wide (excluding cusps), loosely arachnoid-hairy,
with many florets. Involucral bracts about 50; outer bracts 6-7 mm
long, deflexed, narrow-lanceolate, acuminate-spinescent; middle bracts
erect-spreading, about 17 mm long, somewhat attenuate at deflection,
carinate; inner bracts membranous, flat, lanceolate, erect, 17-20 mm
long, short-acuminate; receptacular bristles scabrous. Corolla pale pink.
Achenes obovoid, light gray, glabrous, smooth, about 4 mm long,
roundish above. Flowering VIII; fruiting IX.

305

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Plate XXIV.

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311

is Tschern. and Vved.;
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312

306

In middle mountain zoné.—Soviet Central Asia: Pamiro-Alai
(Shugnan). General distribution: Afghanistan. Described from Afghanistan
(near village of Omar on the left bank of the Pyandzh River). Type in
Leningrad.

Note. A poorly studied species, known so far from three localities,
two of which are in the Gunt River Basin.

215. C. integrifolia Franch. in Ann. Sc. Nat. 6 sér. 16 (1883) 322;
Winkl. in Tr. Peterb. Bot. Sada, XII, 244; XIV, 228; Bornm. in Beih
bot. Centralbl. XXXIV, 2, 162; Juz. in Tr. Tadzh. Bazy Akad. Nauk
SSSR, 8, 559.—C. integrifolia B. heterophylla Bornm. |.c.—Arctium
integrifolium O. Ktze. Rev. Gen. 1 (1891) 308.—Exs.: H.F.A.M. No.
482.

Biennial. Stem 15-50 cm high, erect, usually branched in upper
half, striate-sulcate, arachnoid-hairy; branches one-headed,. leafy, mostly
glabrescent toward end of vegetative growth. Leaves soft, without spines,
entire, decurrent, finely arachnoid-hairy above, finely tomentose beneath;
basal leaves lyrate, less often undivided or lobed, lower cauline leaves
like basal; middle leaves broad lanceolate, acuminate, less often obtuse;
upper leaves oblong; uppermost reduced. Capitula globose, 18—20 mm
wide (excluding cusps), arachnoid-hairy. Involucral bracts numerous,
relatively weak, up to 2 mm wide, with broad, linear basal cusp;
outermost bracts slightly decurved, all others squarrose. Corolla pink;
receptacular bristles scabrous. Achenes obovoid, glabrous, smooth, about
4 mm long. Flowering V—VI-VII; fruiting VII—VIII.

Loamy-stony slopes in middle mountain zone.—Soviet Central Asia:
Pamiro-Alai. Endemic. Described from the Zeravshan River Valley,
Chukalik. Type in Paris.

216. C. divaricata Winkl. in Tr. Peterb. Bot. Sada, XI (1892) 334;
XII, 267; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 559.

Perennial (monocarpic?). Stem erect, 20-35 cm high finely flocculose-
arachnoid hairy, divaricately dichotomous, leafy, branches and branchlets
slender, one-headed. Leaves green and finely arachnoid-hairy above, gray-
tomentose beneath; basal leaves petiolate, lyrate, with ovate, small spinose-
toothed terminal lobe and broadly triangular, acuminate-spinescent, small
spinose-toothed lateral lobes; cauline leaves sessile, semiamplexicaul, with
roundish auricles, ovate-lanceolate, crenate-spinose-toothed, gradually
reduced upward. Capitula ovoid-cylindrical, arachnoid-hairy, 7-11 mm
wide (excluding cusps). Involucral bracts numerous (about 80); outer
bracts ovate, basally appressed, somewhat attenuate above at transition
to cusp, with small thickening on inner side, gradually narrowed in
linear, somewhat flat, decurved, 5-7 mm long cusp; middle bracts like
outer but longer, with wider and longer (15—22 mm long) decurved or

313

307

almost horizontal cusps; subsequent bracts with short (4-5 mm long)
falcate cusps; inner bracts membranous, sometimes purple on outer side,
broad-lanceolate, short-acuminate, smooth along margin; receptacular
bristles glabrous, smooth. Corolla pale purple; anther tube purple.
Achenes three-angled, thickened at angles, with barely noticeable teeth
above, dark gray, 3 mm long and 1.5 mm wide. Flowering VII—VIII;
fruiting VIII.

On gravelly slopes.—Soviet Central Asia: Pamiro-Alai (Darvaz).
Endemic. Described from the Pyandzh River Basin, near Kalaikhumb.
Type in Leningrad.

217. C. centauroides Fisch. and Mey. in Bull. Soc. Nat. Mosc. XII
(1839) 159, (nomen) and ex Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2
(1865) 54; exclus. syn. C. platyacantha; Boiss. Fl. or. II, 493 exclus.
syn. P. platyacantha; Winkl. in Tr. Peterb. Bot. Sada, XII, 236; Kult. in
Fl. Turkm. VII, 206.—Exs.: H.F.A.M. No. 44.—Ie.: Fl. Turkm. VII,
Plate 31.

Perennial (rather monocarpic). Stem with remains of old petioles at
base, distinctly tomentose in-between, erect, 40-50 cm high, usually
divaricately branched from middle, striate-sulcate, partially tomentose
below, slightly glossy, entirely glabrous above, repeatedly branched,
branchlets one-headed. Leaves coriaceous, green and finely loose-
arachnoid hairy above, gray-tomentose beneath, with prominent veins;
basal and lower cauline leaves somewhat short-petiolate, oblong, 4—18
cm long and 2—4 cm wide, unequally pinnatipartite or pinnatilobate,
with spinose-toothed and acuminate-spinescent lateral lobes; cauline
leaves sessile, semiamplexicaul, gradually reduced, oblong-oval or
lanceolate, auriculate, crenate-spinose-toothed, acuminate-spinescent;
uppermost leaves strongly reduced. Capitula ovoid-cylindrical, 10 mm
wide, arachnoid hairy. Involucral bracts about 50, all glandular and
arachnoid-hairy dorsally; outer bracts closely appressed at base,
broadened, gradually acuminate above in stiff triquetrous-subulate,
decurved, 7-8 mm long cusp; middle bracts longest, about 30 mm long,
broadened at base (5 x 4 mm), closely appressed, weakly attenuate above
in carinate-grooved, linear, acuminate-spinescent, 22-24 mm long,
divergent cusps; inner bracts thin coriaceous, linear-lanceolate, acuminate,
scabrous along margin and dorsally, 20-24 mm long; receptacular bristles
smooth; somewhat broadened above. Corolla yellow; anther tube purple.
Achenes obpyramidal, 5 mm long, distinctly ribbed, toothed above.
Flowering VI-VII; fruiting VII.

Slopes in middle mountain zone.—Soviet Central Asia: mountainous
Turkmenia (Great Balkans). Endemic. Described from the Great Balkans.
Type in Leningrad.

314

308

Subsection 3. Kokuria Juz.—Involucral bracts numerous or
comparatively less numerous, long or short acuminate; inner bracts often
with membranous appendages, but not forming “corona;” achenes
glabrous, smooth; receptacular bristles scabrous, less often smooth.
Biennial plants with uniformly or discontinuously winged stems.

Series 1. Pulchellae Juz.—Capitula ovoid or globose-oblong;
involucral bracts numerous, long-acuminate; receptacular bristles
scabrous; basal leaves soft, undivided.

218. C. severtzovii Rgl. in Bull. Soc. Nat. Mosc. XI, 2 (1867) 151;
Winkl. in Tr. Peterb. Bot. Sada, XII, 244 p. p.; XIV, 228; Bornm. in
Beih. bot. Centralbl. XXXIV, 2, 163; Juz. in Tr. Tadzh. Bazy Akad.
Nauk. SSSR, 8, 560.—Arctium sewertzowii O. Ktze. Rev. Gen. 1 (1891)
308.—Exs.: H.F.A.M. No. 487.

Biennial. Stem 30-80 cm high, erect, branched in upper half, more
or less uniformly winged throughout or almost throughout, wings up to
5 mm wide, spinose-toothed. Leaves soft, undivided, very finely
arachnoid-hairy above, finely-tomentose beneath, acuminate; basal leaves
broadly lanceolate, narrowed toward both ends, petiolate; lower cauline
leaves like basal; middle and upper cauline leaves sessile, decurrent,
lanceolate, spinose-toothed; uppermost leaves reduced. Capitula globose-
oblong, 12-15 mm wide (excluding cusps), arachnoid-hairy. Involucral
bracts very numerous, narrow, with thin subulate cusp, squarrose; inner
bracts with narrow-ovate, up to 1 mm wide (in broadest part), gradually
acuminate appendages, not exceeding adjoining middle bracts, usually
dark-colored; receptacular bristles scabrous. Corolla pink. Achenes
obovoid, blackish, glabrous, smooth, lustrous, 3.5—-4 mm long. Flowering
VI-VII; fruiting VIJ-VIII. (Plate XXV, Fig. 1.)

Stony-gravelly slopes in the foothills—Soviet Central Asia:
Tien Shan (western); Pamiro-Alai. Endemic. Described from the
Chirchik River Valley, based on the collections of Severtsev. Type in
Leningrad.

Note. A hybrid has been described under the name C. cana Juz. in
Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3 (1936) 316.—C. ninae Juz. x
C. severtzovii Rgl. from the Alai Range: Taldyk River between Mady
and Lyangar.

Morphologically, it closely resembles C. severtzovii but has more
densely pubescent leaves and stem, broader and less numerous outer
and middle involucral bracts, and inner bracts broadened into ovate-
lanceolate, usually purple, somewhat short appendages.

The numerous hybrids of C. severtzovii and C. microcarpa, as well
as of C. severtzovii and C. arachnoidea, which approach one or the
other species, do not have names.

315

309

219. C. leiocephala (Rgl.) Juz. in Tr. Bot. Inst. Akad. Nauk SSSR,
Ser. 1, 3 (1936) 314.—C. sewertzowii var. leiocephala Rgl. in Tr. Peterb.
Bot. Sada, VI (1880).314; Winkl., ibid. XII (1892) 245; Winkl., ibid.
XIV (1897) 228 p. p.; Bornm. in Beih. bot. Centralbl. XXXIV, 2, 163.

Very similar to C. severtzovii Rgl. but differs from it in the smaller
capitula, almost entirely glabrous involucres and broader (up to 2.5 mm
wide) and usually colored and white-bordered appendages of the inner
involucral bracts. Flowering VI-—VII; fruiting VII—VIII.

Stony or loamy mountain slopes, in juniper forests.—Soviet Central
Asia: Tien Shan (central Tien Shan). General distribution: Dzhungaria-
Kashgaria. Described from the Chirchik River Valley (?). Type in
Leningrad.

Note. Since S.V. Juzepczuk did not designate the type while
describing the species but included the variety of Regel among the
synonyms, the plant of Krause should be considered the lectotype, since
Regel established the variety. As indicated on the label, this plant was
collected in the Chirchik River Valley. However, the range of the species,
as presently described on the basis of numerous collections, provides a
basis to conclude that the plant was collected in Mailisai (foothills of
the Chatkal Range), where Krause also made collections.

220. C. pulchella Bge. in Delect. Sem. Hort. Dorpat. (1845) 3;
Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2, 44; Boiss. Fl. or. IL, 494;
Winkl. in Tr. Peterb. Bot. Sada, XII, 245; XIV, 228; Bornm. in Beih.
bot. Centralbl. XXXIV, 2, 163; Juz. in Tr. Tadzh. Bazy Akad. Nauk
SSSR, 8, 560.—C. pulchella var. latifolia Winkl. op. cit. (1892) 246.—
Arctium pulchellum O. Ktze. Rev. Gen. 1 (1891) 308.

Biennial. Stem 25-60 cm high, erect, branched in upper half, more
or less uniformly winged throughout: on cauline leaves 4-5 mm wide,
on peduncules 2-3 mm wide, finely spinose-toothed. Leaves soft,
undivided, finely spinose-toothed, subglabrous or very finely arachnoid-
hairy on both sides, obtuse; basal leaves broadly lanceolate, petiolate;
lower cauline leaves like basal; middle and upper leaves broadly
lanceolate, decurrent; uppermost leaves oblong, reduced. Capitula ovoid,
8-10 mm wide (excluding cusps), arachnoid-hairy. Involucral bracts very
numerous, narrow, with thin subulate cusp, squarrose, inner bracts with
broad ovate, up to 4 mm wide, abruptly acuminate apical appendage,
not exceeding adjacent middle bracts; receptacular bristles scabrous.
Corolla pink. Achenes narrow-obovoid, light gray, glabrous, smooth, 3
mm long. Flowering VI—-VII; fruiting VIJ-—VIII.

Loamy slopes in middle mountain zone.—Soviet Central Asia:
Pamiro-Alai. Endemic. Described from the Karatau Range, near
Samarkand, i.e., from the southern slopes of the Gissar Range. Type in
Paris; cotype in Leningrad.

316

310

Note. Hybrids of C. outichaschensis Franch. and C. pulchella Bge.,
have been described as C. heterogenetos Bornm.; cf. the note to C.
outichaschensis Franch. about them.

Series 2. Pauciramosae Tschern.—Capitula almost globose;
involucral bracts numerous, carinate-grooved, long acuminate-spinescent;
receptacular bristles scabrous; basal leaves soft, pinnatisect.

221. C. pauciramosa Kult. in Bot. Mat. Gerb. Glavn. Bot. Sada,
V, 8-9 (1924) 139. .

Biennial. Stem solitary, erect, 25-40 cm high, shallow-sulcate,
flocculose-arachnoid hairy, densely leafy, discontinuously winged, more
or less branched in upper half; branches short, divergent, leafy, one-headed.
Leaves soft, without spines, early withering, but persistent, arachnoid
hairy-tomentose on both sides, revolute, broadly lanceolate; basal leaves
petiolate, pinnatipartite, lateral segments very soft, unequal, alternate,
linear, short-acuminate, sessile, two-toothed at base or slightly above,
apically linear-excurrent, petiole wide-winged; cauline leaves gradually
reduced, like basal but sessile, short-decurrent in crenate-toothed cauline
wings. Leaves on branches oblong-narrow lanceolate. Capitula 40 mm
long and 22 mm wide (excluding cusps), broadly ovoid or subglobose,
densely arachnoid-hairy, with many florets. Involucral bracts numerous,
carinate-grooved; outer bracts narrow-lanceolate, long acuminate into
triquetrous spine, decurved, 17-18 mm long, up to 2 mm wide at base;
middle bracts like outer, closely appressed at base, acuminate, divergent,
up to 30 mm long; subsequent bracts more flat, erect, 23-25 mm long;
inner bracts thin-coriaceous, somewhat flat, narrow-lanceolate, short-
acuminate, up to 25 mm long; subsequent bracts almost scarious, with

slightly broadened, acuminate, apical appendages; innermost bracts

membranous, linear; receptacular bristles finely scabrous above. Florets
exserted; corolla pink. Achenes oblong-obovoid, roundish above, without
teeth, 44.5 mm long. Flowering VI-—VII; fruiting VII-VIII.

Gravelly slopes in middle mountain zone.—Soviet Central Asia: Tien
Shan (Mogoltau). Endemic. Described from the Mogoltau Mountains.
Type in Tashkent.

Series 3. Regelianae Juz—Capitula flat-globose; involucral bracts
less numerous, short-acuminate; receptacular bristles scabrous; basal
leaves soft, undivided, entire.

222. C. regelii Winkl. in Tr. Peterb. Bot. Sada, X (1889) 574;
Winkl. in Tr. Peterb. Bot. Sada, XII, 252; XIV, 229; Bornm. in Beith.
bot. Centralbl. XXXIV, 2, 171; Juz. in Tr. Tadzh. Bazy Akad. Nauk

317

311

SSSR, 8, 560.—C. centauroides auct. non Fisch. and Mey.: Rgl. in
Bull. Soc. Nat. Mosc. XL, 2 (1867) 152.

Biennial. Stem 15-30 cm high, erect, branched almost from middle,
arachnoid-hairy, more or less winged throughout, wings 3—7 mm wide,
entire; branches leafy, one-headed. Leaves soft, undivided, gray-
tomentose on both sides; basal leaves petiolate, with winged petioles;
cauline leaves sessile, decurrent, oblong-ovate. Capitula flat-globose, 20—
25 mm wide, arachnoid-hairy. Involucral bracts fewer, wide, 5-7 mm -
wide at base, broadly lanceolate, short cuspidate, somewhat flat,
squarrose; inner bracts spatulate, with subobtuse or obtuse appendages
forming corona; receptacular bristles scabrous. Corolla greenish-yellow.
Achenes ovoid, roundish above, glabrous, smooth, lustrous, 3.5 mm long
and 2 mm wide. Flowering V—VII; fruiting VII—VIII.

Gravelly slopes in foothills—Soviet Central Asia: Syr-Darya, Tien
Shan (western), Pamiro-Alai. Endemic. Described from western Tien
Shan, between Boroldai and Bugun. Type in Leningrad.

Series 4. Arachnoideae Juz.—Capitula globose or broadly ovoid;
involucral bracts numerous or relatively less numerous; lanceolate, short-
acuminate; receptacular bristles scabrous; basal leaves withering early,
pinnatisect, lyrate-pinnatilobate or spinose-toothed.

223. C. arachnoidea Fisch. and Mey. in DC. Prodr. VI (1837) 553;
Ldb. Fl. Ross. II, 677; Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2, 44;
Winkl. in Tr. Peterb. Bot. Sada, XII, 247 p. p.; Bornm. in Beih. bot.
Centralbl. XXXIV, 2, 167—C. arachnoidea var. typica and var. spinosior
Rgl. in Bull. Soc. Nat. Mosc. XL, 2 (1867) 151.

Biennial. Stem erect, 20-60 cm high, usually corymbosely branched
above, arachnoid-hairy, usually winged throughout, wings 3-8 mm wide,
crenate-spinose-toothed; branches often short, narrow-winged. Leaves and
wings of main stem and branches finely arachnoid-hairy above, gray-
tomentose beneath; basal and lower cauline leaves oblong-ovate, with
narrow-winged petiole, sparsely short spinose-toothed, withering early;
other cauline leaves oblong-lanceolate, sessile, decurrent, sparsely and
short spinose-toothed, gradually reduced toward stem apex. Capitula
globose, arachnoid-hairy, 10-15 mm wide (excluding cusps). Involucral
bracts numerous, narrow-lanceolate, about 1 mm wide, short-acuminate;
outer bracts usually decurved, others erect-spreading, 7-15 mm long;
inner bracts 15—18 mm long, lanceolate, membranous, usually brownish,
lighter in color and scabrous along margin, acuminate. Corolla pink,
exserted; receptacular bristles scabrous; insignificantly broadened above.
Achenes narrow obovoid, 3 mm long and 1.5 mm wide, glabrous, smooth,
dark gray. Flowering V—VI.

318

312

Foothills, on clayey slopes, apparently as weed.—Soviet Central
Asia: Balkhash Region, Tien Shan. Endemic. Described from the Urdzhar
River Valley. Type in Leningrad.

The species is closest to C. microcarpa Boiss. Its characters are
intermediate between the latter and C. severtzovii Rgl. Hybrids of this
species with C. severtzovii Rgl. are known.

224. C. microcarpa Boiss. Diagn. pl. nov. -2 (1843) 59; Bge. in
Mém. Acad. Pétersb. 7 sér. 9, 2, 44; Boiss. Fl. or. III, 496; Winkl. in
Tr. Peterb. Bot. Sada, XII, 246; XIV, 228; Bornm. in Beih. bot. Centralbl.
XXXIV, 2, 165; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 560;
Kult. in Fl. Turkm. VII, 190.—C. xeranthemoides Bge. l|.c. in syn.—
Arctium microcarpum O. Ktze. in Tr. Peterb. Bot. Sada, X (1887) 197.—
Exs.: H.F.A.M. No. 47.

Biennial. Stem 25-60 cm high, erect, discontinuously broad-winged,
often branched from middle; branches one-headed; stem wings 5—7 mm
wide, spinose-toothed or almost pinnatipartite. Leaves coriaceous, strongly
spinose-toothed, green, finely gray-tomentose beneath, loosely arachnoid-
hairy above; basal leaves lyrate-pinnatilobate; cauline leaves long-
decurrent, oblong, almost lobed; lobes finely short-spinescent; uppermost
cauline leaves reduced. Capitula broad-ovoid, with many florets, 15—25
mm wide (excluding cusps), somewhat arachnoid-hairy. Involucral bracts
green, coriaceous, carinate, acuminate in stiff, short spine; outermost
bracts weakly decurved, 1 mm wide at base; outer and middle bracts
erect, 2-3 mm wide at base; inner and innermost bracts with broadly
ovoid or almost roundish, membranous, often pinkish-purple or pink
appendages; receptacular bristles scabrous. Corolla purple or pink,
exserted. Achenes oblong-obovoid, gray. Flowering V—VI; fruiting VI—
Vile...

Clayey and stony slopes in foothills and middle mountain zone,
often as weed.—Soviet Central Asia: Syr-Darya, Tien Shan (very widely
distributed in western Tien Shan), Pamiro-Alai, mountainous Turkmenia
(Kopetdag, Badkhyz). General distribution: Iran, Afghanistan. Described
from Kabul. Type in Geneva. .

Note. The species is very widely distributed and often grows as a
weed. It forms numerous hybrids with C. outichaschensis, C. sewertzovii,
C. pulchella, C. submutica and C. coronata. One of the hybrids was
described as C. atripurpurea Juz. (in Tr. Bot. Inst. Akad. Nauk SSSR,
Ser. 1, 3, 1936, 313 = C. coronata Franch. x C. microcarpa Boiss.)
from the Kugitang Range (in the ravine opposite the village of Kugitang).
These plants have short-excurrent leaves, with involucral bracts broader
than in C. coronata and narrower than in C. microcarpa, the inner bracts
widening into dark purple appendages, forming a short “corona.” See
notes to the above-named species for other hybrids.

319

313

225. C. ninae Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3
(1936) 315; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 561.—C.
microcarpa var. parviceps Bornm. in Beih. bot. Centralbl. XXXIV, 2
(1916) 166, saltem p. max. p.

Biennial. The plant occupies intermediate position between C.
microcarpa Boiss. and C. arachnoidea Fisch. and Mey. It differs from
the former by the presence of gray-tomentose pubescence of the entire
plant; narrower cauline leaves; smaller (often almost half), 18-28 mm
long and 10-15 mm wide (excluding cusps) capitula; narrower and shorter
involucral bracts, of which inner ones widening into an ovate whitish or
pale brown thin short cuspidate appendage; from C. arachnoidea it differs
by broader and less numerous (almost half) involucral bracts. Flowering
V; fruiting VII.

Mountain slopes, pebble land.—Soviet Central Asia: Pamiro-Alai
(Alai Range), Tien Shan (western). Endemic. Described from the left
bank of the Taldyk River, between the villages of Mady and Lyangar.
Type in Leningrad.

Note. The species needs to be studied further, as the boundaries of
its range are still unclear.

Series 5. Platylepides Juz—Capitula globose; involucral bracts less
numerous, somewhat flat, short-acuminate; receptacular bristles smooth;
basal leaves soft, with remote lobes.

226. C. platylepis Schrenk ex Fisch. and Mey. Ind. sem. hort. Petrop.
9 (1843) 10; Bull. phys.-math. Acad. Pétersb. 3, 108; Ldb. Fl. Ross. II,
681; Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2, 38; Boiss. Fl. or. III, 466;
Winkl. in Tr. Peterb. Bot. Sada, XII, 202; XIV, 219; Bornm. in Beih.
bot. Centralbl. XXXIV, 2, 150; Pavlov in Fl. Tsentr. Kazakhst. III, 296;
Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 561.—Arctium platylepis
O. Ktze. Rev. Gen. 1 (1891) 308.

Biennial. Stem 30-85 cm high, erect, branched from middle,
appressed arachnoid-hairy, more or less winged throughout, wings
crenate-toothed, spinose, 3—7 mm wide, gray-tomentose; inflorescence
branches strongly elongate. Leaves soft, lanceolate, crenate-toothed, gray
arachnoid-hairy above, gray-tomentose beneath; basal leaves oblong,
petiolate, sparsely lobed; lower cauline leaves like basal, but sessile;
middle and upper leaves narrow, reduced, sessile, with remote lobes,
lobes acuminate-spinescent; uppermost leaves strongly reduced. Capitula
globose, small, 7-9 mm wide (excluding cusps), arachnoid hairy-woolly.
Involucral bracts fewer, 31-36, green, short-lanceolate, carinate,
squarrose, acuminate-spinescent; middle bracts somewhat flat, roundish
above, short-acuminate; inner bracts with obtusely broadened, unarmed

320

314

or with short-acuminate membranous, 5 mm wide appendages; innermost
bracts with membranous, lanceolate, up to 2 mm wide appendages;
receptacular bristles smooth. Corolla pink. Achenes obovoid or almost
obpyramidal, with scarcely noticeable longitudinal ribs, roundish above,
3 mm long and 1.5 mm wide, light gray. Flowering VI—VII; fruiting
VI-VIII.

Clayey mounds in foothilis—Soviet Central Asia: Aralo-Caspian
Region, Balkhash Region, Syr-Darya, Tien Shan, Pamiro-Alai. General
distribution: Dzhungaria. Described from the Khantau Mountains. Type
in Leningrad.

227. C. ortholepis Juz. ex Tschern. Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 99.

Biennial. Stem 70 cm high, erect, appressed arachnoid-hairy,
branched from middle, more or less winged throughout, wings crenate-
toothed, spinose, 5-7 mm wide, gray-tomentose; lateral branches short.
Leaves oblong, finely spinose-toothed, gray-tomentose on both sides;
middle and upper leaves sessile; uppermost leaves reduced. Capitula
obconical, 10-12 mm wide (excluding cusps), arachnoid hairy-woolly.
Involucral bracts less numerous, 25-30, carinate, squarrose acuminate-
spinescent; outermost bracts appressed, up to 10 mm long, outer bracts
stiff, up to 20 mm long; middle bracts somewhat flat, acuminate; inner
bracts with obtusely broadened, short cusp, membranous, up to 3 mm
wide, innermost with lanceolate, up to 2 mm wide appendages;
receptacular bristles smooth; corolla pink. Achenes obovoid, 4 mm long,
light gray with dark spots. Flowering VII—VIII; fruiting VIII.

As weed.—Soviet Central Asia: Pamiro-Alai (western spurs of the
Gissar Range). Endemic. Described from the vicinity of the town of
Guzar. Type in Tashkent.

Subsection 4. Subappendiculatae Tschern. in Bot. Mat. Gerb. Inst.
Bot. Akad. Nauk UzbSSR, XVII (1962).—Involucral bracts less
numerous, carinate, constricted in middle, short acuminate or broad above
and short acuminate; inner bracts membranous, usually forming short
“corona;” receptacular bristles smooth or scabrous. Biennial plants with
wingless stems.

228. C. ferganensis Bornm. in Beih. bot. Centralbl. XXXIV, 2
(1916) 195 (“ferghanensis”).—C. popovii Kult. in Tr. Turk. Gos. Univ.
4 (1922) 66.

Biennial. Stem 30-50 cm high, erect, striate-sulcate, partially
arachnoid-hairy, branched from base or middle; branches one-headed,
leafy. Leaves coriaceous with prominent veins, large spinose-toothed,

A15

Plate XXV.
1—Cousinia severtzovii Rgl.; 2—C. peduncularis Juz.

321

322

316

gray-tomentose beneath, green and finely arachnoid-hairy above; basal
leaves lyrate, with large ovate terminal lobe and rotund, broadly sessile
lateral lobes alternating with smaller sessile lobes; lower cauline leaves
oblong, almost lobate, lobes stiff spinescent; middle leaves sessile,
semiamplexicaul, large spinose-toothed, terminating in long, stiff, weakly
curved spine; upper leaves like middle but with more prominent midrib,
attenuate into divergent stiff spine; uppermost leaves reduced, sometimes
into spines. Capitula ovoid-globose, constricted below, corona of inner
involucral bracts, 20-25 mm wide (excluding cusps), weakly arachnoid-
hairy. Involucral bracts green, carinate, constricted in middle; outer bracts
decurved, relatively long acuminate; subsequent bracts squarrose; middle
bracts coriaceous, more flat, roundish above, abruptly acuminate into
short spine, shorter than outer bracts; inner bracts membranous, linear-
lanceolate, two times as large as adjacent middle bracts, forming
“corona;” receptacular bristles scabrous. Corolla pink, usually not exserted
from corona. Achenes oblong-obovoid, dark gray, glabrous, lustrous,
roundish above, 5 mm long. Flowering V—VI-VII; fruiting VI-—VIII.

Stony-gravelly slopes in foothills and lower mountain zone.—Soviet
Central Asia: Pamiro-Alai (Turkestan Range). Endemic. Described from
the vicinity of Sultan-Murad. Type in Leningrad.

229. C. subappendiculata Kult. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, V, 8-9 (1924) 140; Juz. in Tr. Tadzh. Bazy Akad. Nauk
SSSR, 8, 561.

Biennial ? Stem erect, 35 cm high, herbaceous, branched from base,
striate-sulcate, weakly arachnoid-hairy; branches simple, leafy, one-
headed. Leaves almost coriaceous with prominent veins, green, finely
arachnoid-hairy on both sides (more dense beneath); basal leaves petiolate
(petioles somewhat broad, striate-sulcate), oblong-lanceolate, 18 cm long
and 4 cm wide, attenuate towards both ends, acuminate-spinescent,
pinnatisect into narrow triangular lobes, spinose-toothed; middle cauline
leaves 10 cm long and 5 cm wide, sessile, like basal; leaves of branches
sessile, ovate-oblong, acuminate-spinescent, crenate-spinose-toothed along
margin, upper leaves of branches linear, acuminate-spinescent, short
spinose-toothed. Capitula globose, 4 cm long and 5—6 cm wide (including
cusps; 2.5 cm wide excluding cusps), weakly and loosely arachnoid-
hairy. Involucral bracts green, somewhat arachnoid-hairy, scabrous below
along margin; outer bracts lanceolate, 13 mm long and 2 mm wide at
base, carinate, acuminate-spinescent; subsequent bracts 21 mm long and
4 mm wide, divergent, lanceolate, closely appressed at base, weakly
constricted in middle; next bracts 22 mm long, 2 mm wide in middle,
4 mm above, lanceolate, weakly constricted in middle, slightly broadened
above, acuminate-spinescent, divergent; middle bracts 18 mm long,

323

317

2 mm wide in middle, 3 mm above, slightly carinate, narrowed at base,
broadly linear, roundish-obtuse, acuminate-weakly spinescent; inner bracts
linear, 27 mm long and 2-2.5 mm wide, subglabrous, scabrous along
margin, membranous, dark brown above, acuminate with very short and
thin spine; innermost bracts 26 mm long and 2 mm wide, membranous,
narrow-linear; receptacular bristles scabrous above. Corolla pink, exserted
from corona; anther tube pink. Achenes obovoid, roundish above,
indistinctly striate, with dark spots, 4 mm long and 2 mm wide. Flowering
VII; fruiting VII.

On mountain slopes.—Soviet Central Asia: Pamiro-Alai (Alai
Range). Endemic. Described from Akbasaga in the upper reaches of the
Taldyk River. Type in Tashkent. (?).

Note. Material for this species is not available in the collections.
The type specimen of the species is apparently lost. Its original
description is presented here.

230. C. simulatrix Winkl. in Tr. Peterb. Bot. Sada, X (1889) 573;
XII, 271; Bornm. in Beih. bot. Centralbl. XXXIV, 2, 194; Juz. in Tr.
Tadzh. Bazy Akad. Nauk SSSR, 8, 561.

Biennial. Stem 25-50 cm high, striate-sulcate, arachnoid hairy-
flocculose, branched mainly from middle; branches one-headed, leafy.
Leaves whitish-woolly beneath, weakly arachnoid-hairy above, almost
entire; basal leaves petiolate, lyrate or irregularly pinnatipartite, with
larger roundish terminal lobe; lower cauline leaves lyrate or lobed, sessile;
middle and upper leaves sessile, amplexicaul, auriculate, short spinose-
toothed, more or less obtuse, abruptly terminating into short spine;
uppermost leaves reduced. Capitula globose, 25-35 mm wide (excluding
cusps), weakly arachnoid-hairy. Involucral bracts stiff, carinate, with
large apical spine, constricted in middle; outer and middle bracts
squarrose, slightly decurved, 7-8 mm wide at base; subsequent bracts
somewhat flat, somewhat shorter than middle, 6-7 mm wide at base,
constricted apically, with small stiff apical spine; inner bracts coriaceous,
lanceolate, acuminate, 5-6 mm wide at base; innermost: bracts linear,
receptacular bristles smooth. Corolla pink, prominent, numerous. Achenes
obovoid, roundish above, grayish-yellow, indistinctly nbbed, with very
narrow and discontinuously winged ribs, sometimes developed only as
discontinuously thickened ridges, 44.5 mm long and 2:5 mm wide.
Flowering VI—VII; fruiting VII—VIII.

Stony-gravelly slopes in middle mountain zone. = Suite Central
Asia: Pamiro-Alai (Gissar Range). Endemic. Described from*the Karatag
River Valley between Khoshgassan and Khakimi. Type in Leningrad.

231. C. vvedenskyi Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 106.

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318

Biennial. Stem 10—20(40) cm high, striate-sulcate, arachnoid-hairy,
branched from base and middle; branches with one or rarely two capitula,
leafy. Leaves whitish-woolly beneath, weakly arachnoid-hairy above,
finely spinose-toothed; basal leaves petiolate, lyrate, with ovate-rotund,
cordate terminal lobe, and remote ovate lateral lobes; lower cauline
leaves like basal but sessile or short-petiolate; middle leaves sessile,
oblong, semiamplexicaul, auriculate, finely spinose-toothed, acuminate-
spinescent; upper cauline leaves like middle, but reduced; uppermost
leaves sometimes reduced to spines. Capitula oblong-ovoid, narrowed
above, weakly arachnoid-hairy, 15—25 mm wide (excluding cusps).
Involucral bracts green, carinate, short-acuminate, constricted in middle;
outer bracts decurved; middle bracts squarrose, up to 7 mm wide at
base, constricted in middle, carinate, acuminate, spinescent; subsequent
bracts somewhat flat, weakly carinate, 5-6 mm wide at base, somewhat
constricted at apex, subobtuse, short-spinescent; inner bracts coriaceous,
green, weakly carinate, lanceolate; innermost bracts linear-lanceolate,
purple above; receptacular bristles smooth. Corolla pink, exserted from
corona. Achenes obovoid, roundish above. Flowering VI—VII; fruiting
VIII.

Gravelly-stony slopes in middle mountain zone.—Soviet Central
Asia: Pamiro-Alai (Chulbair and Baisuntau mountains). Endemic.
Described from the Chulbair Mountains, vicinity of the village of Seena.
Type in Tashkent.

Section 37. Microcousinia Tschern. in Bot. Mat. Gerb. Inst. Bot.
Akad. Nauk UzbSSR, XVII (1962) 86.—Constrictae Winkl. in Tr. Peterb.
Bot. Sada, XII (1892) 264 p. p—Dichotomae Bge. in Mém. Acad.
Pétersb. 7 sér. 9, 2 (1865) 8 p. p.—Calcitrapae Boiss. Fl. or. III (1875)
460 p. p.; Series Minutae Juz. in Tr. Tadzh. Bazy. Akad. Nauk SSSR,
8 (1940) 544.—Plants profusely dichotomously branched; capitula
solitary; involucral bracts stiff, carinate, somewhat attenuate above in
cusp; corolla yellowish; receptacular bristles smooth; achenes obovoid,
striate-ribbed, finely toothed above. Annual plants with soft leaves,
attenuate toward both ends and finely spinose-toothed.

Type of section: C. minuta Boiss.

232. C. minuta Boiss. Diagn. pl. or. 6 (1845) 117 and 10 (1849)
103 in adnot.; Bge. in Mém. Acad. Pétersb. 7, sér. 9, 2, 8; Boiss. Fl. or.
Ill, 489; Winkl. in Tr. Peterb. Bot. Sada, XII, 265; XIV, 231; Bornm.
in Beih. bot. Centralbl. XXXIV, 2, 194; Juz. in Tr. Tadzh. Bazy Akad.
Nauk SSSR, 8, 544; Kult. in Fl. Turkm. VII, 167.—C. aralensis Bge. in
Mém. Acad. Pétersb. Sav. étrang. 7 (1851) 178.—C. calcitrapaeformis
Jaub. and Spach, III, pl. or. 2 (1844-1846), tab. 158—Arctium minutum

319

O. Ktze. in Tr. Peterb. Bot. Sada, X (1887) 197.—Ic.: Jaub. and Spach
op. cit. tab. 158.

Annual. Stems profusely dichotomously branched night from base,
whitish, glabrous, lustrous, leafy; branches ascending, repeatedly branched
dichotomously, forming somewhat roundish bush, branchlets one-headed.
Leaves soft, sessile, ovate or oblong-ovate, attenuate toward both ends,
short acuminate-spinescent, finely spinose-toothed, arachnoid-hairy above,
gray-tomentose beneath. Capitula 6-7 mm wide (excluding cusps),
subglobose, finely arachnoid-hairy. Involucral bracts less numerous (about
20), stiff, carinate; outer bracts up to 7 mm long, horizontal, narrow-
lanceolate, acuminate; middle bracts lanceolate, 15-20 mm long and 3
mm wide, gradually acuminate into triquetrous stiff spine, almost
horizontal; inner bracts 9-10 mm long, membranous, white, narrow-
lanceolate, acuminate-spinescent. Corolla yellowish; receptacular bristles
smooth. Achenes obovoid, 2.5 mm long, 1.5 mm wide, ribbed-striate,
pitted, rugose, finely toothed above. Flowering I[V—V; fruiting V—VI.

Clayey deserts, foothill plains, in oases—Soviet Central Asia: Syr-
Darya, Pamiro-Alai, Kyzyl-Kum, Kara-Kum, mountainous Turkmenia,
Aralo-Caspian Region. General distribution: Iran, Kashmir. Described
from Tabriz. Type in Geneva.

Section 38. Jurineopsis (Juz.) Tschern. in Bot. Mat. Gerb. Inst.
Bot. Akad. Nauk UzbSSR, XVII (1962) 86—Nudicaules Winkl. in Tr.
Peterb. Bot. Sada, XII (1892) 191 p. p—WMicrocarpa subsect. jurineopsis
Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8 (1940) 518, 554.—Capitula
solitary terminal on long slender branches; involucral bracts long
acuminate or abruptly short acuminate; corolla pink, whitish, or yellow;
receptacular bristles scabrous. Achenes obovoid, glabrous, smooth.
Biennial plants with wingless stems and sessile cauline leaves.

Type of section: C. hastifolia Winkl.

223. C. krauseana Rgl. and Schm. in Izv. Obshch. Lyubit Estestv.
Antrop. Etnogr. 34, 2 (1882) 48; Juz. in Tr. Tadzh. Bazy Akad. Nauk
SSSR, 8, 554.—Arctium krauseanum O. Ktze. Rev. Gen. 1 (1891) 308.

Biennial. Stem erect, 50-80 cm high, glabrous, brown, sparsely
branched in upper half, usually very sparsely leafy above with remote
upper leaves; branches oblong; slender, one-headed. Leaves green, very
finely arachnoid-hairy above, gray-tomentose beneath; basal and lower
cauline leaves petiolate, unevenly pinnatisect in linear, entire lateral
segments; petiole narrow-winged; other cauline leaves abruptly reduced
toward stem apex, sessile, semiamplexicaul; in shape middle leaves like
basal; upper cauline leaves and those on branches lanceolate, entire,
with two or three pairs of finely spinose teeth only at base, strongly

326

320

reduced. Capitula ovoid-cylindrical, finely arachnoid-hairy, 6-7 mm wide
(excluding cusps) and 20 mm long (with corolla), numerous. Involucral
bracts except inner imbricate, with short recurved cusp; inner bracts
erect, somewhat broadened above, membranous, with thin short cusp,
scabrous along margin; receptacular bristles broadened above, weakly
scabrous. Corolla pale yellow or whitish. Achenes obovoid, glabrous,
smooth, about 3 mm long. Flowering V—VII.

Loamy slopes or outcrops of varicolored rocks, in higher foothills
or lower mountain zone.—Soviet Central Asia: Tien Shan (foothills of
the Fergana Valley). Endemic. Described from the vicinity of the Maili
River in “Kokania.” Type in Leningrad.

234. C. submutica Franch. in Ann. Sc. Nat. Bot. 6 sér. 16 (1883)
314;. Winkl. in Tr. Peterb. Bot. Sada, XII, 195; XIV, 192; Juz. in Tr.
Tadzh. Bazy Akad. Nauk SSSR, 8, 555.—C. krauseana Rgl. and
Schmalh. in Izv. Obshch. Lyubit. Estestv. Antrop. Etnogr., 34, 2 (1882)
48 quoad sp. ex Iskander.—C. /Jancifolia Winkl. in Tr. Peterb. Bot. Sada,
X (1889) 572; XII, 199; XIV, 219.—C. krauseana a. genuina f. elata
Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916) 141.—C. krauseana
a. genuina f. humilior Bornm. Ibid. (1916) 141.—C. krauseana B.
submutica Bornm. Ibid. (1916) 142.—C. kokanica f. nowobadensis
Bornm. Ibid. (1916) 144.—C. hastifolia auct. non Winkl.: Bornm. Ibid.
(1916) 143.—C. krauseana auct. non Rgl. and Schmalh.: Winkl. in Tr.
Peterb. Bot. Sada, XIV (1897) 218.—Arctium submuticum O. Ktze. Rev.
Gen. 1 (1891) 308.

Biennial. Stem erect, 20-60 cm high, brown, not glossy, subglabrous
or very finely arachnoid-hairy, branched throughout, more frequently
above, leafy; branches long, one-headed; basal rosettes lax, or entirely
absent. Leaves coriaceous, finely arachnoid-hairy above, white-tomentose
beneath; basal leaves petiolate, unevenly pinnatisect into lanceolate,
almost entire lobes or almost lyrate, with narrow, oblong-ovate terminal
lobe and lanceolate lateral lobes, usually narrowed at base; cauline leaves
usually like basal, gradually reduced toward stem apex but all sessile.
Capitula broadly ovoid, finely arachnoid-hairy, 10—12(15) mm wide
(excluding cusps). Involucral bracts numerous with short, mostly obtuse,
subobtuse or short acuminate cusp; inner bracts with broad-ovate or
almost roundish, dark brown, membranous apical appendage; receptacular
bristles broadened above, scabrous. Corolla pink. Achenes obovoid,
glabrous, smooth, 4-4.5 mm long. Flowering VII-—VIII; fruiting VIII—
IX.

Gravelly slopes in middle mountain zone.—Soviet Central Asia:
Pamiro-Alai (Zeravshan, Gissar, Peter the Great, and Alai ranges).
Endemic. Described from the vicinity of the village of Voru. Type in
Paris.

327

32

Note. Hybrids between C. submutica Franch. and C. alpestris Bornm.
are known, for which see the note to C. alpestris Bornm.

Hybrids between C. submutica Franch. and C. outichaschensis
Franch., described as C. iskanderi Bornm. (B.B.C. 34; 2 (1916) 168),
are also known from the Isander-Darya Valley.

This plant superficially is rather reminiscent of C. outichaschensis
Franch., but is distinguished by the presence of more numerous, shorter
and narrower outer and middle involucral bracts, and by the membranous
appendages of the inner bracts.

235. C. tamarae Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3
(1936) 297; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 555.—C.
krauseana var. microcephala Winkl. in Tr. Peterb. Bot. Sada, XIV (1897)
218.

Biennial. Stem erect, 25-40 cm high, brownish, finely arachnoid-
hairy, usually branched from base, leafy; branches long, one-headed.
Leaves green, finely arachnoid-hairy above, gray-tomentose beneath;
basal leaves petiolate, unequally pinnatisect in oblong segments usually
attenuate toward both ends; cauline leaves gradually reduced, sessile,
narrow-lanceolate, with two or three pairs of narrow lobes at base.
Capitula broadly ovoid, 15-17 mm wide (excluding cusps), finely
arachnoid-hairy. Involucral bracts numerous, closely appressed; outer
and middle bracts with divergent, 3-5 mm long cusp; inner bracts with
broadly lanceolate or almost roundish, membranous, usually pale pink
appendages, exceeding florets; receptacular bristles broadened above,
very weakly scabrous or almost smooth. Corolla pale pink or white.
Achenes obovoid, glabrous, smooth, about 4 mm long. Flowering VII-—
VIU; fruiting VIII.

Gravelly and clayey slopes in middle mountain zone in ravines and
along river banks.—Soviet Central Asia: Pamiro-Alai (Alai Range).
Endemic. Described from the vicinity of the Gulcha River, between the
mouth of the Shart River and the Kyzylbeles Pass. Type in Leningrad.

236. C. omphalodes Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 98.—C. krauseana y. longispinella Bornm.
in Beih. bot. Centralbl. XXXIV, 2 (1916) 142.—Exs.: H.F.A.M. No.
483.

Biennial. Stem erect, 80-90 cm high, very finely arachnoid-hairy,
glabrescent, dark brown, profusely branched from base, branches long,
spreading, slender, one-headed. Leaves coriaceous, green, glabrous above,
gray-tomentose beneath; basal leaves petiolate, unequally pinnatisect,
with narrow-lanceolate, finely toothed lateral segments; lower cauline
leaves like basal but sessile; middle and upper leaves oblong-lanceolate,
acuminate, usually spinose-toothed at base, sessile, semiamplexicaul;

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322

uppermost leaves strongly reduced. Capitula globose, finely arachnoid-
hairy, 11-12 mm wide (excluding cusps) and 22—25 mm long (with
florets), strongly truncate, almost foveate at base. Involucral bracts
numerous, all except inner with horizontal, subulate, 4-5 mm long cusp;
inner bracts erect, slightly broadened above, membranous, acuminate;
receptacular bristles broadened above, weakly scabrous. Corolla yellow.
Achenes obovoid, glabrous, smooth, 3 mm long. Flowering VII—VIII.
(Plate XXVI, Fig. 2).

Sandy and loamy slopes in foothills—Soviet Central Asia: Tien
Shan (foothills of the Fergana Range). Endemic. Described from Achi-
Sai, vicinity of Dzhelalabad. Type in Tashkent.

Note. At the border with the range of C. waldheimiana Bornm.,
this species develops forms resembling C. waldkeimiana Bornm.

237. C. dubia M. Pop. in Pop. and Andr. Rast. Zapovedn. Guralash
(1937) 37; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 556.—C.
jassyensis auct. non Winkl.: Bornm. in Beth. bot. Centralbl. XXXIV, 2
(1916) 142.

Biennial. Stem 40-60 cm high, erect, striate-sulcate, subglabrous,
branched from middle; branches long, spreading, one-headed. Leaves
glabrous above, finely grayish-tomentose beneath; basal leaves petiolate,
lyrate, with large, ovate, entire terminal lobe and oblong lateral lobes;
lower and middle cauline leaves like basal but middle leaves sessile,
broad-lanceolate, with one or two pairs of basal lobes; upper leaves
broad-linear, somewhat shorter than middle; uppermost leaves linear,
reduced. Capitula ovoid, 10-12 mm wide (excluding cusps), weakly
arachnoid-hairy. Involucral bracts stiff, squarrose, often slightly decurved,
with stiff wide cusp; outer bracts carinate, up to 2 mm wide at base and
10-12 mm long; middle bracts coriaceous, somewhat flat, up to 15 mm
long; inner bracts with membranous, oblong, acuminate, 2-3 mm wide
appendages, not forming corona; receptacular bristles flattened above;
weakly scabrous. Corolla pink. Achenes obovoid, glabrous, smooth, light
brown. Flowering VI-—VIII.

Gravelly slopes in upper mountain belt.—Soviet Central Asia:
Pamiro-Alai (Turkestan, Gissar and Zeravshan ranges). Endemic.
Described from the Guralash Ravine. Type in Tashkent.

Note. Plants somewhat different from true C. dubia are found within
the Gissar Range; they have smaller capitula, shorter involucral bracts,
and broader middle and upper cauline leaves.

238. C. kokanica Rgl. and Schmalh. in Izv. Obshch. Lyubit. Estestv.
Antrop. Etnogr. 34, 2 (1882) 49; Winkl. in Tr. Peterb. Bot. Sada, XI],
198; XIV, 219; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 555.—C.

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323

kokanica var. virgata Winkl. op. cit. (1897) 219; Bornm. in Beih. bot.
Centralbl. XXXIV, 144.—C. kokanica a. genuina Bornm. Ibid. 143.—
Arctium kokanicum O. Ktze. Rev. Gen. 1 (1891) 308.

Biennial. Stem erect, arachnoid hairy-tomentose, usually corymbosely
branched above or from base, sparsely leafy, with dense rosette of leaves
20-35 cm high; branches slender, long, one-headed. Leaves finely
arachnoid-hairy above, white-tomentose beneath, lanceolate, pinnatisect
or almost bipinnatisect, petiole narrow-winged or almost wingless, lateral
segments approximate, narrow-linear, almost entire, weak, usually weakly
crispate in basal leaves; basal leaves petiolate, usually 7—8(13) cm long,
with 8-12 pairs of segments; cauline leaves reduced, 2 to 5 mm long,
sessile, like basal leaves but with two to five (seven) segments; leaves
on branches strongly reduced. Capitula globose or somewhat flat-globose,
weakly or distinctly foveate at base, 10-12 mm wide (excluding cusps),
finely arachnoid-hairy. Involucral bracts numerous, linear-subulate, all
bracts closely appressed, only short bracts about 2—3 mm long, cusps
arcuately recurved; inner bracts scarious, flat, short acuminate, not
forming “corona,” about 15 mm long; receptacular bristles more or less
broadened above, scaberulous. Corolla pale pink. Achenes narrow
obovoid, glabrous, smooth, about 4 mm long. Flowering VII-—VIII;
fruiting VIII.

Stony-gravelly dry slopes from middle to upper mountain zone——
Soviet Central Asia: Pamiro-Alai (Alai Range, Shugnan, Pamir).
Described from Alai. Type in Tashkent.

239. C. hastifolia Winkl. in Tr. Peterb. Bot. Sada, X (1889) 571;
XII, 195; XIV, 208; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8,
555.—C. taldykensis Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3
(1936) 299; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 555.

Biennial? Stem erect, 25—80 cm high, finely arachnoid-hairy,
glabrescent; brownish, leafy, profusely branched from base; branches
spreading in turn, branched in one-headed branchlets, entire plant with
many capitula. Leaves coriaceous, green, glabrous above with network
of prominent veins, white-tomentose beneath; basal leaves petiolate or
pinnatisect in lanceolate, sparsely toothed, sessile lateral segments on
winged rachis or almost lyrate, with large, oblong-ovate, acuminate,
hastate, excurrent terminal lobe and rather small, narrow-lanceolate,
usually slightly decurved, many (up to 10 pairs) lateral lobes, usually
withering early; lower cauline leaves like basal but somewhat smaller;
other leaves sessile, lanceolate or subovate, with roundish auricles, sparsely
finely spinose-toothed, gradually reduced toward stem apex. Capitula
ovoid, very finely arachnoid-hairy, 8-12 mm wide (excluding cusps) and
20-25 mm long (with florets). Involucral bracts numerous, all except

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324

inner lanceolate, with somewhat short, 3—4 mm long, erect, subulate
cusp; inner bracts slightly broadened above, with lanceolate, light brown,
usually erect, membranous, somewhat acuminate appendages; receptacular
bristles broadened above, scabrous. Corolla purple or pink, exserted.
Achenes obovoid, glabrous, smooth, about 4 mm long. Flowering VII-—
VIII. (Plate XX VI, Fig. 1.)

Stony slopes and loamy screes in middle mountain zone.—Soviet
Central Asia: Pamiro-Alai (Alai, Gissar, Zeravshan, Peter the Great,
and Vakhsha ranges, Roshan). Endemic. Described from Karategin,
Pumbach. Type in Leningrad. ;

Note. The plants from the Peter the Great Range differ somewhat
from typical C. hastifolia Winkl. by broader leaves (the lateral lobes of
the basal leaves are broadly ovate or even round.) Further study of the
material from this region is necessary.

The species was described from three localities. I propose to
designate as lectotype the plant with the label: “Pumn-Baczi. 1878, 21,
VIII. Newessky,” preserved in Leningrad.

240. C. transiliensis Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser.
1, 3 (1936) 298.

Biennial. Stem erect, 45-55 cm high, brownish, finely arachnoid-
hairy, striate-sulcate, branched from base, more frequently in upper half;
branches long, one-headed. Leaves thin-coriaceous, green, finely
arachnoid-hairy above, finely tomentose beneath, sparsely finely spinose-
toothed; basal leaves petiolate, almost lyrate, withering early; lower
cauline leaves like basal but short-petiolate; other cauline leaves ovate
or broadly ovate, sessile, gradually reduced toward stem apex. Capitula
broadly ovoid, 17 mm wide (excluding cusps) and 30 mm long (with
florets), finely arachnoid-hairy, weakly truncate at base or somewhat
attenuate. Involucral bracts numerous, all except inner closely appressed,
terminating in thin, subulate-triquetrous, 5-6 mm long erect cusp; inner
bracts membranous, erect, broadened above, lanceolate, acuminate in
thin cusp, scaberulous along margin; receptacular bristles subglabrous,
sometimes broadened above and weakly scabrous. Corolla pink. Achenes
(immature) obovoid, glabrous, smooth, about 4 mm long. Flowering .
VI-VII.

In foothills——Soviet Central Asia: Tien Shan (Trans-Ili Alatau).
Endemic. Described from the Bakhtiar River. Type in Leningrad.

241. C. waldheimiana Bornm. in Beih. bot. Centralbl. XXXIV, 2
(1916) 140.

Biennial. Stem erect, 30-80 cm high, leafy, subglabrous, finely
arachnoid-hairy only below, somewhat brownish, corymbosely and

325

Plate XXVI.

1—Cousinia hastifolia Wink|.; 2—C. omphalodes Tschern.

331

332

326

sparsely branched from middle branches long, one-headed. Leaves
coriaceous, glabrous and green above, grayish-tomentose beneath; basal
leaves petiolate, pinnatisect into narrow lanceolate, fast withering
revolute, lateral segments, terminal segment lanceolate, acuminate,
excurrent on narrow-winged rachis; cauline leaves gradually reduced
toward stem apex, sessile, semiamplexicaul, narrow-lanceolate or
lanceolate, attenuate toward apex, amost entire, usually with two to
four pairs of teeth only at base; leaves on branches strongly reduced.
Capitula ovoid-conical, weakly arachnoid-hairy, 10-12-15 mm wide
(excluding cusps) and 25-30 mm long (with florets), somewhat truncate.
Involucral bracts numerous, all except inner with more or less arcuate
cusp; inner bracts erect, lanceolate, membranous above, acuminate in
very thin and short spine, usually brownish; receptacular bristles
broadened above, scabrous. Corolla pink or purple. Achenes obovoid,
glabrous, smooth, 3.5—-4 mm long. Flowering VI-—VIII.

Gravelly-loamy dry slopes in middle mountain zone.—Soviet Central
Asia: Tien Shan (Chatkal, Uzun-Akhmat and Fergana ranges). Endemic.
Described from Uzun-Akhmat. Type in Leningrad.

Note. C. waldheimiana Bornm. and C. margaritae Kult. are very
close species, differing mainly in the color of the corolla (whitish or
pale yellow in C. margaritae). However, the color of the corolla is
poorly preserved in the herbarium, which makes identification of plants
difficult. This is why the geographic delimitation of these species is
unclear at the present time. In collecting material for these species,
special attention must be paid to the color of the corolla in nature.

242. C. margaritae Kult. in Tr. Sredneaz. Gos. Univ. Ser. 1, VIllc,
ViIN@io29)e5:

Biennial. Stem erect, 20-40 .cm high, finely arachnoid-hairy, leafy,
sparsely branched, branches one-headed. Leaves green, glabrous above,
finely white-tomentose beneath; basal leaves petiolate, pinnatisect into
narrow-lanceolate, early-withering lateral segments; lower cauline leaves
like basal; other leaves gradually reduced toward stem apex, sessile,
pinnatisect or pinnatilobate, lateral segments narrow-lanceolate, remote
on narrow-winged rachis; uppermost leaves lanceolate, spinose-toothed
at base, acuminate-spinescent. Capitula ovoid-globose, arachnoid-hairy,
truncate at base 11-18 cm wide (excluding cusps), 25-30 mm long
(with exserted corollas). Involucral bracts numerous, all except inner
with subulate, arcuately decurved or squarrose cusp; inner bracts scarious
above and slightly broadened in somewhat short, almost ovate, brownish
spinescent appendages; receptacular bristles broadened above, weakly
scabrous. Corolla whitish or pale yellowish. Achenes obovoid, glabrous,
smooth, 4 mm long. Flowering VI.

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S27

Limestone screes or loamy slopes from foothills to middle mountain
zone.—Soviet Central Asia: Tien Shan (Talass Alatau). Endemic.
Described from the mouth of the Beshtash River. Type in Tashkent.

Note. Cf. note to the previous species.

243. C. jassyensis Winkl. in Tr. Peterb. Bot. Sada, X (1889) 472.—
C. kokanica 6. perapressa Bornm. in Beth. bot. Centralbl. XXXIV, 2
(1916) 144.

Biennial. Stem erect, 30-80 cm high, usually branched from base,
leafy, finely arachnoid-hairy, brownish; branches long, one-headed.
Leaves coriaceous, green above, gray-tomentose beneath; basal leaves
petiolate, pinnatipartite into narrow-lanceolate, acuminate terminal lobe,
and almost linear, almost entire, acuminate, short-spinescent lateral lobes;
lower cauline leaves usually like basal but sessile; other cauline leaves
lanceolate, sessile, acuminate, entire, with two or three pairs of small
teeth at base, auriculate, gradually reduced toward stem apex. Capitula
narrow-lanceolate, 9-10 mm wide and 22 mm long (with florets), very
finely arachnoid-hairy. Involucral bracts numerous, very closely
appressed, short cuspidate or entirely without cusp; inner bracts with
short, narrow-lanceolate, almost membranous, short acuminate
appendages; receptacular bristles weakly broadened above, slightly
scabrous. Corolla whitish or yellowish; anthers purple. Achenes broadly
obovoid, glabrous, smooth, light gray, 3.5 mm long. Flowering VII-
VIII.

Gentle slopes in foothills —Soviet Central Asia: Tien Shan (Fergana
Range). Endemic. Described from the Yassy River Valley near Uzgent.
Type in Leningrad.

Note. Hybrids of this species with C. ninae Juz. are known.

Section 39. Tenellae Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2 (1865)
8.—Psilacanthae Boiss. Fl. or. II] (1875) 461 p. p.—Jnermes Winkl. in
Tr. Peterb. Bot. Sada, XII (1892) 196 p. p—Series Tenellae Juz. in Tr.
Tadzh. Bazy Akad. Nauk SSSR, 8 (1940) 544.—Capitula solitary, with
many florets; involucral bracts narrowed into subulate, weakly divergent
cusp; corolla pink; receptacular bristles scabrous; achenes obovoid, with
scarcely noticeable longitudinal ribs, denticulate above.

Annual plants with soft, almost entire leaves.

Type of section: C. tenella Fisch. and Mey.

244. C. tenella Fisch. and Mey. Ind. sem. hort. Petrop. I (1834) 25;
Ldb. FI. Ross. II, 677; Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2 (1865)
8; Boiss. Fl. or. IJ, 494; Winkl. in Tr. Peterb. Bot. Sada, XII, 197;
XIV, 219; Bornm. in Beih. bot. Centralbl. XXXIV, 2, 145; Grossh. FI.

328

Kavk. IV, 165; Takht. in Tr. Arm. Fil. Akad. Nauk SSSR, Biol. Ser. 2,
177; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 544; Takht. in FI.
Erev. 317; Kult. in Fl. Turkm. VII, 163; Fl. Azerb. VIII, 362.—Lappa
tenella Fisch. and Mey. ex DC. Prodr. VI (1837) 662.—Arctium tenellum
O. Ktze. in Tr. Peterb. Bot. Sada, X (1887) 197.

Annual. Stem erect, 8-40 cm high, whitish, smooth, lustrous, leafy,
dichotomously branched from very base; branches leafy, long, in turn
dichotomously branched, branchlets one-headed. Leaves soft, sessile, very
finely arachnoid-hairy above, almost smooth, gray-tomentose beneath,
oblong-ovate or lanceolate, attenuate toward both ends, entire or with
small teeth. Capitula 4 mm wide (excluding cusps), almost globose,
glabrous. Involucral bracts numerous, all similar, narrow-linear, squarrose,
finely cuspidate, weakly uncinate, 5-7 mm long. Corolla pink;
receptacular bristles scabrous. Achenes obovoid, 2 mm long and 1 mm
wide, with scarcely noticeable longitudinal ribs, gray, punctate, matty,
with small teeth above. Flowering IV—V; fruiting V—VI.

Foothills and high deserts, as weed.—Soviet Central Asia: Pamiro-
Alai, Syr-Darya, Tien Shan, Balkhash Region, Aralo-Caspian Region,
Kara-Kum, mountainous Turkmenia (Kopetdag, Badhkyz); Caucasus:
Southern and eastern Transcaucasia. General distribution: Iran,
Afghanistan, Dzhungaria-Kashgaria, Armenia-Kurdistan. Described from
the Nakhichevan District. Type in Leningrad.

Section 40. Glaphyrocephalae Tschern. in Bot. Mat. Gerb. Inst.
Bot. Akad. Nauk UzbSSR, XVII (1962) 87.—Capitula solitary, on short
slender peduncles; involucral bracts carinate, gradually acuminate into
thin, falcate cusp; receptacular bristles smooth; corolla pink; achenes
obovoid, smooth, roundish above. Perennial plants with greatly incised
leaves with linear, short, revolute lateral segments, on narrow-winged
rachis.

Type of section: C. glaphrocephala Juz.

245. C. glaphrocephala Juz. ex Tschern. in Bot. Mat. Gerb. Inst.
Bot. Akad. Nauk UzbSSR, XVII (1962) 94.

Perennial. Stems several, erect, short, 15-20 cm high, finely
tomentose, branched from middle; branches slender, short, one-headed.
Leaves arachnoid-hairy above, white-tomentose beneath; basal leaves
short-petiolate, pinnatisect, lateral segments linear, short, revolute,
acuminate, rachis narrow-winged, entire, revolute; cauline leaves
gradually reduced, like basal sessile, very short-decurrent; leaves on
branches lanceolate, with one or two pairs of teeth at base. Capitula
ovoid, 20 mm long and 7 mm wide (excluding cusps), arachnoid-hairy.
Involucral bracts about 40, carinate; outer bracts narrow-lanceolate,

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329

acuminate into thin spine, falcately decurved, about 5 mm long; middle
bracts narrow-lanceolate, acuminate-spinescent, falcate, 10-12 mm long;
subsequent bracts lanceolate, short acuminate, erect, about 9 mm long;
inner bracts almost scarious, lanceolate, slightly broadened toward apex
and acuminate into thin spine, dark purple in middle, about 10 mm
long; receptacular bristles smooth. Corolla exserted, pink. Achenes
obovoid, smooth, roundish above. Flowering VI-—VII; fruiting VIJ—VIII.

Middle mountain zone.—Soviet Central Asia: Pamiro-Alai. Endemic.
Described from Suzystau (Novo-Bulak). Type in Tashkent.

Section 41. Alpinae Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2 (1865)
45 p. p.—Odontocarpae Winkl. in Tr. Peterb. Bot. Sada, XII (1892)
253 p. p.—Acanthotoma Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8
(1940) 522, 563.—Alpinae auct. non Bge.: Boiss. in Fl. or. HI (1875)
461 p. p——Capitula solitary terminal on stem and branches or aggregated
in inflorescences; involucral bracts acuminate-cuspidate; receptacular
bristles scabrous, occasionally smooth or weakly scabrous; corolla yellow,
yellowish, whitish or less often pink; achenes obovoid, glabrous, smooth,
with narrow-winged narrow ribs or almost triquetrous, with broad winged
ribs. Perennial plants with many-headed root and pinnatisect basal leaves;
leaf segments palmatipartite in linear lanceolate or almost linear entire
lobes, seemingly forming whorl over almost entire length of uniformly
winged leaf rachis.

Lectotype of section: C. verticillaris Bge.

246. C. magnifica Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1,
3 (1936) 319; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 563.

Perennial. Root thick, woody, with numerous remains of previous
year’s leaves at base. Stem 80—135 cm high, erect, glabrous in lower
part, finely appressed-tomentose above, densely white-tomentose in
inflorescence, sparsely branched from middle. Leaves with usual structure
for the section, with narrow-winged, glabrous, weakly tomentose rachis
in cauline leaves and long, linear, acuminate-spinescent, undivided,
revolute lateral segments, bright green, scabrous above due to small
spines, finely white-tomentose beneath; basal leaves petiolate; petioles
glabrous, broadened at base, margin spinose; cauline leaves gradually
reduced, sessile, semiamplexicaul, short-decurrent, with wider-winged
rachis and wider lateral lobes; inflorescence almost racemose, with
spreading many-headed branches. Capitula numerous, on short peduncles,
broadly ovoid to almost turbinate, 25-30 mm long and 25—30 mm wide
(at base), finely arachnoid-hairy. Involucral bracts numerous (100-120),
somewhat flat, compactly imbricate; all bracts erect, dorsally scabrous
due to dense spines, with weakly divergent cusp; outer bracts lanceolate,

330

up to 10 mm long, acuminate; middle bracts like outer, but up to 20
mm long; inner bracts thin-coriaceous, broadly linear, acuminate-
spinescent, 25-30 mm long; receptacular bristles strongly scabrous.
Corolla yellow, about 30 mm long (including anther tube). Achenes
numerous, somewhat flat, obovoid, 12-14 mm long, 5-7 mm wide, two
or three ribbed, with somewhat broad, fragile, two or three wings, and
two or three obtuse teeth at apex. Flowering VII; fruiting VIII. (Plate
XXVII, Fig. 1.)

Stony mountain zone.—Soviet Central Asia: Pamiro-Alai (south).
Endemic. Described from the area between the Shurak and Kyzylsu
rivers, on the western slopes of the Korimonak Mountain. Type in
Leningrad.

247. C. lanata Winkl. in Tr. Peterb. Bot. Sada, XIII (1894) 238;
Winkl. ibid., XIV, 204; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8,
563.

Perennial. Root thick, woody, stems solitary, occasionally two or
three, with numerous remains of previous year’s leaves at base, erect,
100-135 cm high, thick, about 15 mm wide below, striate-sulcate,
tomentose-arachnoid hairy, sericeous, less often arachnoid-hairy,
occasionally glabrous, densely leafy, branched from base; branches short,
sometimes reduced to peduncles; inflorescence long, spicate or subspicate.
Leaves typical of the section, with narrow-winged, rather wide but
strongly broadened toward base, densely woolly rachis due to long silky
hairs and linear-subulate entire revolute lateral segments, reduced toward
rachis base to thin spines, green, glabrous and smooth above or sericeous
along midrib, white-tomentose beneath with long silky hairs; basal leaves
short-petiolate; cauline leaves gradually reduced toward stem apex,
sessile, short-decurrent. Capitula numerous, on short peduncles, young
capitula almost sessile, broadly cylindrical, deflorate capitula almost
obconical, up to 30 mm long and 20-25 mm wide, densely woolly.
Involucral bracts numerous (about 100), somewhat flat, compactly
imbricate, erect; outer bracts up to 10 mm long, lanceolate, short
acuminate-spinescent, scabrous dorsally due to numerous spinules; middle
bracts like outer but up to 20 mm long, 5 mm wide at base, attenuate
from middle; inner bracts thin-coriaceous, linear, acuminate-spinescent,
25-30 mm long, 2 mm wide at base; receptacular bristles smooth or
some weakly scabrous. Corolla yellow, about 15 mm long. Achenes
numerous, obovoid, winged, 10-12 mm long and 6 mm wide (including
wings), somewhat flat or almost triquetrous, wings 1.5 mm wide, weakly
bulged at apex, forming pit lodging pappus bristles. Flowering VII—
VII; fruiting VII-IX.

Outcrops of varicolored rocks, loamy slopes from the middle to the

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331

lower boundary of the upper mountain zone.—Soviet Central Asia:
Pamiro-Alai. Endemic. Described from Musabazar. Type in Leningrad.

248. C. pseudolanata M. Pop. ex Tschern. in Bot. Mat. Gerb. Inst.
Bot. Akad. Nauk UzbSSR, XVII (1962) 104.

Perennial. Root woody, thick. Stem erect, with numerous remains of
previous year’s leaves at base, usually tall, thick, striate-sulcate;
tomentose, leafy; inflorescence oblong, narrow spicate. Rosette leaves
glabrous beneath, glandular above or densely pubescent on both sides
pinnatisect in numerous, almost acicular lateral segments, sessile on
narrow-winged rachis, in whorl, rachis basally broadened, lateral segments.
reduced to thin spines. Capitula (previous year’s) on short peduncles,
almost turbinate, up to 20 mm wide (excluding cusps), numerous,
pubescent. Involucral bracts numerous, somewhat flat, smooth, erect; outer
bracts narrow-lanceolate, long acuminate, 22 mm long, up to 3 mm wide
at base; middle bracts like outer but longer, 25 mm long; inner bracts
thin-coriaceous, linear, acuminate-spinescent; receptacular bristles weakly
scabrous. Achenes obovoid, almost triquetrous 5 mm long and 3 mm
across, narrow-winged, apically terminating in three teeth.

Lower part of upper mountain zone.—Soviet Central Asia: Pamiro-
Alai (town of Nuratau). Endemic. Described from Aktau. Type in
Tashkent.

Note. We have the tip of an inflorescence with the previous year’s
capitula and numerous sterile rosettes of leaves. However, even from
such inadequate material it is apparent that this is an entirely unique
species of the section. Additional collections of this species are necessary
to determine more precisely its position in the section.

249. C. adenophora Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser.
1g8@i936)2320)

Perennial. Roots thick, woody, many-headed, bearing sterile rosettes
of leaves and stems, caudices densely covered with remains of previous
year’s leaves, usually gray-tomentose. Stems erect, 40-50 cm high, almost
smooth, glabrous or sparsely glandular, spreadingly branched, rather
densely leafy. Leaves green, scabrous above due to glandular hairs,
grayish-tomentose beneath, pinnatisect in numerous, linear-acicular,
revolute segments, on almost wingless leaf-rachis, whorled, rachis
broadened at base and lateral segments reduced to thin spines; cauline
leaves gradually reduced toward stem apex, sessile, short-decurrent.
Capitula on glandular-hairy peduncles, narrow ovoid, 10—15 wide
(excluding cusps), without arachnoid-hairy indumentum. Involucral bracts
about 30-40, somewhat flat, coriaceous, scabrous due to dense cover of
glandular hairs; outer bracts 8-15 mm long, divergent, lanceolate, with
prominent midrib, terminating into stiff spine; middle bracts erect,

338

332

20-30 mm long, with prominent midrib, terminating into stiff cusp;
inner bracts membranous, lanceolate, acuminate-spinescent. Corolla
yellow; receptacular bristles scabrous. Achenes obovoid, almost three-
angled, narrow winged, 2-3 dentate above, 6-7 mm long. Flowering
VII-VIN; fruiting VITI-IX.

Stony slopes in upper mountain zone.—Soviet Central Asia: Pamiro-
Alai (Samarkand Mountains). Endemic. Described from Manys Pass.
Type in Leningrad.

Note. The species was described with an indication of its having
originated from Manas Mountain in the Talass Alatau. This report is
erroneous because the plant was collected, as is clear now, in Manys
Pass in the Samarkand Mountains.

250. C. splendida Winkl. in Tr. Peterb. Bot. Sada, XIII (1894) 240
(incl. var. komarovii); XIV, 234; Bornm. in Beih. bot. Centralbl. XXXIV,
2, 198; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 563.—C. lipskyi
Winkl. op. cit. XIV (1897) 237 excl. var. approximata.

Perennial. Root woody, many-headed; caudices with numerous
remains of previous year’s leaves. Stem erect, 50-70 cm high, finely
striate-sulcate, glabrous, leafy, sparsely branched; branches and stem
one-headed. Leaves coriaceous, glabrous and bright green above, white-
tomentose beneath; basal leaves lanceolate, pinnatisect, 4-7 mm wide,
somewhat flat, lateral segments on broad-winged 3-6 mm wide rachis
with flat wings as if in whorl, petiolate, petioles thickened at base,
somewhat flat, ciliate; lower cauline leaves like basal, but with short
petioles, semiamplexicaul; other cauline leaves like basal but gradually
reduced toward stem apex, sessile, ‘semiamplexicaul, short-decurrent.
Capitula broadly cylindrical, 18-25 mm wide (excluding cusps), glabrous.
Involucral bracts numerous, up to 100, all except inner very similar;
outer bracts light green, squarrose, with prominent vein on dorsal surface,
broadly lanceolate, acuminate-spinescent, ciliate, cilia 12-15 mm long;
middle bracts similar but wider and longer, up to 18 mm long and 6
mm wide, more densely ciliate; inner bracts somewhat flat, 20-23 mm
long, with membranous appendage above, unevenly divided along margin,
acuminate-spinescent. Corolla yellow; receptacular bristles scabrous.
Achenes obovoid, 7 mm long, ribbed, narrow-winged along ribs.
Flowering VIII-IX; fruiting IX. (Plate XXVIII, Fig. 1.)

Gentle and loamy slopes in upper mountain zone.—Soviet Central
Asia: Pamiro-Alai (Zeravshan and Gissar ranges). Endemic. Described
from the vicinity of Lake Iskanderkul. Type in Leningrad.

Note. Three plants were cited in the description of this species; |
propose to consider as the lectotype the plant with the label: “Iskander
kul. alt. 7000 pd. 1893 10 VII legit. V.L. Komarov.”

339

Bie)

251. C. xanthina Bornm. in Beih. bot. Centralbl. XXXIV, 2 (1916)
190; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 563.—C. curviloba
Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3 (1936) 321; Juz. in
Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 563——C. acanthophylla Kult. ex
Zak. in Tr. Uzb. Gos. Univ. Nov. Ser. 89 (1958) 158, nomen et in herb.

Perennial. Root thick, woody, many-headed, caudices covered with
numerous remains of previous year’s leaves. Stem several, erect, 25-40
cm high, shallow striate-sulcate, finely arachnoid-hairy aboye, almost
always one-headed, very sparsely branched in upper one-third part, leafy.
Leaves coriaceous, green above, scabrous due to glandular hairs, green
beneath or white-tomentose only along margin; basal leaves short-
petiolate, petioles sheath-like broadened, lanceolate, pinnatisect in
lanceolate, somewhat flat, rather broadened, 3-6 mm wide, gradually
acuminate-short spinescent, sessile lateral segments, in whorls on
somewhat flat, broad winged, 2-3 mm wide rachis, less often lateral
segments and leaf rachis, somewhat narrower; cauline leaves gradually
reduced toward stem apex, like basal leaves, but sessile, short-decurrent;
uppermost leaves reduced. Capitula broadly lanceolate, 15-23 mm wide
(excluding cusps), finely white arachnoid-hairy. Involucral bracts about
50, somewhat flat, sparsely glandular; outer bracts horizontal, ‘slightly
decurved; lanceolate, 3-4 mm wide at base, 17 mm long, gradually
acuminate-short spinescent; middle bracts erect-spreading, lanceolate,
short acuminate, up to 20 mm long and up to 5 mm wide at base; inner
bracts almost membranous, lanceolate, dorsally scabrous due to spinules,
broadened above in appendages up to 4 mm wide, usually short and
unevenly divided along margin. Corolla pale yellow; receptacular bristles
scabrous. Achenes oblong-obovoid, 6-7 mm long, with scarcely
noticeable ribs, with small teeth above. Flowering VIII-IX; fruiting IX.

Loamy slopes in upper mountain zone.—Soviet Central Asia: Pamiro-
Alai (Alai Range, northern slopes of Turkestan Range; Zeravshan Range;
in the upper reaches of the Pasrut River). Endemic. Described from the
vicinity of Kalai-Makhmud (the Sokh River system), climbing from the
Kul area to Kyzyl-Uchuk. Type in Leningrad.

Note. True C. xanthina Bornm. is known from the region from
where C. curviloba Juz. was described. Considering that C. curviloba
differs from C. xanthina only by narrower, somewhat recurved lateral
leaf segments, it is difficult to recognize it as a separate species.

The hybrid of C. xanthina Bornm. x C. laetevirens Winkl. is known
from the upper course of the Pasrut River (a tributary of the Fandarya
River).

252. C. trachyphylla Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8
(1940) 524, 563.

340

334

Perennial. Root thick, woody, many-headed; caudices with numerous
remains of previous year’s leaves. Stem erect, about 50 cm high, striate-
sulcate, sparsely glandular, glandular-arachnoid-hairy above, leafy, usually
branched in upper one-third part. Leaves coriaceous, green and scabrous
above due to glandular hairs, white-tomentose beneath; basal leaves
petiolate, with thickened wide petioles, lanceolate, pinnatisect in
lanceolate, somewhat flat, 5-6 mm wide, gradually acuminate-short
spinescent, lateral segments in whorls on somewhat flat, broad-winged,
2—4 mm wide rachis; cauline leaves gradually reduced toward stem apex,
like basal, but sessile, semiamplexicaul, short-decurrent; uppermost leaves
almost pinnatilobate, lobes lanceolate, with wider wings of rachis and
wider lateral segments. Capitula broadly ovoid, about 17 mm wide
(excluding cusps), rusty-arachnoid-hairy. Involucral bracts about 50,
somewhat flat, glandular; outer bracts slightly decurved, 15—17 mm long,
oblong-lanceolate, acuminate-spinescent, with prominent midrib on dorsal
side; middle bracts oblong-lanceolate, erect, about 25-30 mm long, with
prominent midrib on dorsal side, acuminate-spinescent, usually exceeding
inner bracts; inner bracts membranous above, broadly lanceolate, short
acuminate-spinescent, scabrous along dorsal surface and margin, about
18-20 mm long. Corolla whitish; receptacular bristles scabrous. Flowering
VII-IX; fruiting IX.

Loamy slopes in upper mountain zone—Soviet Central Asia: Pamiro-
Alai (town of Kugifrush). Endemic. Described from Kugifrush,
Alchakhon area. Type in Leningrad.

Note. The species is quite close to C. xanthina Bornm., from which
it differs by having a whitish corolla, rusty-pubescent capitula, and leaves
that are whitish-tomentose on the lower surface.

253. C. chrysantha Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIIIc,
VI (1929) 25 incl. var. aspera.—C. xanthina 8. subvestita Bornm. in
Beih. bot. Centralbl. XXXIV, 2 (1916) 191. -

Perennial. Root thick, woody, many-headed; caudices with remains
of previous year’s leaves. Stems several, erect, 20-50 cm high, glabrous
or sometimes, especially in upper part, sparsely glandular, simple or
branched in upper one-third part; branches one-headed, leafy. Leaves
coriaceous, green and glabrous above, or sometimes with sparse and
fine sessile glandular hairs, white-tomentose beneath; basal leaves
lanceolate, pinnatisect in 4-7 mm wide, somewhat flat lateral segments,
on wide winged 3-5 mm wide rachis with somewhat flat wings, as if in
whorl, petiolate, petioles somewhat flat, broadened at base; lower cauline
leaves like basal but with short, semiamplexicaul, usually ciliate petioles;
other cauline leaves like basal but gradually reduced, sessile, short-
decurrent. Capitula broadly cylindrical, sparsely arachnoid-hairy, 15—25

335

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34] Plate XXVII.

1—Cousinia magnifica Juz.; 2—C. chlorantha Kult.

342

336

mm wide (excluding cusps). Involucral bracts about 90; outer bracts
slightly decurved, 7-19 mm long, lanceolate, short-acuminate; middle
bracts erect, 12-23 mm long, somewhat flat, broadly lanceolate, with
somewhat prominent midrib on dorsal side, terminating in cusp; inner
bracts with membranous, almost roundish, short and unevenly divided
appendage, usually recurved at anthesis. Corolla yellow; receptacular
bristles scabrous. Achenes obovoid, about 6 mm long, indistinctly ribbed,
very narrow-winged along ribs. Flowering VIII-IX; fruiting IX.

Gravelly slopes in upper mountain zone.—Soviet Central Asia: Tien
Shan (western). Endemic. Described from the vicinity of Karatyube.
Type in Tashkent.

Note. Two plants were mentioned in the description of this
species; I consider as the lectotype the first of these with the label:
“In distr. Czimkent, in montibus prope Kara-Tjube. 22 IX 1921.
Kultiassov.”

254. C. stephanophora Winkl. in Tr. Peterb. Bot. Sada, X (1887)
575; XII, 260; XIV, 230; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8,
563.—C. lipskyi var. approximata Winkl. op. cit. (1897) 237.—C.
xanthina y. approximata Bornm. in Beih. bot. Centralbl. XXXIV, 2
(1916) 192. C. xanthina 8. subvestita auct. nom Bornm.: Bornm. in
Beih. bot. Centralbl. XXXIV, 2 (1916) 290.

Perennial. Roots thick, woody, many-headed; caudices with numerous
remains of previous year’s leaves. Stem numerous, 20-40 cm high, erect,
simple or sparsely branched above, usually appressed gray arachnoid-
hairy throughout, glabrescent, less often glabrous below, appressed-hairy
above, leafy; branches and stems one-headed. Leaves not glandular above,
usually finely appressed-hairy or almost glabrous, white-tomentose
beneath; baal leaves petiolate, usually with weakly broadened, somewhat
flat petioles, pinnatisect, with 3-6 mm wide segments, in turn,
palmatipartite in linear-lanceolate entire lobes, as if forming whorl almost
throughout on uniformly winged rachis with flat wings; lower cauline
leaves like basal but sessile, short-decurrent; other cauline leaves
gradually reduced toward apex, sessile, short-decurrent, with wider leaf
segments and rachis wings. Capitula cylindrical, (10)15—18(20) mm wide
(excluding cusps), very finely arachnoid-hairy. Involucral bracts about
50-60; outer bracts lanceolate, recurved, 10-16 mm long, carinate, short-
acuminate; middle bracts lanceolate, almost horizontal, 15-20 mm long,
acuminate; subsequent bracts somewhat flat, broadly lanceolate, short-
acuminate, usually dark-colored dorsally, 12-20 mm long, erect; inner
bracts membranous, broadened above, scabrous along margin, 15-18
mm long, terminating in short spine. Corolla white or pale yellow; anther
tube pink; receptacular bristles scabrous. Achenes obovoid, 5-7 mm

343

330,

long, longitudinally ribbed, narrow-winged along ribs, with four teeth
above. Flowering VII—VIII; fruiting VIII-IX.

Stony loamy slopes, in upper mountain zone.—Soviet Central Asia:
Pamiro-Alai. Endemic. Described from the region between Tavildar and
Sagyrdasht. Type in Leningrad.

Note. Three plants from different places were mentioned in the
description of this species; I consider as the lectotype the plants with
the following label: “Darwas: nérdl. Pass. zw Tevildara et Sagridascht
an einem sudl. Zufluss des Wakisch. 8-9000’ 30 VHI-9 IX 1881. Regel.”

255. C. semilacera Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1,
3 (1936) 322; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 564.—C.
lachnopoda Juz. op. cit. (1940) 525, 564.—C. jacea Rech. f. and
Edelberg in Dan. Biol. Skr. 8, 2 (1955) 104, fig. 74, 78F.—Ic.: Rech. f.
and Edelberg. op. cit.

Perennial. Root thick, woody, many-headed, bearing stems and
rosettes; caudices densely covered with remains of previous year’s leaves,
usually white-tomentose. Stem erect, 30-55 cm high, white-tomentose
below, especially near base, glabrous above, smooth or fine-tomentose,
leafy throughout, usually sparsely branched in upper half; branches one-
headed. Leaves typical of section, with almost wingless or very narrow-
winged rachis, lateral segments narrow-lanceolate, acuminate-spinescent,
revolute, glabrous and green above or finely arachnoid-hairy, then
grayish-green, white-tomentose beneath, leaf rachis gray-tomentose; basal
leaves petiolate, petioles ciliate; cauline leaves gradually reduced, sessile,
semiamplexicaul, very short-decurrent, rachis of upper leaves narrow-
winged. Capitula almost broadly cylindrical, turbinate at fruiting, glabrous
or finely arachnoid-hairy, 12—15(20) mm wide (excluding cusps).
Involucral bracts about 60, outer and subsequent bracts 7-12-15 mm
long, coriaceous, green, somewhat flat, divergent, broadly lanceolate,
almost ovate, with prominent midrib on dorsal side, terminating in short
stiff spine, entire or slightly divided with narrow scarious border; middle
and inner bracts erect, 15—20(22) mm long, with membranous, broadly
ovate, pale brown, yellowish or finely and unevenly toothed appendages
with prominent midrib dorsally, cuspidate. Corolla pink, sometimes (less
often) whitish, about 17 mm Iong; receptacular bristles broadened above,
scabrous. Achenes obovoid, 6-7 mm long, glabrous, smooth. Flowering
VII; fruiting VIII-IX.

Gravelly slopes in middle mountain zone.—Soviet Central Asia:
Pamiro-Alai. General distribution: Iran, Afghanistan. Described from
Dabasta. Type in Leningrad.

Note. The species has quite constant characters; only the degree of
pubescence of the plant varies. The most pubescent plants were described

344

338

as C. lachnopoda Juz., which.were also found to have longer outer
involucral bracts. Study of extensive herbarium material showed variation
in these characters throughout the range, and I do not consider it possible
to separate these species. In the region of its contact with C. rubiginosa
Kult., this species forms. hybrids with it, which have characters that are
closer to one or the other parent species. In the original description it is
said that the flowers apparently are pale yellow. Numerous collections
from the classic localities have helped to correct this error, as the corollas
of this species are pink, a fact reported incidentally by the author himself
in a later publication (1940).

256. C. allolepis Tschern. and Vved. in Bot. Mat. Gerb. Inst. Bot.
Akad. Nauk UzbSSR, XVII (1962) 91.

Perennial. Roots woody, many-headed, bearing relatively lax sterile
rosettes of leaves and stems; caudices densely covered with remains of
previous year’s leaves. Stem erect, 20-55 cm high, shallow-sulcate,
glandular-hairy, finely tomentose at base, branched from middle; branches
slender, further branched in one-headed branchlets. Leaves green, strongly
scabrous above due to numerous glandular hairs, gray-tomentose beneath,
pinnatisect with narrow-lanceolate revolute acuminate-spinescent
segments, on narrow-winged rather densely pubescent rachis not in one
plane, appearing in whorls; outer basal leaves with sheath-like broadened
petioles and short, linear-cristate lamina; inner basal leaves petiolate,
petioles flattened toward base, with numerous thin spines along margin;
cauline leaves gradually reduced upward, like basal but sessile,
semiamplexicaul, short-decurrent, with wider rachis wings. Capitula less
numerous, on densely glandular-hairy peduncles, broadly cylindrical, 17
mm wide (excluding cusps). Involucral bracts crisped-glandular-hairy,
about 35, weakly carinate, usually dark colored above; outer bracts
horizontal, lanceolate, up to 17 mm long, acuminate-spinescent, broad
at base, 3-5 mm wide, sometimes with one to three pairs of lateral
spines along margin; middle bracts erect-spreading, lanceolate, cuspidate-
spinescent, 17-18 mm long, up to 4 mm wide at base; inner bracts
membranous, linear, acuminate, very short and stiff spinescent. Corolla
yellow, 14-15 mm long; anther tube pale purple; receptacular bristles
scabrous. Achenes oblong-obovoid, glabrous, smooth, 5—6 mm long.
Flowering VIII; fruiting IX.

Stony slopes in upper mountain zone.—Soviet Central Asia: Pamiro-
Alai. Endemic. Described from the Tamshush River Basin. Type in
Tashkent.

257. C. newesskiana Winkl. in Tr. Peterb. Bot. Sada, XI (1890)
333; XII, 258; XIV, 209; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8,
564.

345

339

Perennial. Root thick, woody, many-headed; caudices with numerous
remains of previous year’s leaves. Stem erect, 20-40 cm high, striate-
sulcate, arachnoid-hairy, sparsely branched in upper one-third part;
branches sparsely leafy, one-headed; inflorescence lax paniculate. Leaves
appressed arachnoid-hairy, grayish-green, and sparsely glandular above,
white-tomentose beneath; outer (lower) basal leaves strongly reduced,
with sheath-like broadened petioles; inner (upper) basal leaves short-
petiolate, with somewhat flat, finely spinose petioles, lanceolate,
pinnatisect in narrow linear, 1—-1.5 mm wide, subulate, revolute lateral
segments, as if in whorl on very narrow-winged, densely seriaceous
rachis; cauline leaves gradually reduced toward stem apex, like basal
leaves but sessile, short-decurrent, with wider winged leaf-rachis;
uppermost leaves strongly reduced. Capitula hemispherical-cylindrical,
about 12-15 mm wide (excluding cusps), sparsely arachnoid-hairy.
Involucral bracts about 60, carinate; outer bracts horizontal or weakly
decurved, lanceolate, acuminate-spinescent, 12—15 mm long;- other bracts
erect, closely appressed, slightly divergent only in upper one-third part;
middle bracts lanceolate, acuminate-short-spinescent, 12 mm long; inner
bracts almost membranous, lanceolate, acuminate, finely toothed, scabrous
dorsally, not exceeding middle bracts, corolla yellow; receptacular bristles
broadened above, scabrous. Achenes obovoid. Flowering VII—VIII; fruiting
1X

Stony screes from middle to upper mountain zone——Soviet Central
Asia: Pamiro-Alai. Endemic. Described from the Chakmankuidy Ravine
and Lagorimurd (Nevesskii). Type in Leningrad.

258. C. trichopora Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIllIc,
VI (1929) 26; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 566.

Perennial. Roots woody, many-headed; caudices with numerous
remains of previous year’s leaves. Stem erect, 25-40 cm high, densely
sericeous, branched in upper half; branches short, densely leafy, branched
further in short one-headed branchlets; inflorescence paniculate. Leaves
covered above with papilliform glandular hairs, alternating with long,
silky simple hairs, gray-tomentose beneath; outer basal leaves with
sheath-like broadened petioles, with numerous thin spines along margin,
pinnatisect in considerably smaller lateral segments, borne in whorl on
leaf rachis; inner basal leaves short-petiolate; pinnatisect in narrow-linear,
up to 1.5 mm wide, revolute lateral segments, as if in whorl on narrow-
winged revolute rachis, densely sericeous below; cauline leaves gradually
reduced upward, like basal leaves but sessile, short-decurrent. Capitula
ovoid-globose, about 12-18 mm wide (excluding cusps), very densely
pubescent with rusty, silky hairs. Involucral bracts about 50, carinate;
outer bracts horizontal, lanceolate, usually as long as middle; middle

346

340

bracts erect, divergent above, lanceolate, acuminate-spinescent, 7-15 mm
long; inner bracts almost membranous, narrow-lanceolate, acuminate,
not exceeding middle bracts. Corolla yellow; receptacular bristles
broadened above, scabrous. Achenes oblong-obovoid, 5 mm long, usually
not ribbed. Flowering VII—VIII; fruiting VIII-IX.

Stony and gravelly slopes in upper mountain zone.—Soviet Central
Asia: Pamiro-Alai. Endemic. Described from mountain slopes in the
vicinity of Yakkabag. Type in Tashkent.

Note. This species is very close to C. newesskiana Winkl. and forms’
hybrids with it where the species are sympatric.

259. C. schugnanica Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8
(1940) 525, 564.—C. newesskiana auct. non Winkl.: Rech. f. in Dan.
Biol. Skr. 8, 2 (1955) 132 (?).

Perennial. Root thick, woody, many-headed; caudices densely
covered with numerous remains of previous year’s leaves. Stems erect,
relatively weak, 20-30 cm high, finely arachnoid-hairy, leafy, usually
sparsely branched from middle or above; branches one-headed. Leaves
typical of section, with almost wingless, densely gray arachnoid-hairy
rachis; lateral segments almost linear, terminating in yellowish, short,
stiff cusp, sparsely arachnoid-hairy and grayish-green above, almost
white-tomentose beneath; basal leaves short-petiolate, others sessile, short-
decurrent. Capitula cylindrical, 15-18 mm wide (excluding cusps), finely
rusty-tomentose. Involucral bracts about 80, all except inner very similar,
narrow-lanceolate, weakly erect spreading, coriaceous, rather weak,
slightly carinate, terminating in thin yellowish cusp, 7-11 mm long,
usually dark colored; inner bracts flat, narrow-lanceolate, membranous,
short acuminate, scabrous along margin, sparsely scabrous dorsally.
Corolla yellow, about 17 mm long; anther tube pale pink; receptacular
bristles strongly scabrous. Achenes obovoid, narrow-ribbed, 6 mm long.
Flowering VIII; fruiting VIII-IX.

Gravelly slopes in upper mountain zone.—Soviet Central Asia:
Pamiro-Alai. General distribution: Afghanistan. Described from the
Vakhan-Ishkashim District, slopes of the Nishgor River. Type in
Leningrad.

Note. The species was described without specific citation of the
type; therefore, Juzepczuk’s reference to the “Slopes of the Nishgor and
Mats rivers” pertains to the two plants collected by P.N. Ovczinnikov.
I am selecting as the lectotype the specimen with the label “Vakhan-
Ishkashin District, eastern slope of the Nishgor River, 5—6 km from its
mouth, 14 VIII 1935. Ovczinnikov and Afanas. ev No. 1955,” because
a careful study of the characters, given in the key, leads to the conclusion
that the plants particularly from the mouth of the Nishgor River should

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341

be considered as C. schugnanica. The plants from the vicinity of Mats
River have a whitish corolla, and it is better to refer them to C.
rubiginosa.

260. C. sororia Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8 (1940)
525, 564.

Perennial. Root woody, many-headed; caudices densely covered with
remains of previous year’s leaves. Stem erect, about 50 cm high, finely
tomentose, glabrescent, leafy, sparsely branched above, with short one-
headed branches. Leaves typical of section, with narrow-winged white-
tomentose rachis, lateral segments almost linear, acuminate into stiff
short spine, revolute, green and glabrous above, gray-tomentose beneath;
basal leaves petiolate; cauline leaves gradually reduced, sessile,
semiamplexicaul, short-decurrent. Capitula almost cylindrical, turbinate
at fruiting, 12-15 mm wide (excluding cusps), subglabrous. Involucral
bracts about 60; outer bracts lanceolate, coriaceous, weakly carinate,
acuminate, almost horizontal or slightly decurved, 7-15 mm long; middle
bracts erect, somewhat flat, 12-15 mm long, broadly lanceolate, obtuse,
spinescent, with membranous border; inner bracts erect, up to 20 mm
long, with compact scarious, pale straw-colored appendages at apex,
unevenly divided along margin and short spinescent. Corolla yellow,
16-18 mmelong; receptacular bristles broadened, scabrous. Flowering
VII.

_ Gravelly slopes.—Soviet Central Asia: Pamiro-Alai. Endemic.
Described from Bogushdar. Type in Leningrad.

Note. Only one location of this species is known, whose separate
status is somewhat doubtful. This plant is very similar in its characters
to C. semilacera Juz., from which it differs only by the color of the
corolla (yellow!). The discovery of taller C. schugnanica Juz. around
Pyandzh, which also has deep yellow corollas, leads to the hypothesis
that C. sororia Juz. is of hybrid origin, which can be confirmed only by
further study of plants from this region. At Gunt we find plants that
very much resemble C. sororia, but have smaller capitula and a pale
yellow corolla. Could this also be a hybrid of C. semilacera with some
other yellow-flowered cousinia from the Shugnan Range? The paucity
of material makes it impossible at present to solve this question.

261. C. abbreviata Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 87.

Perennial. Roots woody, many-headed, bearing sterile rosettes of
leaves and stems; caudices densely covered with remains of previous

year’s leaves. Stem erect, finely tomentose, 20-30 cm high, leafy,

branched above, with short one-headed branches. Leaves typical of

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342

section, with almost wingless rachis and numerous narrow-lanceolate,
convolute, acuminate-spinescent lateral segments, green and very finely
and sparsely arachnoid-hairy above with fine glandular hairs, white-
tomentose beneath; basal leaves short-petiolate; cauline leaves gradually
reduced, sessile, semiamplexicaul, short-decurrent. Capitula narrow-
cylindrical, 13-14 mm long (excluding corolla) and 8-10 mm wide
(excluding cusps), finely arachnoid-hairy. Involucral bracts somewhat
flat, with erect-spreading cusp, dark-colored; outer and middle bracts
similar, 5-11 mm long, lanceolate, acuminate; inner bracts up to 14
mm long, scariously broadened above and acuminate-short spinescent,
dorsally glabrous. Corolla yellow, 12-13 mm long; receptacular bristles
scabrous. Flowering VIII.

Stony slopes in upper mountain zone.— Soviet Central Asia: Pamiro-
Alai (Pamir). Endemic. Described from the slopes of the pass into Garm-
Chashma. Type in Leningrad.

262. C. ramulosa Rech. f. in Dan. Biol. Skr. 8, 2 (1955) 149, fig.
101, 103F.—Iec.: Rech. f. Ibid.

Perennial. Roots woody, many-headed; caudices with few remains
of previous year’s leaves. Stem erect, finely flocculose-tomentose, 40—
55 cm high, profusely branched; branches slender, spreading, usually
exceeding main stem and further branched into one-headed branchlets,
forming lax, almost corymbose inflorescence. Leaves typical of section,
with narrow-winged, rather weak rachis and relatively less numerous
narrow-lanceolate, short and thin acuminate lateral segments, green and
glabrous above, white-tomentose beneath; basal leaves petiolate, petioles
broadened at base; cauline leaves reduced, with wider winged rachis
and wider lanceolate lateral segments, sessile, short-decurrent, leaves
on branches strongly reduced, sessile, with wide and large terminal
segment. Capitula narrow-cylindrical, 15-17 mm long (excluding
corollas) and 8—-9(11) mm wide (excluding cusps), finely arachnoid-hairy,
glabrescent; capitula on main stem usually larger than those on branches.
Involucral bracts about 40—50, all coriaceous, compactly imbricate, with
somewhat short, about 2-3 mm long, divergent cusp; outer and middle
bracts 3-10 mm long; inner bracts 12-15 mm long, strongly scabrous
due to dense spinules; involucral bracts of capitula larger on main stem,
outer divergent, 14-16 mm long, middle 12-13 mm long, erect, inner
up to 17 mm long, strongly scabrous due to dense spinules. Corolla
sulfur-yellow; anther tube pink, about 16 mm long; receptacular bristles
scabrous. Achenes obovoid, 5—6 mm long, narrow-ribbed, finely toothed
above, gray. Flowering VIII; fruiting VIII-IX.

Stony-loamy slopes in middle mountain zone—Soviet Central Asia:
Pamiro-Alai (Pamir). General distribution: Afghanistan. Described from
Afghanistan (Kash). Type in Vienna.

349

343

Note. Our plants differ somewhat from the type specimen of C.
ramulosa by the sparser pubescence of capitula.

263. C. acrodroma Tschern. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XVII (1962) 89.

Perennial. Roots woody, many-headed, bearing sterile rosettes of
leaves and stems; caudices densely covered with remains of previous
year’s leaves. Stems numerous, erect, simple, one-headed, less often
two-headed, leafy, flocculose-arachnoid, 27-40 cm high. Leaves typical
of section, with stiff, narrow-winged, gray-tomentose rachis and narrow-
lanceolate, almost linear, acuminate-spinescent, revolute lateral segments,
finely appressed arachnoid-hairy and grayish-green above, white-
tomentose beneath; basal leaves petiolate, petioles short, broadened
basally, ciliate; cauline leaves sessile, very short-decurrent, gradually
reduced upward. Capitula globose, weakly truncate, 12-15 mm wide
(excluding cusps) and 12-15 mm long (excluding corolla), arachnoid-
hairy. Involucral bracts dark-colored, somewhat flat, with prominent
midrib, short-acuminate; outer and middle bracts similar, lanceolate,
slightly divergent, 7-9 mm long; inner bracts erect, broadened above in
membranous, brownish, up to 15 mm long appendage somewhat incised
along margin. Corolla yellow, about 15 mm long, receptacular bristles
scabrous. Flowering VII. (Plate XXVIII, Fig. 2.)

Stony slopes.—Soviet Central Asia: Pamiro-Alai (Pamir). Endemic.
Described from the Sharf-Dara Valley. Type in Leningrad.

Note. This species is quite close to C. schugnanica, but is well
distinguished by the strong, erect, one-headed stems and by the presence
of the membranous appendages of the inner involucral bracts.

264. C. rubiginosa Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIIIc,
VI (1929) 23; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 564.

Perennial. Roots thick, woody, many-headed, bearing sterile leaf
rosettes and stem; caudices white-tomentose, densely covered with
remains of previous year’s leaves. Stem erect, 30-40 cm high, striate-
sulcate, finely flocculose-tomentose, white-tomentose below; sparsely
glandular under pubescence, leafy. Leaves green above, sparsely
glandular, white-tomentose beneath, pinnatisect in narrow-lanceolate or
almost linear, revolute, acuminate-spinescent segments, in turn,
palmatisect, on narrow-winged petiole not in one plane, as if in whorl,
petioles ciliate along margin; other leaves sessile, semiamplexicaul.
Capitula globose, 2 cm wice (excluding cusps), usually densely rusty-
tomentose, somewhat truncate basally. Involucral bracts numerous; outer
bracts 15 mm long, deflexed from deltoidly broadened appressed base,
terminating in short triquetrous spine; middle bracts 18 mm long, 4—6
mm wide at base, divergent from appressed base, terminating in stiff,

350

344

somewhat short, triquetrous spine, dorsally scabrous, densely rusty-
tomentose; subsequent bracts erect, acuminate-spinescent, scabrous along
margin, about 20 mm long; inner bracts about 20 mm long, erect,
scariously broadened above, erose, densely scabrous dorsally and along
margin up to scarious border, scarious part glabrous dorsally with
prominent vein, terminating in very short spine. Corolla whitish (7),
17-18 mm long; anther tube purple; receptacular bristles scabrous.
Achenes obovoid, ribbed, narrow-winged along us. Flowering VII-”%
VIN; fruiting VIII.

Stony and gravelly slopes in middle mountain zone.—Soviet Central
Asia: Pamiro-Alai (Vakhsh, Rushan, Shugnan). Endemic. Described
from the region of the Odudi Pass (slope of Kala-i-Vamar). Type in
Tashkent.

Note. Forms hybrids with C. semilacera Juz.

265. C. multiloba DC. Prodr. VI (1837) 554; Bge. in Mém. Acad.
Pétersb. 7 sér. 9, 2 (1865) 46; Boiss. Fl. or. III, 499 p. p. (excl. syn. C.
pycnoloba Boiss.); Winkl. in Tr. Peterb. Bot. Sada, XI (1890) 136; XII,
260; XIV, 231.—C. palmatiloba Jaub. and Sp. Ill. or. (1844-1846) tab.
162.—Arctium multilobum O. Ktze. in Rev. Gen. 1 (1891) 308.—Ie.:
Jaub and Sp. op. cit. tab. 162.

Perennial. Roots woody, many-headed; caudices densely covered
with remains of previous year’s leaves. Stem erect, appressed white-
tomentose, leafy, 50-60 cm high, weakly branched in upper haif, leafy;
branches one-headed. Leaves typical of section, with narrow-winged,
white-tomentose petioles and almost linear, acuminate-spinescent,
revolute lateral segments, appressed arachnoid-hairy and grayish green
above, appressed white-tomentose beneath; basal leaves petiolate, petioles
white-tomentose, somewhat broadened at base and spinose; lower cauline
leaves like basal but with short and wider petioles; other cauline leaves
sessile, semiamplexicaul, short-decurrent, gradually reduced toward apex;
uppermost leaves strongly reduced, with long, acuminate spinescent
terminal lobe. Capitula ovoid, 13-15 mm wide (excluding cusps), densely
white arachnoid-hairy. Involucral bracts about 70; outer bracts almost
horizontal, 12-16 mm long, lanceolate, scabrous dorsally above,
acuminate; middle bracts about 20 mm long, erect-spreading, otherwise
like outer; inner bracts narrow-lanceolate, 15-17 mm long, short-
acuminate, scabrous dorsally above. Corolla pink; receptacular bristles
scabrous; achenes obovoid, 6 mm long, longitudinally narrow-ribbed,
dentate above. Flowering VII—VIII; fruiting VIII-IX.

Stony slopes.—Soviet Central Asia: mountainous Turkmenia
(Kopetdag). General distribution: Afghanistan, Iran. Described from Iran.
Type in Paris.

345

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Plate XXVIII.
1—Cousinia splendida Wink|.; 2—C. acrodroma Tschern.

351

352

346

266. C. eriotricha Juz. in Tr. Bot. Inst: Akad. Nauk SSSR, Ser. 1,
3 (1936) 321; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 564.

Perennial. Roots thick, woody, many-headed; caudices densely
covered with remains of previous year’s leaves. Stem erect, 25-40 cm
high, striate-sulcate, finely white-tomentose, with long entangled hairs,
leafy, usually weakly branched above; branches short, one-headed. Leaves
typical of section, green above, white-tomentose beneath, densely covered
on both sides with fine, long, silky, entangled, divergent hairs, especially
on petiole; petiole almost entirely wingless with linear, acuminate-
spinescent, revolute, lateral segments; basal leaves short-petiolate; cauline
leaves sessile, short-decurrent. Inflorescence less-branched, corymbose-
paniculate; capitula oblong-ovoid, 12-14 mm wide (excluding cusps),
arachnoid-hairy or more or less finely white-tomentose. Involucral bracts
80-90, linear-subulate, erect-spreading; outer bracts 10-12 mm, middle
up to 18 mm long; inner bracts flat, membranous, linear-lanceolate, 20—
22 mm long, acuminate, scabrous dorsally and along margin. Corolla
pink, about 20 mm long; receptacular bristles somewhat broadened above,
scabrous. Achenes obovoid, 7 mm long, glabrous, smooth. Flowering
VII-VIII; fruiting VIII.

Gravelly slopes in upper mountain zone.—Soviet Central Asia:
Pamiro-Alai (Turkestan Range). Endemic. Described from the north slope
of the Turkestan Range, near the village of Bachmazar. Type in
Leningrad.

Note. Several plants from different localities were mentioned in the
description of this species. I consider as the lectotype the plant with the
label “North slope of the Turkestan Range, ravine near the village of
Bachmazar. 1914, VII, 2. Michelson.”

Hybrids of C. eriotricha Juz. with C. verticillaris Bge. are known
from the eastern part of its range.

267. C. laetevirens Winkl. in Tr. Peterb. Bot. Sada, X (1887) 576;
XII, 261; XIV, 231; Bornm. in Beih. bot. Centralbl. XXXIV, 2, 188;
Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 564.—C. laetevirens var.
monocephala Winkl. op. cit. (1887) 576.—C. polyothrix Winkl. ibid.
(1887) 577; XII, 260.—C. bonvalotii var. poliotrichodes Bornm. in Beih.
bot. Centralbl. XXXIV, 2 (1916) 185-186.

Perennial. Roots woody, many-headed, bearing sterile rosettes of
leaves and stems; caudices densely covered with numerous remains of
previous year’s leaves. Stems numerous, erect, 30-50 cm high, striate-
sulcate, finely arachnoid hairy-pubescent below, leafy, very sparsely
branched above; branches usually short. Leaves green, sparsely glandular,
or finely arachnoid-hairy and not glandular above, gray-tomentose
beneath, pinnatisect with many narrow-linear, acuminate-spinescent,

347

353 revolute lateral segments, as if in whorl on almost wingless rachis; basal
leaves petiolate; cauline leaves gradually reduced, sessile, short-decurrent.
Capitula cylindrical, 10-15 mm wide (excluding cusps), pubescent.
Involucral bracts 30-40, green, sometimes dark-colored; outer and
subsequent bracts distinctly different from others, carinate, usually
divergent, often horizontal, acuminate into short cusp, 15—20(30) mm
long, 2-3 mm wide at base, 1-1.5 mm in middle; middle bracts
coriaceous, somewhat flat, short-acuminate, 8—10(12) mm long; inner
bracts membranous, lanceolate, acuminate-spinescent, scabrous dorsally
and along margin, 10—12(15) mm long. Corolla pink, 16-18 mm long,
exserted; receptacular bristles slightly broadened above, scabrous.
Achenes obovoid, 6—7 mm long, glabrous, smooth. Flowering VII-VIII;
fruiting VIII-IX.

Stony slopes in upper mountain zone.—Soviet Central Asia: Pamiro-
Alai (Zeravshan Range, Gissar Range and its southwestern spurs up to
Kugitang, Peter the Great Range). Endemic. Described from Malguzar
(Rgl.). Type in Leningrad.

Note. The species is very close to C. verticillaris Bge., from which
it differs only by smaller capitula and quite long, horizontally spreading
outer involucral bracts, equaling or exceeding the diameter of the
capitulum.

It forms hybrids with this species where they are sympatric.

The species was based on plants collected from two localities; |
propose to designate as the lectotype of C. laetevirens Winkl. the plant
with the label “Sarawschan-Gebiet: Margusar am Pasrut. 9000’ 21, VI-
8 VII, 1882. Regel.”

The plants from Kugitan and the southwestern spurs of the Gissar
Range differ somewhat from typical C. laetevirens Winkl. by more
densely pubescent capitula and strongly rugose outer involucral bracts;
it is possible that further study of additional material from these regions
will permit us to separate them as independent species.

268. C. verticillaris Bge. in Mém. Acad. Pétersb. sav. étrang. 7
(1851) 357; Bge. in Mém. Acad. Pétersb. 7 sér. 9, 2, 46; Boiss. FI. or.
III, 500 (excl. C. verticillaris var. major Boiss.); Winkl. in Tr. Peterb.
Bot. Sada, XII (1892) 259 (quoad sp. Lehmann); XIV, 230; Bornm. in
Beih. bot. Centralbl. XXXIV, 2, 186; Juz. in Tr. Tadzh. Bazy Akad.
Nauk SSSR, 8, 566.—C. hissarica Winkl. op. cit. (1887) 92; XII, 259;
XIV, 209; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 565.—C.
verticillaris var. leiocephala Bornm. op. cit. (1916) 187.—C. poliothrix
Winkl. op. cit. (1897) 230.—C. pseudo-bonvalotii Juz. op. cit. (1940)
525, 565.—C. poliothrix f. albiflora Bornm. op. cit. (1916) 187.—
Arctium verticillare O. Ktze. Rev. Gen. 1 (1891) 308.—Cousinia

354

348

laetevirens auct. non Winkl.: Winkl. op. cit. (1897) 231 (quoad pl. e
Revout).—C. verticillaris var. major. auct. non Boiss.: Bornm. op. cit.
(1916) 187.—C. poliothrix var. robusta auct. non Winkl.: Bornm. Ibid.
187.

Perennial. Roots woody, many-headed, bearing sterile rosettes of
leaves and stems; caudices densely covered with numerous remains of
previous year’s leaves. Stem erect, 10-40 cm high, striate-sulcate,
pubescent, sparsely branched above, leafy. Leaves green and appressed
arachnoid-hairy above, gray-tomentose beneath, pinnatisect in many,
narrow-linear, acuminate-spinescent, revolute, lateral segments in whorl
on almost wingless rachis; basal leaves petiolate; cauline leaves gradually
reduced upward, like basal but sessile, short-decurrent. Capitula broadly
cylindrical, 15-22 mm wide (excluding cusps), pubescent. Involucral
bracts 50-60, carinate; outer bracts squarrose, sometimes weakly
decurved, 7-15 mm long and 2 mm wide at base; middle bracts like
outer, 10-15 mm long, up to 3 mm wide at base, spinescent like outer;
inner bracts membranous, narrow-lanceolate, scabrous dorsally and along
margin, terminating in thin short cusp, 15-17 mm long. Corolla pink,
somewhat exserted, 15-17 mm long; receptacular bristles weakly
broadened above, scabrous. Achenes obovoid, 6-7 mm long, glabrous,
smooth. Flowering VII—VIII; fruiting VIII-IX.

Gravelly slopes in upper mountain zone.—Soviet Central Asia:
Pamiro-Alai. Endemic. Described from the Zeravshan River basin
(Karatau). Type in Paris.

Note. Until recently, C. verticillaris Bge., described from Lehmann’s
collections from the Karatau Range, was considered a doubtful species.
S.V. Juzepezuk expressed the view that C. hissarica Winkl. was described
under this name, which is fully confirmed by study of the photograph
of the type specimen sent from Paris.

Hybrids of C. verticillaris Bge. and C. laetevirens Winkl. are known
from the areas where they are sympatric.

269. C. bonvalotii Franch. in Ann. Sc. Nat. 6 sér., 16 (1883) 320;
Winkl. in Tr. Peterb. Bot. Sada, XII, 258.—Arctium bonvalotii O. Ktze.
Rev. Gen. | (1891) 307.—Exs.: H.F.A.M. No. 42.

Perennial. Roots woody, many-headed; caudices with numerous
remains of previous year’s leaves. Stems several, erect, 20-40 cm high,
finely tomentose, arachnoid-hairy above, weakly branched; branches
short, one-headed. Leaves typical of section, with broadly lanceolate,
broadly excurrent lateral segments on narrow-winged rachis (wings 1—
1.5 mm wide); finely appressed-pubescent or sparsely glandular above,
gray-tomentose beneath; basal leaves short-petiolate; cauline gradually
reduced upward, sessile, short-decurrent, with wide-winged rachis.

355

349

Capitula cylindrical, 8(12) mm wide (excluding cusps), finely arachnoid-
hairy. Involucral bracts relatively less numerous, about 40, carinate; outer
bracts horizontal, narrow-lanceolate, up to 7 mm long, sometimes up to
2 mm wide at base; middle bracts erect-spreading, narrow-lanceolate,
10-12 mm long; inner bracts almost membranous, linear, acuminate,
sometimes blackish dorsally in middle, 10-12 mm long. Corolla pink;
receptacular bristles somewhat broadened above, weakly scabrous.
Achenes oblong-obovoid, compressed dorsally, roundish above, about 5
mm long. Flowering VII—VIII; fruiting VIII.

Gravelly-stony slopes in upper mountain zone.—Soviet Central Asia: -
Tien Shan (western). Endemic. Described from the vicinity of Ona-
Oulgan. Type in Paris.

270. C. ceratophora Kult. in Tr. Sredneaz. Gos. Univ. Ser. VIIIc,
VI (1929) 24; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 565.
Perennial. Roots thick, woody, many-headed, bearing sterile rosettes
of leaves and stems. Stem erect, 35-40 cm high, glabrous, leafy, very
sparsely branched above, branches one-headed. Leaves typical of section,
with almost wingless, rather strong rachis and more or less linear, short-
acuminate, revolute lateral segments, green, very finely glandular above,
gray-tomentose beneath; basal leaves petiolate, petioles robust, armed,
slightly thickened at base; cauline leaves sessile, gradually reduced, short-
decurrent. Capitula 12-20 mm wide (excluding cusps), about 20 mm
long (excluding corolla), globose, glabrous. Involucral bracts less
numerous, 25-30, flat; outer bracts lanceolate, about 15 mm long and
5—8 mm wide at base, decurved; middle bracts broadly lanceolate,
carinate above, 20—25(28) mm long and 7-10 mm wide at base, more
or less erect-spreading; inner bracts with brownish, membranous, scabrous
on margin apical appendage, lanceolate, erect, acuminate-short cuspidate.
Corolla whitish (?); anther tube purple; receptacular bristles scabrous.
Achenes obovoid, light gray, with dark spots, 7 mm long, slightly
compressed dorsally, longitudinally narrow-ribbed. Flowering VIII;
fruiting IX.
Stony-loamy slopes in upper mountain zone.—Soviet Central Asia:
Pamiro-Alai (Darvaz). Endemic. Described from the Viskharvi Pass. Type
in Tashkent.

271. C. serawschanica Winkl. in Tr. Peterb. Bot. Sada, IX (1886)
424; XII, 261; XIV, 210; Bornm. in Beih. bot. Centralbl. XXXIV, 2,
188; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 565.—C.
sarawschanica B. elegantula Bornm. op. cit. (1916) 188.—C. laetevirens
var. flava Winkl. op. cit. (1897) 231.—C. flava Bornm. op. cit. (1916)
280; Juz. op. cit. 565.

350

Perennial. Roots thick, woody, many-headed, bearing rosettes of
sterile leaves and stems; caudices densely covered with remains of
previous year’s leaves. Stem erect, 15—40 cm high, shallow striate-sulcate,
glandular above, gray-pubescent below, sometimes white-tomentose,
leafy. Leaves usually green and sparsely glandular above, less often
finely gray-pubescent, gray-tomentose beneath, especially along midrib,
pinnatisect in narrow-lanceolate, revolute, acuminate-spinescent, almost
linear segments on very narrow-winged rachis, not in one plane, but as
if in whorl; basal and lower cauline leaves on short, weakly sulcate,
ciliolate, usually gray-pubescent petioles; other leaves gradually reduced
toward apex, semiamplexicaul, uppermost leaves decurrent. Capitula
numerous, on glandular-hairy peduncles, oblong-cylindrical, 8—12(15)
mm wide (excluding cusps). Involucral bracts glandular, pubescent; outer
bracts carinate, 10-30 mm long, horizontal, gradually acuminate from
closely appressed base to grooved cusp, terminating in short yellowish
spine; middle bracts like outer but slightly shorter and usually somewhat
erect, sometimes only slightly divergent; inner bracts somewhat flat,
11-16 mm long, broadly lanceolate, acuminate-spinescent. Corolla
yellow, 12-15 mm long; anther tube pale purple; receptacular bristles
scabrous. Achenes obovoid, glabrous, smooth, 6—7 mm long. Flowering
VII-VIII; fruiting IX.

Stony slopes in upper mountain zone.—Soviet Central Asia: Pamiro-
Alai. Endemic. Described from Santuruch near the pass from Kshtut to
Lake Kulikalon. Type in Leningrad.

Note. Pink florets are mentioned in the original description of C.
serawschanica, which is definitely wrong. This error was committed
apparently, because Winkler saw only young plants and determined the
color of the corolla from buds, which, probably, because of the purple
anther tubes, appeared pinkish. Numerous specimens from the region of
origin of the type specimen confirm the presence not of pink but yellow
corollas in C. serawschanica.

272. C. caespitosa Winkl. in Tr. Peterb. Bot. Sada, X (1887) 93;
XII, 259; XIV, 230; Bornm. in Beih. bot. Centralbl. XXXIV, 2, 186;
Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8, 565.—Exs.: H.F.A.M.
No. 43.

Perennial. Roots woody, many-headed; caudices with remains of
previous year’s leaves. Stems few, erect, white arachnoid-hairy or white-
tomentose, 15-40 cm high, usually simple, one-headed. Leaves soft,
gray appressed-arachnoid hairy on both sides, sometimes upper surface
more sparsely arachnoid-hairy, greenish; basal leaves short-petiolate,
pinnatisect in narrow-lanceolate, acuminate-spinescent, decurved lateral
segments in one plane, leaf rachis narrow-winged, wings about 2 mm

357

351

wide, revolute; cauline leaves gradually reduced toward stem apex,
sessile, semiamplexicaul, short-decurrent, almost pinnatipartite with
entire, broadly lanceolate lateral lobes. Capitula cylindrical or almost
ovoid, 10-15 mm wide (excluding cusps), densely pubescent, less often
finely arachnoid-hairy. Involucral bracts about 100, all except inner
similar, narrow-lanceolate, weakly divergent; outer bracts usually
decurved, 7-17 mm long, inner bracts erect, 16-18 mm long, narrow-
lanceolate, compact-scarious above, whitish, glabrous dorsally, smooth,
weakly scabrous along margin. Corolla pink; receptacular bristles
somewhat broadened above, more or less scabrous. Achenes obovoid,
6 mm long, with filiform longitudinal ribs. Flowering VII—VIII; fruiting
VIII.

Gravelly and stony slopes in upper mountain zone.—Soviet Central
Asia: Tien Shan (predominantly western Tien Shan), Pamiro-Alai (Alai
Vailey). Endemic. Described from the Susamyr Mountains. Type in
Leningrad.

DOUBTFUL SPECIES

1. C. bucharica Winkl. in Tr. Peterb. Bot. Sada, X (1889) 477;
Winkl. ibid., XII, 196; Juz. in Tr. Tadzh. Bazy Akad. Nauk SSSR, 8,
555.—Arctium bucharicum O. Ktze. Rev. Gen. 1 (1891) 307.

Described from the region between the city of Kabadian and
Kurganbulak spring. Type in Leningrad.

This species is based on the tip of a stem of a very young plant
and even now, when we lay out the extensive herbarium material
accumulated during recent years from the range of this species, it is
difficult to judge from the description and type specimen to which species
this name applies. Its inclusion under the subsection Jurineopsis, as done
by S.V. Juzepezuk, also seems doubtful.

2. C. trachylepis var. integrifolia O. Ktze. in Tr. Peterb. Bot. Sada,
X (1887) 197.

Described from Kopetdag, vicinity of Ashkhabad. This plant is not
in the herbaria of Leningrad. Winkler also did not see it; therefore, it is
difficult to express a definite opinion about it.

3. C. xiphiolepis Boiss. Diagn. pl. or. nov. ser. 1, 6 (1845) 121; O.
Ktze, in Tr. Peterb. Bot. Sada, X, 197.

This species was described from Iran. We could not ascertain which
material from Turkmenia was reported under this name by Kuntze
because the plant is not in the herbarium and Winkler also did not
see it.

358

352
GENUS 1579. Lipskyella Juz.':2

Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3 (1936) 323.—
Cousinia sect. Dichacantha Lipsky. Bot. Eksk. za Kaspii (1889) 13.—
Cousinia subgen. | Dichacantha Winkl. in Tr. Peterb. Bot. Sada, XII, 2
(1892) 184.

Capitula homogamous, all florets bisexual, tubular; involucral bracts
imbricate, usually all membranous, very short-spinescent, divergent at
fruiting; receptacle flat, setose, achenes several times as large as
receptacular bristles; filaments glabrous, smooth; basal appendages of
anthers barbed; achenes with straight attachment, narrow-prismatic,
transversely alveolate-reticulate, two or three winged, terminating into
two or three teeth above; pappus bristles scabrous, all equal, persistent
in fruits or subsequently shedding individually.

Annual herb with almost undivided leaves, usually spiny throughout,
mainly along veins.

Type species: L. annua (Winkl.) Juz.

1. L. annua (Winkl.) Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser.
1, 3 (1936) 324.—Cousinia annua Winkl. in Tr. Peterb. Bot. Sada, X
(1887) 88; XII, 184; Kult. in Fl. Turkm. VI, 163——C. dichacantha
Lipsky, Bot. Eksk. za Kaspii (1889) 12.—Ic.: Kult. op. cit. Plate 24.

Annual. Stem solitary, erect, 15—30 cm high, whitish, glabrous, leafy,
branched; branches short, bearing numerous capitula in almost spicate
inflorescence. Leaves green, broadly lanceolate or almost ovate, attenuate
toward both ends, with isolated 3-4 mm long spines along veins on
upper surface, slightly arachnoid-hairy beneath, almost undivided, with
spines in pairs along margin; basal leaves short-petiolate; cauline leaves
sessile. Capitula on very short peduncles, campanulate-cylindrical, 8—10
mm wide, 12-15 mm long; involucral bracts membranous, lanceolate,
transparent-fimbriate, obtuse, abruptly acuminate into thin, 3-5 mm long,
divergent cusp, sometimes almost horizontal; receptacular bristles smooth,
1—1.5 mm long; corolla whitish (?), with very short limb; achenes 7
mm long, narrow-prismatic, with 1-1.5 mm wide wings. Flowering V;
fruiting VI.

On loose sand.—Soviet Central Asia: Kyzyl-Kum, Kara-Kum.
Endemic. Described from the Rafatak Well in the Chardzhon Region.
Type in Leningrad.

'Treatment by O.V. Tscherneva.
*Named after the famous Russian botanist V.I. Lipsky.

353

=
=e
=

SN

=

NS

‘S

WY.

Plate XXIX.

1—Hypacanthium echinopifolium (Bornm.) Juz.; 2—Anura pallidivirens (Kult.) Tschern.

359

354
GENUS 1580. Hypacanthium Juz.' ”

Juz. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3 (1936) 324.

Capitula homogamous, all florets bisexual, tubular; involucral bracts
imbricate, many-rowed; acuminate-spinescent; receptacular setose, bristles
shorter than achenes, finely serrate-scabrous; filaments glabrous, anther
tube exserted, basal appendages of anthers short, broadened, apically
pointed, fimbriate along margin; styles strongly exserted, deeply bifid,
with weakly divergent branches; achenes with straight attachment,
glabrous, obovoid, almost roundish above; pappus bristles unequal, inner
bristles long, flattened, serrulate, outer gradually reduced, somewhat
scabrous, shedding singly. A perennial herb.

Type species: H. echinopifolium (Bornm.) Juz.

1. H. echinopifolium (Bornm.) Juz. in Tr. Bot. Inst. Akad. Nauk
SSSR, Ser. 1, 3 (1936) 324.—Cousinia echinopifolia Bornm. in Beth.
bot. Centralbl. XXXIV, 2 (1916) 192.—Alfredia? talassica Korov. ex
Ijin in Byull. Glavn. Bot. Sada, 23, 2 (1924) 136.

Perennial. Stems thick, 10—12 mm in diameter, longitudinally ribbed,
flocculose-tomentose, distinctly tall, leafy, sparsely branched in upper
half, with rather long, one-headed branches. Leaves grayish-green and
arachnoid-hairy above, white-tomentose beneath with prominent veins;
basal leaves petiolate, up to 60 cm long (with petiole), oblong,
bipinnatisect, pinnae sessile, decurrent on rachis, acuminate-spinescent,
pinnules narrow-lanceolate, acuminate-spinescent; cauline leaves like
basal but sessile, semiamplexicaul, with few stiff spmnes on both sides
at base, gradually reduced toward stem apex. Capitula flat-globose, 25—
35 mm wide (excluding cusps), strongly foveate at base, with many
florets. Involucral bracts numerous, outer (one row) pinnate-spinulose,
with stiff lateral spines and stiff subulate-triquetrous cusp; other bracts,
except inner, coriaceous, closely appressed, abruptly attenuate at
divergence into stiff, subulate-triquetrous cusp; inner bracts membranous,
brownish above, short-acuminate. Corolla violet-purple, about 18 mm
long. Achenes obovoid, indistinctly three-ribbed, roundish above, dark
gray, 5—5.5 mm long, transversely rugose. Flowering VII—VIII; fruiting
VIUI-IX. (Plate XXIX, Fig. 1.)

Stony screes in upper mountain zone.—Soviet Central Asia: Tien
Shan. Endemic. Described from the vicinity of Namangan. Type in
Leningrad.

'Treatment by O.V. Tschemeva.
*From Greek words: hypo—under, and acantha—spine.

36

—

355
GENUS 1581. Schmalhausenia Winkl.!': 2

Winkl. in Tr. Peterb. Bot. Sada, XII (1892) 281.

Capitula homogamous, all florets bisexual, tubular cupuliform;
involucral bracts imbricate, densely tomentose, acuminate-spinescent;
receptacle flat, setose, bristles smooth, two-fifths as long as achene;
filaments free, glabrous; anthers with short basal appendages; achenes
with straight attachment, five or six ribbed, glabrous, five or six-toothedl
at apex; pappus bristles unequal, readily shedding singly, outer bristles
thinner and shorter, two times as long as achene, short-plumose, inneg
wider, slightly grooved, three times as long as achene, short-plumose.
A prickly perennial herb.

Type species: S. nidulans (Rgl.) Petrak.

1. S. nidulans (Rgl.) Petrak in Allg. Bot. Zeitschr. 20 (1914) 117.—
Cirsium nidulans Rgl. in Bull. Soc. Nat. Mosc. XL, 2 (1867) 160.—
Jurinea horrida Rupr. in Mém. Acad. Pétersb. 7 sér. 14, 4 (1869) 57.—
Cousinia eriophora Rgl. and Schmalh. in Tr. Peterb. Bot. Sada, VI
(1880) 313; Hoffm. in Pflanzenfam. IV, 4-5 (1897) 319.—
Schmalhausenia eriophora Winkl. in Tr. Peterb. Bot. Sada, XII (1892)
281.—Arctium eriophorum O. Ktze. Rev. Gen. 1 (1891) 307.—Carduus
horridus B. Fedtsch. in O. and B. Fedtsch. Perech. Rast. Turk. IV (1911)
278.

Perennial. Root vertical, black-fibrous. Stems erect, 20-40 cm high,
with blackish remains of old petioles at base; densely arachnoid hairy-
tomentose, striate-sulcate, densely leafy, bearing 5—10 capitula in dense
heads. Leaves gray-tomentose on both sides, especially dense along
usually dirty purple midrib; basal leaves oblong-lanceolate, petiolate,
petioles spinose, bipinnatisect, with lanceolate, acuminate-spinescent
pinnules often modified into stiff purple spines at base; cauline leaves
sessile, otherwise like basal. Capitula 4—5 cm wide (excluding cusps),
dirty white-tomentose; involucral bracts narrow-lanceolate, long
acuminate with stiff, subulate, divergent, usually purple, cusp; corolla
purple. Achenes obovoid, transversely rugose. Flowering VII-—VIII;
fruiting VIII-IX.

Alpine and subalpine meadows in upper mountain zone.—Soviet
Central Asia: Tien Shan. Endemic. Described from the Kok-Dzhar Pass.
Type in Leningrad.

'Treatment by O.V. Tscherneva.
*Named after the famous Russian botanist I.F. Schmalhausen.

362

356
GENUS 1582. Saussurea DC.':2

DC. in Ann. Mus. Paris, XVI (1810) 198; Benth. and Hook. f. Gen.
pl. Il (1876) 471; O. Hoffm. in Pflanzenfam. IV, 5 (1894) 320.—
Heterotrichum MB. FI. taur.-cauc. III (1819) 551.—Bennetia S.F. Gray.
Nat. Arr. Brit. PI. II (1821) 440.—Lagurostemon Cass. Dict. sc. nat.
LHI (1828) 466; Wydl. in Linnaea, V, 3 (1830) 425.—Theodorea Cass.
Dict. sc. nat. LIII (1828) 463.—Eriostemon Less. Synops. Compos.
(1832) 12.—Aplotaxis DC. in Guill. Arch. Bot. II (1833) 330; Ej. Prodr.
VI (1837) 538.—Haplotaxis Endlich. Gen. pl. (1836-1840) 468; Ldb.
Fl. Ross. II, 2 (1845-1846) 672.—Cyathidium Lindl. in Royle, Ill. Bot.
Himal. (1839) 251.—Aucklandia Falc. in Trans. Linn. Soc. XIX (1842)
23.—Frolovia (DC.) Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVI (1954) 461.

Capitula homogamous, with many similar florets, in narrow corymbs,
or broad panicles, sometimes solitary. Florets tubular, bisexual, all fertile.
Involucre globose, ovoid, campanulate or cylindrical; involucral bracts
many-rowed, usually imbricate, appressed or outer bracts divergent or
deflected, acute, acuminate, obtuse or subobtuse, often without
appendages, less often with membranous colored appendages, sometimes
with herbaceous saber-shaped appendages. Receptacle flat or convex,
densely or sparsely covered with free scales or less often with yellowish
papillae, sometimes connate to apex, glabrous in some species. Corolla
pink, purple, or dark violet, less often white, with narrow tube and
swollen throat, limb five-fid from middle or below. Anthers with long,
acute apical appendages, and two bristles at base, basal appendages
ciliate, hairy or with woolly tuft; filaments glabrous, free. Stigmas long,
bifid, branches linear, obtuse, dorsally papillate. Achenes glabrous, with
truncate apex, sometimes with fimbria (corona) usually smooth, less
often spinose along ribs, especially in upper part; slightly compressed at
base; cylindrical, or four-angled, straw-colored, brown, with black-purple
dots, less often black, often mottled. Pappus one- or two-rowed; outer
bristles short, toothed or finely plumose, readily shedding, inner always
long-plumose, connate basally into ring, persistent. Perennial or less
often biennial herbs. Sometimes semishrubs, short or tall, branched or
simple. Leaves alternate, often more or less coriaceous, hard or soft, entire
or toothed, crenate-toothed, crenate-lobate, or pinnatifid, glabrous or
pubescent (sometimes woolly); often in same species entire, lobed, or
pinnatifid. The genus Saussurea includes more than 400 species, growing

'Treatment by S.J. Lipschitz.
?Genus named in honor of Theodore de Saussure (1767—1845), Professor of Mineralogy
in Geneva, the famous explorer of the Alps.

363

357

in Eurasia and North America. The main centers of species variation
are the Himalayas, Tibet, China (about 280 species), Korea and Japan.
Within the USSR: Siberia and Far East, Soviet Central Asia.

The unstable nature of the series of characters and the presence of

a large number of hybrids and their segregate offspring make it difficult
to identify some members of the genus.

Type species: S. alpina (L.) DC.

. Achenes spinose, especially in upper part, forming corona (fimbria) .

of long, hard, acute teeth at apex. (Subgenus Stephanodontos
lipschy) colwiicn’ Haleveleosent J, 1. S. carduicephala (Iljin) Iljin
Achenes smooth, without spines, not forming corona; if present,
of different shape (not of long, hard, acute teeth; however, compare
wathinS sechondrilloides,.Winkl.).....<:21s-0eend:.0e MOR dk 2.

. Corymb of numerous capitula enclosed in light-colored scaly

bracteal leaves, in two rows. Root collar stupeous due to fibrous
remains of previous year’s leaves. (Subgenus Amphilaena (Stschegl.)

leipschipiss..2sesionccaent.. 2. S. involucrata Kar. and Kir. ex Maxim.
Whitish scaly bracteal leaves enclosing corymb absent. Root collar
MOGUSTE PONS AcE. cncicennnsoasaneeauoetyestnan eon emma ist Ae chawenu ds adtetesen: 3.
. Stem simple, with solitary terminal capitulum (less often, when
stenvébranched, tcapitulapseveral) fi... Wk OE ask 4.
Capitula aggregated in large or small number in corymbs,
slomeruless pamiclesmori racemes (hice hese eed... 14.

. Plants pulvinate with short, 1.5—2 cm long stems, bearing small

solitary capitula. Basal leaves in rosettes small, pinnate. Pappus-
bristles thickened at tip. (Kuldzha Region, outside the USSR)....
Lalweathcied Jobeiaegho) sn one. leone 33. C. pulviniformis Wink.
Plants of different shape, usually with more or less distinct stem.
Capitula larger. Leaves lyrate-pinnatisect or undivided, only
sometimes weakly toothed, variable in shape (linear, etc.) ...... ah

. Capitula cupuliform, large, (2.5)3—6 cm wide, often drooping.

Involucral bracts lanceolate-linear, outer bracts usually decurved.
Leaves lyrate, broadly ovate or almost triangular. Florets large,
1.5—3 cm long. Receptacle densely-scaly, scales 8-15 mm long.
Achenes usually with border (corona). (Subgenus Frolovia (DC.)
Iipschr)inatiseleet ett loves he eh OY. SER AL cok HAM Res! 6.
Capitula of different shape, smaller, up to 2 cm wide, erect.
Involucral bracts appressed, outer bracts usually not decurved.
Leaves undivided or weakly toothed. Florets not more than 1.5
cm long. Receptacular scales shorter. Achenes without fimbria
(corona) une AS MAL: AMG eRe .. AROIOLO). MIQUE >, 8.

. Stem branched. Involucral bracts light green. Florets pink or dark

358

red, more than 2 cm long including narrow corolla tube (plants of
Soviet: Gentralé Asia). Oe: seieh el. oT, enirelennsht seit. Te
Stem simple. Involucral bracts dark green. Florets dirty violet, about
1.5 cm long including narrow corolla tube (plants of Siberia and
Marbagatal)oarwstts. weeds. see he body 3. S. frolovii Ldb.

. Rhizome tuberous. Basal leaves ovate or almost triangular, crenate-

lobate. Capitula 2—2.5(3) cm wide. Florets 2.3-2.5 cm long
including narrow corolla tube and limb. Achenes mature, lustrous,
about.5 mm long excluding fimbria (corona) ................:cccccceseeeees
Lise aie eae a ae AI, 3 5. S. sulcata [Ijin
Rhizome not tuberous. Basal leaves lyrate. Capitula very large, 3—
6 cm wide. Florets large, 3—3.2 cm long including narrow corolla
tube and limb. Achenes about 7 mm long, with short but distinct

fimibria? (corona), cf ee. TR 4. S. asbukinii [ljin
. Involucral bracts imbricate, unequal; outer bracts ovate or almost
triangular scene EGS hoe ae ee ae Be Rel Tester -. 9.
Involucral bracts almost equal; outer bracts lanceolate or ovate-
Lane eOlatS Hts. A a PAYS «Oooo oaciccncwncnncongnrccand (oan S .. 13.

. Leaves tomentose above along veins, bright green (plants of Altai)

Zach eM ets eer er conse hha i tf ems AN 35. S. jadrincevii Kryl.
Leaves not tomentose on upper surface along veins, either entirely
tomentose-pubescent, or glabrous (plants of eastern Siberia and

Rar; Bast)raseeen. ane ee eeinl as eee ek et ae 10.
. Capitula about 1.5—2 cm wide. Involucral bracts distinctly
acuminate, cusps often uncate ................ 36. S. tomentosa Kom.
Capitula about | cm wide. Involucral bracts somewhat obtuse, erect
DAU os. SERB GY piped Eerie A A te Denon) sabia. ie
. Leaves more or less pubescent beneath (plants of Far East)..... 12

Leaves glabrous beneath, green (plants of East Siberia; Sayans)
Se eB ah eee GY cl te ee 39. S. sajanensis Gudoschnikov

. Basal leaves distinctly cordate at base. Pubesence of leaf dense

beneath, white-tomentose (plants of Sikhote-Alin) .......................
obs Spree. Std. Belin. eee ete irs 37. S. porcellanea Lipsch.
Basal leaves cuneate or roundish at base. Pubescence of leaf not
dense beneath, grayish arachnoid-hairy (plants of Sikhote-Alin and
Sakhalin) .aineelan).. 38. S. kitamurana Miyabe and Tatewaki

. Leaves 2-10 mm wide, green, convolute, glabrous or somewhat

tomentose. Stem (4)10-30 cm long. Involucre, like stem, usually
dark colored.......... 11. S. schanginiana (Wydl.) Fisch. ex Herd.
Leaves 1.5—2 mm wide, grayish-pubescent or white-tomentose
above, white-tomentose beneath. Involucral, like stem, light colored
or entirely colorless. Prostrate plants, 2-10(12) cm long..............
cn sakeehee sundgenes el caters eebel phates daey 12. S. leucophylla Schrenk

14(3).

365 ar

16.

la

18(14).

+

19.

+

20.

359

Capitula about 1.5 cm wide, in racemose or corymbose-paniculate
inflorescence. Involucral bracts almost equal ............... eee 15:
Capitula smaller, in corymbs, sometimes if stem branched,
individual corymbs clustered in panicles. Involucre imbricate, in
several rows of bracts, gradually reduced toward periphery... 18.

. Receptacle covered with cartilaginous yellow papillae, sometimes

connategalmost,up, tOstip jeszeeess.-cietat. cinch. Letanhienh.. 16.
Receptacle covered with long, white, linear-subulate, stiff, lustrous
ET ERR a oR OE EES: RONG sete neey™ OR Teer CIM Pee Ns 17.

Stem up to 1 m long, about 1.5—2 cm wide. Capitula in dense,
wide, racemose, inflorescence, often half or one-third as long as
stem. Involucre densely woolly-tomentose. Veins on lower leaf
surface, especially at base, very broad, up to 1—-1.5 cm wide, straw-
yellows . os. as....earth.oe. ebelnsenal 4 9. S. dorogostaiskii Palibin
Stem 15-60 cm long, about 0.5—1 cm wide. Capitula, in less dense
and wide racemose inflorescence, usually much shorter than
halflength of stem, less often solitary. Involucre almost glabrous
or weakly pubescent. Veins on lower leaf surface, especially at
base, much narrower, up to 5 mM WIde............ cc eeeeeeeeeeeeeeeeeeereees
D, cxmeashsadunerd monet... Gb co nnaiice 8. S. baicalensis (Adams) Robinson
Inflorescence racemose, less often capitula solitary. Leaves narrow-
lanceolate, 1-1.2(1.5) cm wide, attenuate upward, sparsely toothed
(plants of Altai)... 10. S. krylovii Schisch. and Serg.
Inflorescence corymbose-paniculate, capitula less often less
numerous, sometimes solitary. Leaves broadly lanceolate, 2.5—7
cm wide, ovate or ovate-oblong, more closely toothed (plants of
Soviet Central Asia)...........0..ccee 13. S. sordida Kar. and Kir.
Receptacle glabrous, i.e., entirely without scales or less often with
numerous, sometimes shedding scales.............::cssccseeeeeeseeeeeeee’ Le
Receptacle more or less densely scaly; scales linear-subulate,
lustrous, persistent, of variable length in different species ..... 24.
Plants rosette type, almost without stem or with strongly reduced
SEEM... ance secseevsacuvanecesiea de teRheaaeteat sae. .uwiaranet fs emcee. motions tame ats rem 20.
Plants with distinctly stem, stems more or less tall ................ 21.
Leaves hard, orbicular, almost entire, with distinct short cusp.
Capitula few, up to 3. Outer involucral bracts strongly attenuate,
acuminate. Receptacle entirely glabrous (plants of the Darvaz
Range) setsrce. «ctenerevtdee «. 24de 105. S. caprifolia Iljin and Zapriagaev
Leaves herbaceous, soft, broadly lanceolate or ovate, with indistinct
cusp, shallow but distinctly crenate-toothed. Capitula usually
numerous. Outer involucral bracts weakly attenuate, acuminate.
Receptacle glabrous or with few short shedding scales (plants of
Pamiro-Alai, Dzhungarian Alatau).... 91. S. kuschakeviczii Wink.

360

21.

366

22

23)

24(18).

Involucral bracts blackish or violet; outer bracts equal or almost
equal or almost equal to inner, glochidiate with rusty woolly hairs.
Leaves fleshy, sparsely toothed or distinctly crenate-toothed.
Receptacle entirely glabrous (plants of Uda, Okhotsk, Anadyr, and
Kamchatka regions of the Flora of the USSR .........cccccccccees Jan

Involucral bracts distinctly unequal, i.e., outer much shorter than
inner, not glochidiate; if glochidiate, hairs white and not rusty ..
Siivdduaedneadsasdeou dived ea Cedi icb ste ee ALOE OLA ONE 22.
Leaves ovate, 2—2.5 cm wide. Outer iiivislubial bracts almost
herbaceous, acuminate, cusp often corniculate. Receptacle with very
fine scales (about 1 mm long) (plants of Alai) ............. ee
CEST INST coh aah ce tee ee tee ee es 92. S. ovata Benth.

Plants caespitose. Root collar densely covered with hardened
remains of previous year’s leaves, often fibrous. Stem prostrate or
ascending. Leaves lanceolate, toothed, less often pinnatipartite.
Receptacle variable: glabrous or scaly (plants of Pamir and central
even t Sivan), PERRI i Oe sizes 98. S. faminziniana Krassn.
Plants not caespitose. Root collar without numerous hardened
remains of previous year’s leaves. Stem erect. Leaves linear or
lanceolate-linear, entire, scarcely convolute. Bracteal leaves one
to three, approximate with inflorescence, usually exceeding it.
Receptacle glabrous (plants of Chukotka) ..............:cceeseeeseeeeees
GLE IO OE IR OI eee Se 104. S. tschuktschorum Lipsch.
At least inner (and partly also outer) involucral bracts with apical
membranous, usually colored, appendage. (Subgenus Theodorea
(Cass!) “Eipseht:)idisseiss oe eee eee eee steve ee enn coerce ener: 25.
Involucral bracts without membranous appendage or with narrow,
herbaceous, saber-shaped appendage .................:cecccssseeesseeeseees 36.

. Teeth on leaves and sometimes on outer involucral bracts with

cartilaginous cusp. Leaves pinnatifid ..............c:cccceseerereeeee 26.
Teeth on leaves and on involucral bracts without cartilaginous cusp.
Leaves in some species sometimes undivided.................:000+ Bul.

. Plants with distinctly winged stem due to decurrent leaves, or stem

WINeTESS) ECE: CORTE ORES SEs eee, eee eeeeeeee Diy.
Plants almost without stem, glaucous, strongly pubescent (plants
of Tuva Region) 22222022 See. 113. S. ceterachifolia Lipsch.

. Stem wingless. Leaves with narrow, linear or lanceolate lobes.

Outer involucral bracts attenuate, herbaceous, often as long as inner
OPIEVEMMLONGER ert et eee teak seen sete ates 111. S. runcinata DC.
Stem strongly or weakly winged. Outer involucral bracts distinctly
Shorter tham IMMer seecccoscceteesa ce aecosaczsoses tess tccanese-vetenennet eee reuecnadisuce 28.

367

30.

Sl:

2 Ja

a3"

34.

361

. Leaves pinnati- or bipinnatifid .......... ee eeeeesreeeeneeeeeeeeeeeees Z0:

Leaves entire or crenate-toothed, especially in lower part (plants
Of-Zaisan. Repion)<i04.).. 38.1. 208. 94. __S. paradoxa Lipsch.

. Capitula up to 1.5 cm wide. Leaves bipinnatifid, i.e., lobes deeply

and finely divided in small lobules with numerous, distinct,
cartilaginous cusps. Outer involucral bracts in several rows with
acute decurved cusp. Outer row of pappus bristles short-plumose
(plants of Kazakhstan Alltal) ........0.00000.. 112. S. laciniata Ldb.
Capitula up to 1 cm wide. Leaves pinnatifid, primary lobes only
toothed-crenate, not so deeply and finely divided as in previous
species, cartilaginous cusps poorly developed. Outer row of pappus
bristles serrate (plants of Trans-Baikal Region) ...................... 30.
Plants tall, more or less pubescent and scabrous. Stem 20—35(50)
cm high, strongly winged in type specimen. Involucral bracts
arachnoid-hairy or tomentose, glochidiate ....... 109. S. alata DC.
Plants shorter, almost glabrous and smooth. Stem 5—10(15) cm
high, somewhat winged. Involucral bracts almost glabrous. ..........

Pest IG OE Ee ate 110. S. crepidifolia Turcz.
Outer involucral bracts with subulate cusp, sometimes strongly
dee unvicd Bete Ot 0. BOLI RE BORIS. dale Sed. SD a2:
Outer involucral bracts without subulate cusp.................c000 33:
Capitula smaller (with florets 1.5-2 cm long). Stem prostrate or
ascending (erect only in var. erecta Winkl.), poorly branched,
scabrous due to dense, flexuous, soft acerate hairs, with few or
total absence of vegetative branches in axils of lower cauline leaves.
Leaves usually deeply pinnatifid. Outer involucral bracts weakly
subulately acuminate, not or slightly deflexed (plants of central
MSA Shani ee htc ON cc cnete 115. S. prostrata Wink.
Capitula larger (with florets 2.5—3 cm long). Stem always erect,
strongly branched, weakly scabrous, usually with larger number of
vegetative branches in axils of lower cauline leaves. Leaves in
type specimen undivided or with few teeth, less often pinnatilobate.
Outer and middle involucral bracts, at least in four rows, long,
acute-subulate, with strongly decurved cusp ............:::cccceseeeeeeees

enenentinnisa unin Merete yer et, ON BAT O OI, See 114. S. robusta Ldb.
Lower leaves ovate-oblong or lanceolate-oblong, coarsely crenate
or irregularly toothed, less often almost entire. Involucre turbinate
adschn host dudnbe dabehcenteestce, xccatl. Site caom eA eee 106. S. amara (L.) DC.
Lower leaves always pinnate. Involucre h: nispherical (plants of
caste Siberia and’ Par East) sf) noes. WAT en 8 a ee oe 34.
Plants tall, stems up to 1.5 m tall. Leaves large, individual leaves
reaching 25 cm in width, with prominent midrib. Involucre strongly
arachnoid-hairy (plants of Sakhalin and lower reaches of Amur)

matin st athein dannieirh datetiennann sn snr cotnerenneted Seaeae 108. S. neopulchella Lipsch.

368

362

+

35

+

36(24).

Si.

38.

30.

40.

41.

Plants shorter, stems 25—75(100) cm tall. Leaves less wide, with
less prominent midrib. Involucre weakly pubescent or glabrous
(plants of eastern Siberia and Far East)... ee eeeeeeeeeeeees 35%
All leaves hard, pinnate, with linear lanceolate-linear or oblong,
acuminate, up to 10 pairs of lateral lobes, scabrous above due to
spinules and sessile glandular hairs, with short pubescence and
numerous sessile glandular hairs beneath (plants of Trans-Baikal
andsalan: Bash) eisai if ee 107. S. pulchella Fisch.
Only lower leaves pinnate, middle and upper leaves undivided or
toothed, lanceolate or oval; lamina more SOft...............ccccccecceeeee
Pepieh arch ise. Se AUN a 107. S. pulchella var. subintegra Rel.
Semishrubs with numerous, strongly and repeatedly branched,
virgate, erect, more or less glandular-hairy stems. Leaves less
numerous, small (plants of Pamiro-Alal) .......0.........ceeseeeeeeeeeeeeeees
Bes acces PMN Roca. con conti ee Maan Pe obi: 100. S. chondrilloides Winkl.
Perennials, sometimes with woody bases, but then without virgate,
erect stems. Leaves usually numerous ...............cccccceesseeeeeseees 37h
Involucral bracts coriaceous, straw-yellow, with distinct, brownish,
rather wide border along margin. Receptacle with very long scales
(up to 2 cm long), scales as long or almost as long as inner
mVolucral: bractsys ayant ei pak Ret a 38.
Involucral bracts not coriaceous, without distinct, brownish wide
border along margin. Receptacle with much shorter scales; scales
shorter than inner involucral bracts ..............::eseseceeeeeeeeeeeereeeees 39.
Leaves bicolorous: green above, snow-white and appressed-
fOMENtOSE. beneath). 2.41.0) -.feee ese 101. S. masarica Lipsky.
Leaves concolorous on both sides: green, finely punctate-glandular
beneathipasyigs. ace et 102. S. tadshikorum [ljin and Gontsch.
Small (1.5—6 cm) alpine plants, densely leafy; upper leaves
surrounding dense, capitate, almost hemispherical inflorescence of
numerous sessile capitula. Bracteal leaves covered with brownish
or whitish pubescence. Involucral bracts of almost equal length.
(Subgenus Eriocoryne Wall. ex DC.) (plants of Soviet Central Asia
anidwAtltat) ioanincntiv, Re SOCIAL, segs Aa Vee a A 40.
Plants with different combination of characters ..............0..05 4}.
Leaves tomentose on both sides, grayish in type specimen.
Involucral bracts light colored. Pappus brownish-red ....................
Pilate Gt Okt tel ates 7. S. gnaphalodes (Royle) Sch. Bip.
Leaves velutinous above due to thick tomentum of white hairs,
less densely pubescent beneath, sometimes almost glabrous, reddish-
violet or green in type specimen. Involucral bracts reddish-violet.
Pappuspayw lite ents cvs desrsaresseeaseavaste oar otateraneo ye: 6. S. glacialis Herd.
Roots fibrously branched. Basal appendages of anthers with tuft of
longpiloceulase=woolllys Waits... osceeessrseroadoresseerorssounccasslecaestensser ds 42.

370

45.

46.

47.

48.

49.

363

Roots not fibrously branched (however, compared with S.
ussuriensis Maxim. and S. maximoviczii Herd.). Basal appendages
of anthers weakly or finely hairy, ciliate in some species,

flocculose-woolly only in S. maximoviczii Herd. .............000. 63.
. Involucral bracts with distinct long aristate cusp...........0..0... 43.
Involucral bracts somewhat obtuse or acuminate, in latter case
without distinct long aristate CUSDP .............ceeeeseeeesecesreeeeseeeeees 46.
Cusp of, mvolucral, bracts vuneate LON A ae 44.
Cusp of involucral bracts straight........ PEO OO ONS A 45.

. Stem tall (up to 80 cm high), strong, angular, more or less winged

due to decurrent leaves, not woody at base. Inflorescence
paniculate, strongly branched, with numerous capitula (plants of
Sémipalatii/ Reston) ee as ake ae 14. S. rigida Ldb.
Stem much shorter (20-40 cm high), cylindrical, not winged,
becoming woody at base. Inflorescence weakly branched, with
fewer (five to eight) capitula (plants of Dzhungarian Alatau)......
RIOR Sookie, Some worgusertemen neem VERE Ts 15. S. ninae Iljin
Corymbs erect. Pointed tips of involucral bracts sometimes fragile.
Florets in capitulum (10)12—13. Leaves white- or grayish-tomentose
beneath O}.9e> Gioia Oe eis Be Bees 31. S. larionovii Winkl.
Corymbs drooping. Pointed tips of involucral bracts not fragile.
Florets in capitulum 19-20. Leaves glabrous............... isc
AQ A AER Ua, 32. S. alberti Rgl. and Winkl.
Perennials; stem distinctly woody at base ........... eee 47.
Perennials; stem not woody or weakly woody at base............ 49.
Leaves 3—6 mm wide, oblong, crenate-toothed, some almost entire.
Capitula one to three (plants of Syr-Darya Karatau)...........00..0....
PTO MABE | GOMES CON BOE oe 18. S. mikeschini []jin
Leaves 1—2 cm wide. Capitula usually many (plants of Dzhungarian
ATE) WA LAME LE LA MAU AES EN TY, Ro oct ee a 48.
Leaves distinctly pinnatifid in type specimen. Capitula in lax
inflorescence, five to eight. Involucral bracts somewhat acuminate
ETE QAERGe | Wart AC OUISING 30 QMsarennt]. Siblanda*Schrenk
Leaves almost entire to weakly crenate-toothed. Capitula in compact
inflorescence, more or less numerous, up to 15. Involucral bracts
Strongly “acuminate .. 22 16. S. coronata Schrenk
Very tall plants, stem up to 1.5—2 m high, about 1 cm thick in
lower part. Inflorescence robust, paniculate. Leaves green beneath
BORO ARDS lh slice salud ducer tala tess ea oS 29. S. elata Ldb.
Plants short, stem up to | m high, usually much shorter; stem
GUTTA S, sR V IA VE 3 og PARIS 9 QB a OR BT, 50.

. Plants forming more or less dense turf. Leaves densely canescent

On bothy Gidesyceterrnt OI Eas HORT AR Sa Ea iia cemnerend Sit
Plants not forming dense turf. Leaves white-tomentose beneath or

364

53.

54.

55):

56.

371 =F

58.

less often grayish due to arachnoid hairy-tomentose pubescence,

sometimes, preenuon jboth sides)... °...) 2) .. agree eee 54.
. Leaves entire in type SPECIMEN .0.......... es cetccceeeseeecessnesseseeosenees 52.
Leaves distinctly pinnatifid or crenate-toothed................00 5D:

. Involucral bracts (especially outer) rather obtuse. Capitula small,

narrow-cylindrical, in dense erect corymbs. Leaves linear ...........
U sasvade cen canceansugcenenenetsettat cata Rey: eee bGE. REERe aes 22. S. cana Ldb.
Involucral bracts distinctly acuminate. Capitula large, broad,
campanulate, one to three, at stem apex, not forming erect corymbs.
Leaves narrow-lanceolate................6. 27. S. vvedenskyi Lipsch.
Involucral bracts light colored, weakly pubescent. Stem in typical
specimen 1.5—-4 cm high, in var. major Winkl. up to 10-12 cm.
Capitula one to three on each stem (plants of Kuldzha District—
Sairam, Kookamyn)::.../ dace apes ee eee 26. S. canescens Winkl.
Involucral bracts dark, densely arachnoid-hairy. Capitula three to
five on each stem, number of capitula on entire plant large (plants

GieGentralPPienm Siam). es... wees eee 25. S. caespitans Iljin
Leaves concolorous, i.e. green on both sides. Stem somewhat woody
Ab DASE fassesee. -dee «=a derises cote hati des fede le Sok Ok ceRP epee phe ane Do.

Leaves green above, white or grayish beneath due to tomentose or
arachnoid hairy-tomentose pubescence. Stem not woody at base
vemos teesadenasausccsaea geep@ehneees «Ques Raa I OR AEN: seeteceetsb chiens gaa ap esemenn.. 36.
Leaves in type specimens lanceolate, acuminate, undivided or finely
toothed-crenate, weakly scabrous. Capitula smaller and narrower,
5—7 mm wide. Involucre dark colored. Pappus dirty-smoky (plants
of, western, Dien,shan)) oiinses..-celto 19. S. ispajensis [jin
Leaves in type specimen ovate, basal and lower cauline leaves
obtuse, upper cauline leaves acuminate, usually more deeply
toothed-crenate, strongly scabrous. Capitula somewhat larger and
wider, 7-10 mm wide. Involucre white. Pappus white (plants of
Dzhungarian Alatau (Chulak) and eastern Tien Shan) ..................
sosvasn cages iapeeeeere: pagieaeh - Haber Meee 28. S. salemanni Wink.

vice tok cabagtaceeae ). «cut ltaawo te Renate. So ena See, eae: ential ene es SIs
All leaves undivided or crenate-toothed ..................:c:seseeeeeeeeeees 59.

. Plants up to 60 cm high; stem strongly branched above. Leaves up

to 14 cm long and 4 cm wide, pinnatipartite into lanceolate or
linear, 1.5-4 cm long and 2-5 mm wide lobes. Involucral bracts
almostgelabrous i512. ee 24. S. serratuloides Turcz.
Plants shorter, up to 10-15(25) cm high, with weakly branched
stem. Leaves smaller, variously divided; pinnatipartite,
pinnatilobate, or crenate-toothed with shorter lobes and 1-9 mm
long teeth. Involucral bracts pubescent ................cccceceeeeereeeee 58.
Capitula in somewhat lax corymbs, not erect. Leaves strongly

59:

60.

61.

62.

63(41).

+

64.

365

unequally divided; pinnatipartite or sinuate-toothed, only some

entire (plants of Siberia)... 23. S. sumneviczii Serg.
Capitula in compact corymbs, erect. Leaves pinnatifid (plants of
Kazakhistan) tennis. .Mumety uel ltr taunt. ily 22. S. cana Ldb.

Basal and lower cauline leaves narrow, usually 2-8 mm wide (not
wider than 1 cm), entire, bright green above and white-tomentose

beneath (plants of Siberia) .................. 20. S. salicifolia (L.) DC.
Basal leaves often withering before flowering; lower cauline leaves
waders 5-2(3). cmywide).ecc.: hole. contona: te shone) .. 60.

Plants usually with large number of short vegetative shoots in axils
of cauline leaves. Stem densely leafy. Leaves strongly scabrous
above and along margin, gray-pubescent beneath, sometimes very
insignificantly, or white-tomentose. Capitula 5-8 mm wide, in wide
corymbose-paniculate inflorescence ............c.ccccccssseseeeeseeeseeeeees 61.
Plants without short vegetative shoots in axils of cauline leaves,
or vegetative shoots scarcely noticeable ...............ccccecceseeeees 62.
Leaves more or less gray-pubescent beneath, sometimes very
weaKdya pubescent, ttc. .etasals.. aun. quiet 30. S. elegans Ldb.
Leaves distinctly white-tomentose beneath ..............ccccceeeeeeeees
Riur..2ohosied, LePage 30. S. elegans Ldb. var. nivea Lipsch.
Capitula small, 5—8 mm wide; involucral bracts acuminate, without
black border. Inflorescence in type specimen depauperate, terminal,
usually compact, corymb, not broadly corymbose-panicle; less often
several lateral corymbs besides terminal. Stem sparsely leafy .....
eeabe bees cleo, ioe... 30. S. elegans ssp. turkestanica Lipsch.
Capitula rather large, 1 cm wide, with obtuse, roundish at apex
involucral bracts, with black border (plants of Tien Shan, Kunges,

and outside the USSR) ...........ce 21. S. lomatolepis Lipsch.
Leaves white or gray beneath due to distinct tomentum or
arachnoid-hairyopubescericenen..£.78...cist. ia DES BON... 64.

Leaves green or glaucescent beneath, glabrous or sparsely
pubescent, sometimes indistinctly arachnoid hairy-pubescent, and

PHEM, {BRAVES oes ccnnccecesncseancencernrvbactonroacveA Ret: AUREL ee 82.
Lower leaves in type specimen deeply cordate or hastate, and wide
at base\..bes. .aueey.. th. gutele) aaretidiy a dul ier bay BNE) | 65.
Lower leaves cuneate and narrow at base ...............ccccccseeeeees 74.
. Leaves on both sides, as also stem and involucre white-tomentose
oF sitongly;pubescentiui.s.c 04 -.420.. A1 ek .. Se. 66.
Leaves green above, white-tomentose beneath. Stem glabrous or
WW CAL OPUI DES CERHMGEME Be) Bene cnson sachatctonseonencnimnecamreccesuitveeeer ts 67.

. All plants snow-white due to dense tomentose pubescence. Capitula

ovoid, larger; involucres up to 12 mm long; involucral bracts
densely white-tomentose, short spinescent. Florets pink (plants of
FaneBast)its..8........28 Ba cbsaarstidttn tn Gh eat earls 40. S. sovietica Kom.

373

366

67.

68.

69.

70.

7

TQ

Plants more weakly pubescent, leaves arachnoid hairy-woolly above
and not tomentose, weakly crenate-sharp toothed. Capitula smaller;
involucres 7-8 mm long; involucral bracts not densely white-
tomentose, with long, usually strongly squarrose, cusps. Florets
violet (plants of eastern Sayans)............ 41. S. squarrosa Turcz.
Upper cauline leaves approximate with inflorescence. Leaves
strongly crenate-sharp toothed. Stem often flattened, thick.
Involucral bracts with strongly squarrose cusp in type specimen
(plants\/oF ‘eastern /Sayans) es. .scc..--5 chee sods user teen fe ces ecns
Le aa LEER Pee eee boot 42. S. pseudosquarrosa M. Pop. and Lipsch.
Upper cauline leaves not approximate with inflorescence. Stem
never flattened. Involucral bracts without squarrose cusp, if cusp
present, then*strarplitqe See OP es, cree oe UR REO IS 0 68.
Stem very tall (up to 1 m or higher). Lower cauline leaves long-
petiolate, rather large and wide, up to 16 cm long excluding petiole
and up to 10 cm wide. Petiole of middle and lower cauline leaves
often winged. Inflorescence-robust panicle. Peduncles brownish due
to dense glandular hairs. Florets pink (plants of Far East—
Birobidzhan)).::2xtiawad: ake aon 49. S. splendida Kom.
Stem less tall. Lower leaves medium. Petioles wingless.
Inflorescence less robust. Peduncles gray arachnoid-hairy pubescent,
not clandularshairyseigh et ie ROR SNE RS... 69.
Lower leaves ovate, oblong-ovate or oblong-lanceolate (2.5 to 8
cm wide), deeply cordate at base in type specimen................ 70:
Lower leaves narrower, 1-2 cm wide, not deeply cordate at base
b: Tao, ee ie omar ere 1 One MER Ee, « Ty,
Stem 10—20(15) cm high, ascending. Inflorescence usually compact,
with small number of capitula (plants of the Carpathians)...........
EEE | TORI OE PE Boe 3 44. S. discolor (Willd.) DC.
Stem 25-80 cm high, erect from base. Inflorescence usually lax,
with larger number of capitula (plants distributed from Ural to
Mongolia)sind. enndoces ctoaiteiban: ns 43. S. controversa DC.
Beavestdeeplystoothicd le .).-.cw.ve-cten-cosucectcncunstrpmee eee Meee aataea ns (2
Leaves scarcely toothed, gradually acuminate. Involucral bracts
obtuse or somewhat obtuse (plants of Yakutia and Ayan District)
eS iconcctwnewtcccece ORO acc MELDNIOSET URE, EOE ce, MOTUS. Tad Mena hac 73.
Leaves lanceolate-linear or oblong-lanceolate, abruptly long
acuminate. Capitula on long peduncles, somewhat remote.
Involucral bracts acuminate (plants of Irkutsk Region).................
yas Caen aad Malia ara up Or 47. S. lenensis M. Pop. ex Lipsch.
Leaves ovate or ovate-oblong, long acuminate, somewhat hastate
or roundish-obtuse at base. Capitula in compact inflorescence, less
often solitary. Involucral bracts obtuse or somewhat obtuse (plants
OLADRAR)S STI. csoresealersdoadinnaradsataeseades 48. §. uralensis Lipsch.

74(64).

374

LB:

367

Plants 10—15(20) cm high. Capitula usually solitary, sometimes
two or three on short peduncles in compact corymbs (plants of
Avan District) vcd 23.3... tity 46. S. ajanensis (Rgl.) Lipsch.

+ Plants taller, stem 25—40 cm high. Capitula two to five, in corymbs;

7.

76.

ills

78.

19.

usually stem more or less’ strongly branched in upper part, and
then, besides terminal corymb, lateral capitula present at different
heights on long floriferous branches (plants of Yakutia) ..............
LcaseeT crae) arate Sere tay 45. S. hypargyrea Lipsch. and Vved.
Involucral bracts narrowed from broader base to distinct, usually
incurved cusp or small, herbaceous, also incurved, appendage ....
OME OES OTE ROU OMIA) at ihe, OU eet 75.
Involucral bracts without incurved cusp or appendage............ ite
Stem wingless. Capitula larger and wider, 1—-1.4 cm long and 1
cm wide. Involcural bracts usually dark-colored, with long cusp,
TAI Nye COUT VIC GI Ui Ruins an Natta ake Saute rans Ue apts ir poo. 76.
Stem more or less winged. Capitula 8 mm long and about 7 mm
wide. Involucral bracts light-colored with small, herbaceous, usually
violet, recurved appendage (plants of Trans-Baikal, Yakutia and
| PIE Ge] SES Paar en hy eh a oe A ae te 62. S. duvia Freyn.
Stem up to 15 cm high. Lower leaves 1.2-1.5 cm wide. Cusps of
involucral bracts less developed, straight. Pappus slightly exserted
from involucre (plants of Sikhote-Alin) .......0...ccccccesesscseeseeeeees
sh oe es en eel el Btls itt a a Pe 79. S. socezavae Lipsch.
Stem more than 15 cm high. Lower leaves up to 5-6 cm wide.
Cusps of involucral bracts long, usually recurved. Pappus strongly
exserted from involucre (plants of Kamchatka and Commander

ESLAMGS ere sn eer eee tne ee eee 80. S. pseudo-tilesii Lipsch.
Stem tall, usually more than 35 cm high ...............cccceeeeeeees 78.
Stem short or medium, usually less than 35 cm high............. 12.

Outer involucral bracts usually turning black; entire involucre
(especially inner bracts) in type specimen velutinous-woolly. Leaves
hard, white-tomentose beneath, somewhat convolute. Florets pink
Bae at Sine ti eae edie duces Caen Nedieatinge 59. S. amurensis Turcz.
Outer involucral bracts usually not turning black; entire involucre
in type specimen pubescent but not velutinous-woolly. Leaves
softer, gray- or grayish-arachnoid hairy-tomentose beneath, not
convolute: Flonets,deepried Glways 7). 2. ...c.-dcccacecadceanins sohocurneestoves
Reet aa Nee aa Med nad ena neck atncsok 85. S. stubendorffii Herd.
Stem densely leafy up to inflorescence. Outermost involucral bracts
lanceolate, as long as or somewhat shorter than innet............ 80.
Stem sparsely leafy up to inflorescence. Outer most involucral
bracts distinctly shorter than inner, often hardening later, somewhat
acuminate or somewhat obtuse. Capitula mostly on long peduncles,

368

80.

81.

82(63)

83.

usually in somewhat lax terminal corymb (plants of eastern Siberia)
LEAL: EL Laas menheganns tes 6 83. S. sukaczevii Lipsch.
Plants almost stem-less or with 1-10 cm long, less often up to 15
cm stem. Upper leaves lanceolate or lanceolate-linear. Basal leaves
usually not withering before flowering and often reaching up to
inflorescence? onkexcee dinghies ke teh elle petal 81.

Plants with 15-30 cm long stem. Stem leafy up to top. All leaves
oblong-ovate or elliptical, with frequent sharp teeth. Basal leaves
usually withering before flowering (plants of Altai and Saur) .....

gsernepiee~. hancsanye) ee 74. S. foliosa Ldb. var. altaica Turcz.
Capitula in dense capitate corymbs, often subtended by one to
three upper cauline leaves, either longer or as long as capitulum.
Outer involucral bracts shorter than inner. Receptacular bristles up
to 6 mm long. Florets 1—1.4 cm long (plants of the Siberian Arctic)
SEEING. IGN A ar Os BT MR AO Tae Le ge Dili 78. S. tilesii Ldb.
Capitula in dense, but not capitate corymbs, not subtended by upper
cauline leaves. Outer involucral bracts almost as long as innermost
or slightly shorter; all involucral bracts often dark-colored (red-
violet). Receptacular bristles 2-3 mm long. Florets about 8 mm
long (plants of Altai and Sayans).............. 75. S. pricei Simpson
Leaves distinctly punctate-glandular beneath, glandular hairs not
lustrous. Outer involucral bracts as long as or almost as long as
inner. All involucral bracts narrowed from wider base to long
Subtulates Cuspy ia. c hates 7s Deere take See mr eee ae 83.
Leaves not distinctly punctate-glandular beneath, if glandular hairs
present, then scattered, lustrous. Involucre imbricate, consisting of
several rows of unequal bracts: outer short, inner longer. Involucral
bracts without appendage or with narrow saber-shaped appendage
se dul as jetta ton en nae ica idijcas ble aet ma nioe tae ee ON 84.
Alpine plants, small, often almost stemless, stem 2—8 cm high.
Leaves thicker, hard, oblong-lanceolate, 0.5—1 cm wide, shallow
CHEM AEC = FOOURE Gone ca araecmeccee soo cee aes 76. S. congesta Turcz.
Plants of valleys, larger; stem 10-20 cm high. Leaves thin, ovate
or ovate-triangular, 2-5.5 cm wide, deeply crenate-toothed with
sharper teeth (plants of the Vitim Plateau) .............eeeeeeeeeeeeeeeees

seadscevengsiadi.Caaesaer ote aa dca eee ae ence 77. S. poljakovii Glehn

. Involucral bracts with saber-shaped, straight or decurved appendage

SUTURE CRIA ER” S50), DSIRE oe een ee Sere 85.
Involucral bracts without saber-shaped appendage, obtuse or
ACUMUMALS 5 .s.c-ceccseresantegeseonns-ctesneettue ROR ERS. eaten cee eee a2
} eaves entire or toothediine.....ise Scat ween: eee eet eee 86.
Leaves deeply pinnatisect or pinnatilobate «2.0.00... eee 90.

. Middle cauline leaves decurrent on stem, as a result, stem more or

VESSU WINERY. © cee scortncnntebectecncheneecsessstetts socec tees ee eee eae ee 87.

376

87.

88.

89.

Oi,

369

Stem winless yrshec snares eee actus sap ccebonsucelaane caauliatees beegt 3 89.
Capitula in compact terminal corymbs, 5—7 mm wide. Stem narrow-
Aerie ell. hee heey ts baler greta dees oct, Paatrssiesie's, . eases eR 88.
Capitula in racemose panicle or lax equally long corymb, 10—14
mm wide. Stem with broader toothed or entire wing. Saber-shaped
apical appendages of outer and middle involucral bracts dark red,
more or less recurved (plants of Trans-Baikal Region and Far East)
Le RR BRE TCMIG OR SERS, ROTA 63. S. umbrosa Kom.
Leaves aeruginous, oblong, glabrous, usually glossy on both sides,
entire or weakly toothed; basal leaves narrower (about 2 cm wide),
petiolate. Saber-shaped apical appendages of involucral bracts
always distinctly developed, long (plants of eastern Siberia) .......
Pea RR ER AIS Uae aE 57. S. acuminata Turcz.
Leaves green, oblong-lanceolate, or ovate-oblong, glabrous above,
sparsely arachnoid hairy-pubescent beneath, distinctly toothed-
crenate, teeth acute; basal leaves 3-4.5 cm wide, with triangular-
hastate (sometimes almost cordate) base, narrowed into petiole.
Saber-shaped apical appendages of involucral bracts sometimes
poorly developed and short (plants of Sakhalin) «0.0.0.
see Rane MESS ea tReet) Vite ckcer eomam Ret 58. S. sachalinensis F. Schm.
Capitula together with florets about 2 cm long and up to | cm
wide, usually numerous (up to 15), less often only three, clustered
at stem apex. Involucral bracts glabrous, dark-colored in type
specimen, with thin recurved apical appendage, less often without,
but then apices recurved and appressed. Leaves thick, hard, oblong-
(0) Sta ae ante ene iene erie, Act 51. S. recurvata (Maxim.) Lipsch.
Capitula one to three, less often more. Involucral bracts (especially
outer) velutinous-woolly, less often subglabrous, with thin, up to
1-1.25 cm long recurved apical appendage. Leaves thin,
OVdle-thlanewlar OF AMMOSE taNSUlar .1...2 nce. tse. -o--cnseqcscsseesecaresuae ce
Sigg SASH CSU Nat alte erate Repeater voe i 65. S. subtriangulata Kom.

. Primary lobes of leaves usually more than 10 pairs, narrow, long,

large toothed. Upper cauline leaves oblong, cristate-pinnate.
Involucral bracts with herbaceous, cristate-toothed, erect or recurved
ES) OLN Sp Raga Na 34. S. odontolepis Sch. Bip.
Primary lobes of leaves less than 10 pairs, wide. Lower leaves
pinntilobate, upper cauline leaves oval or lanceolate. Involucral
bracts with undivided appendages. Capitula at least 1.2-1.5 cm
| OYSTERS ANE Stee a Nt ey RI EE OEE 91.
Peduncles slender, somewhat drooping. Leaves thin, oval-oblong,
with wide, acute lobes. Involucral bracts puberulent or weakly
arachnoid-hairy, green, with saber-shaped, 5-8 mm long, recurved
Of, Straight AP MENGAGES a onc on case anes nennspeecduads 66. S. sinuata Kom.

370

92(84)

377

95.

96.

ae

98.

Peduncles thick, short, straight. Leaves, ovate-oblong, cuspidate,
deeply pinnatisect. Involucral bracts glabrous, usually dark-colored,
with or without recurved appendages, but then tip of involucral
bractsyrecunved {iene a8 51. S. reeurvata (Maxim.) Lipsch.
Involucre glabrous; outer bracts like true floral bracts herbaceous,
green; subsequent bracts green, ovate at base, attenuate-long
acuminate; inner bracts oblong or lanceolate, red in upper part,
long acuminate, exserted from outer bracts. Leaves fleshy, glabrous,
quite variable in shape, size and especially in serration. Capitula
two to seven on each stem, solitary or less often three, sessile, at
apices of stem and lateral branches, protruding from axils of upper
leaves (plants of Kara-Archa area in central Tien Shan)..............
pe a ebereea her, See se th LES EM 99. S. karaartscha Saposhn.
Involucre glabrous or pubescent; outer involucral bracts not like

true bracts. Plants with different characters..............:cccscere 22),
. All or at least lower cauline leaves pinnatisect or pinnatilobate..
baw adllpe b= We drekvavioecita’ ts Sommdieh Pg eaeeeeecs ace tee eae etre aire, ie ge 94.
All eavesundividedyor*toothed?.... 505 tance: fees eee 103
Plants of saline lands, slightly bitter to taste. Leaves fleshy ........
flee SAR ROL Pee ae bee i ena Mbt ok ater errs PPR EnY qd bie aman d A, 95.
Plants not from saline lands but other habitats. Leaves not fleshy
Aes aepabitia nut Pace AUR lee A bgt. Ase tebe: ride Mb ce yA Goilhe Meinl da 99"

Stem prostrate or ascending, caespitose. Leaves narrow (2-5 mm
wide) pinnatifid, with few or more triangular or linear lateral lobes,
sometimes leaves almost undivided. Capitula usually drooping,
sometimes in secund inflorescence (plants of Pamir and Tien Shan)
eR des Nie SEALING ANB 98. S. faminziniana Krassn.
Stem straight and not prostrate or ascending, usually solitary.

Capitularercet sand “notidroopine seis eae neeeasee eeeeee eeeeen oe 96.
All leaves deeply pinnatisect, usually bipinnatisect, then secondary
lobes Nineata en. ene teeta tee 96. S. turgaiensis B. Fedtsch.
Leaves” mott Bip ininatisectr sss earrou ees Cee ne ea ee nee Ths

Basal leaves lyrate-pinnatisect, with large, almost sagittate, and
crenate-large toothed terminal lobe; middle and ypper cauline leaves
almost undivided or toothed-lobed. Capitula small. Inflorescence
branched in type specimen. Involucral bracts pale. Outer pappus
Dristles* not Persistent sre ccc, eee ek ee cae Sea ee wens meee ee 98.
Basal and cauline leaves crenate to toothed-lobed. Capitula at
fruiting 1.6-2 cm long and 1—1.2 cm wide, in dense corymbs at
stem apex. Inflorescence unbranched or weakly branched. Involucral
bracts deeply colored (red-violet). Outer pappus bristles persistent.
Leaves and peduncles strongly scabrous (plants of Tien Shan,
GUESTS UME OS SR ire se tee eerente cert 97. S. kaschgarica Rupr.
Stem 15-50 cm high. Leaves decurrent, as a result stem sometimes

378 ty

103(93)

104.

105.

37]

winged. Capitula numerous, lax paniculate, of many corymbs.
Achenes without border (corona)....... 93. S. salsa (Pall.) Spreng.
Stem 4—10(15) cm long. Entire plant usually more glaucous. Leaves
not decurrent on stem. Capitula terminal on main stem and
branches, in dense, often globose corymbs of 4-10 capitula.
Achenes with narrow border (corona).... 95. S. dehurica Adams

- Capitulayat least, 1.2—1.5.cm long dante aeons 100.

Capitula not more than 1 cm ong ........... ccc eeeeeeeeseeeseeeeeees 101.

. Peduncles thick, short. Leaves ovate-oblong, cuspidate, deeply

pinnatisect. Involucral bracts glabrous, usually dark, not recurved
igdeaapen «delastieae! nmt 50. S. elongata DC. var. polypodiifolia DC.
Involucral bracts recurved .... 51. S. recurvata (Maxim.) Lipsch.

. Basal leaves usually large and wide, lyrate-pinnatisect, with

prominent terminal lobe, generally appressed to stem; lobes
lanceolate or oblong-lanceolate, cuspidate, weakly cuneate-decurrent
at transition to petiole. Involucral bracts ciliate or weakly tomentose
along margin. Basal appendages of anthers comose......................
BS aca ct yates tp neana ce ceumeebaabemesen 53. S. maximoviczii Herd.
Basal leaves oblong or ovate, cordate or hastate at base, without
prominent terminal (upper) lobe; lateral lobes broadly oblong,
terminating in wide somewhat obtuse teeth. Involucral bracts
arachnoid-hairy. Basal appendages of anthers ciliate ............ 102.

. Leaves glabrous beneath, weakly scabrous or smooth...................

ial saat, Nas Sete sh te 52. S. ussuriensis Maxim. var. incisa Maxim.
Leaves arachnoid hairy-pubescent beneath, harder, strongly scabrous
Se eer sete ba 52. S. ussuriensis Maxim. var. firmifolia Lipsch.
Capitula cupuliform, peduncles up to 2 cm long and 1 cm wide,
3=15,,tatpstem, apices: (plants,of Kar East) tj. 2t.21eletee tian) 8.28
Site oe tpee, «Pui viata: seeded, Naps ieee N elem sad 64. S. grandifolia Maxim.
Capitula smaller, narrowly or broadly cylindrical, infundibuliform
or campanulate ...... gay yes perieegre cael easels bas weat. Loeb etl, 104.
Leaves with frequent and sharp teeth, green on both sides, glabrous
beneath or sparsely covered with short, somewhat stiff hairs; middle
cauline leaves broadly ovate, ovate or elliptical .................... 105.
Leaves grayish beneath due to arachnoid-hairy pubescence,
sometimes green and glabrous (but then different in shape than
reported above); middle cauline leaves lanceolate, ovate-lanceolate,
triangular or triangular-ovate, hastate, undivided or weakly,
Sometimes! sstronglysitoothied i.e... weer ei ee MO, 106.
Leaves strongly decurrent, as a result stem distinctly winged.
Involucre often glossy; involucral bracts almost glabrous in type
specimen, weakly pubescent only along margin and apex; outer
bracts ovate, shortest. Large, 40-120 cm high forest plants.........
NOS ERLE oe cctnte Ms. Meat incey oe racrl nea iceuic waste vas nesliae 73. S. latifolia Ldb.

37/8)

372

108.

109.

2.

Leaves weakly decurrent on stem. Involucre not glossy; involucral
bracts usually pubescent; outermost bracts lanceolate, longer than
subsequent ovate bracts; inner bracts densely covered with long,
somewhat stiff, appressed hairs. Short, 15—35 cm high, alpine and
subalpine plants .............. 74. S. foliosa Ldb. var. glabrata Kryl.

. Middle leaves decurrent, as a result stem winged in type specimen;

wings unequal in width 0.00.00... eeeeeeeeneeeeeeee An Be 107.
Middle leaves not decurrent or very weakly decurrent, as a result
stem’ winglessiin' type*specimen? 2.2 2A eae 114.

. Stem in upper part and subtending leaves, when present, glandular-

hairy. Inflorescence of numerous capitula. Capitula up to 1 cm

_ wide (plants of Sakhalin) .......... ITE RR,

LARA RENOIR IN Tu 54. S. nupuripoensis Miyabe and Miyake
Glandular pubescence absent. Inflorescence usually comprising
numerous capitula. Capitula 5-6 mm wide .............. eee 108.
Leaves hard, 3.5—8 cm wide, strongly attenuate. Involucral bracts
green, in type specimen, only sometimes pigmented at apex. Pappus
weakly "exserted: from’ ‘Ca pituliin vsscccerccesesevscoreeeeesz eect 109.
Leaves fleshy, 1.5—3.5 cm wide, somewhat attenuate-acuminate.
Involucral bracts usually dark-colored, less often green. Pappus
distinctly Texserted: from’ capitulunite 2202s... RAR 110.
Stem 50-100 cm long. Leaves glaucous or somewhat glaucescent
beneath, glabrous, strongly toothed, distinctly cuneate at base,
narrowed into rather long winged-petiole (plants of eastern Trans-
Baikal Region and continental part of Far East) ..0..... eee
bc EE SSSR PER 2 Ws PE NS ER A 71. S. neoserrata Nakai
Stem 150-200 cm long. Leaves grayish-pubescent beneath,
somewhat toothed, with roundish base, narrowed in short winged-
petiole (plants of southern Kuril Islands) «00.0.0... eee eee
FEN SRE, EE 72. S. fauriei Franch.

. Involucral bracts glarbous, sometimes weakly pubescent only along

Tarp. EA I ls, OR UN BBD 111.
Involucral¢bractsspubescentieiey. Boa! le eee ee 12

. Leaves and stem green. Stem narrow-winged in type specimen.

Leaves small toothed, not convolute (plants distributed from Ural
to Trans-Baikal Region) ................. 69. S. parviflora (Poir.) DC.
Leaves and stem dark colored, purple. Stem very weakly winged
in type specimen. Leaves almost entire, somewhat convolute (plants
of :-Trans-Baikal«Region)e..ie a eae 5 oases
te 69. S. parviflora subsp. purpurata (Fisch. ex Herd.) Lipsch.
Leaves more or less pubescent beneath, with glossy, more numerous
or less numerous glandular hairs. Stem narrow winged, sometimes
Wihplessae LA Se. Ba Loh. PRR tee ee 113.

+

AL 3s

BTS.

Leaves entire, glabrous, glaucescent beneath, eglandular. Stem
broad-winged. Involucral bracts somewhat obtuse, velutinous or
arachnoid-hairy dorsally (plants of the Carpathians) .....................
pte het ceeds eminent, Sn taal illo Reeth 2d 70. S. porcii Degen
Stem weakly leafy. Leaves sparse, up to 1, less often 1.5 cm wide.
Involucral bracts, especially outer, acute, pubescent, with narrow
violetphonder sth. 2. heise. cadens nds 61. S. shiretokoensis Sugawara
Stem strongly leafy. Leaves usually densely arranged, up to 2 cm
wide. Involucral bracts strongly arachnoid hairy-pubescent,
glochidiate, without narrow violet border... cee eeeee esses
jetta. car anise, botaeds wmoticeeel silat 60. S. duiensis Fr. Schm.

114(106) Stem and leaves purple. Stem very weakly winged or wingless.

LS)

Leaves almost entire, weakly convolute ..............ccscecssceeeseeeeeeeees
. 69. S. parviflora subsp. purpurata (Fisch. ex Herd.) Lipsch.
Stemandyleavespereceniewtneheuslomereilecrs PIES. 115.

. Lower leaves long-petiolate, wide, triangular, ovate-triangular or

hastate, large toothed, cordate, emarginate or distinctly cuneate at

baSeenevwdie Oak t blac, nhs, dodem cad cea maces abba) 116.
Leaves of other shape, often, but not always, lanceolate, narrow;
not cordate nor distinctly cuneate at base .............ceeeeeeeeees 121.
. Lower leaves ovate or ovate-triangular. Upper cauline leaves
approximate with and usually exceeding inflorescence ......... olive
Lower leaves cuneate. Upper cauline leaves not approximate with
and not exceeding inflorescence ...........cccccscssscessseseseeeeeeeeeeeeees 119.

. Stem in upper part, cauline and subtending leaves beneath, as well

as petiole flocculose-arachnoid hairy-woolly (plants of Kuril Islands)
wa retese sd ecenss RRO: EE Oe ores. De, cost 55. S. kurilensis Tatevaki
Stem, cauline and bracteal leaves beneath, as well as petiole not
arachnoid hairy-woolly pubescent ........0.....eeseesceeeseeeesseeeeees 118.
Leaves thin. Inflorescence of numerous capitula. Capitula narrow
(about 5-6 mm wide). Stem pubescent in uppermost part with
stiff, not glandular hairs (plants of Kamchatka, Kuril Islands, and
SAMI) acca suacsvndetanenaaeiecarest MRR RUM 56. S. riederi Herd.
Leaves thick. Inflorescence of less numerous capitula. Capitula
more than 1 cm wide. Stem sometimes weakly winged due to
decurrent leaves in uppermost part, and bracteal leaves, especially
along margin, glandular-hairy (plants of Sakhalin).........000.00000..
i (deed Rabnal ele \ethheat 54. S. nupuripoensis Miyabe and Miyake
Leaves, even uppermost cauline, oval or broadly lanceolate, toothed.
Involucre’ arachnoid: hairy-puibescent 20.0... 2 okie ekicehidee ie
BA Ce, ean eal Bis ba eg) 52. S. ussuriensis var. genuina Maxim.
Upper leaves linear-lanceolate, narrow, entire. Involucre glabrous ~
ORSUbSABKOMS ARTE ARC Rs Te EAS a aE 120.

374

WA) Bs

122.

123)

+

381 124.

+

125.

126.

. Involucre broader and shorter, of three or four rows. Inflorescence

comprising fewer (4—15) capitula in type specimen, compact......

silane anne RAR ee ENS | BaD 67. S. triangulata Trautv. and Mey.
Involucre narrow and oblong, of five to seven rows. Inflorescence
lax, long paniculate, usually comprising numerous narrow capitula;
floriferous branches in type specimen bent ................ccccccssceeeseeeees

2 Sas RAR TNR MEIN 2 ENN a ccaitscimwgennenaien 68. S. manshurica Kom.
Outer involucral bracts wide, ovate, ovate-lanceolate or oblong-
ovate, somewhat obtuse; inner bracts strongly pilose. Leaves
arachnoid hairy-pubescent, grayish beneath, less often glabrous...

Laceeha papers AoA. ood systineneneney cau te ier casks cena cade. 122.
Outer involucral bracts less numerous, narrow, long acuminate,
almost as long as subsequent bracts; inner bracts glabrous or
subglabrous, white-villous only at tip. Leaves entirely glabrous on
lower surface; lower cauline leaves lanceolate or ovate-lanceolate,
irregularly toothed, teeth sharp (plants of Kamchatka) .................

ih Resid EUAERTERURR eta, SSRN EDI, SRR IRI IR 81. S. oxyodonta Hult.
Capitula numerous, large and wide, (1)1.5—2 cm wide; central
capitulum of corymb usually larger than laterals...

dscsppeduiereead ss Begete pte, QR i ER 90. S. pseudoangustifolia Lipsch.
Capitula numerous in type specimen, narrower; central capitulum
ofi,corymb)not)largenthan lateralsiis.8. ee ee 123%
Leaves (especially along margin), as well as partly stem (in upper
part) glandular due to many-celled glandular hairs. Involucre of

two or three rows (plants of Chukotka)........ 89. S. viscida Hult.
Leaves as well as stem not glandular. If glandular, then glands
sessile, lustrous. Involucre of three to five rows ..................+ 124.
Plants ewoody. .A%...2) ee wpe Ea RARER Ee OER 125):
Plants), herbaceous |..g.20h..cd ROI RE OOO ERG - OE Ee. 126.

Caudices absent. Leaves linear, narrow, 2-6 mm wide, entire,
usually convolute, glabrous on both sides or sparsely flocculose
beneath. Receptacular bristles up to 5-6 mm long (plants of
thestislands, offithe Bering Strait) :....3...c.scececeoeetecnvee Re OE
ssh eiCeeh ah . mae tt ier cattle Bae, 88. S. angustifolia (Willd.) DC.
Caudices numerous, densely covered with hardened remains of
previous year’s leaves. Leaves broad (1.5—2 cm wide); lower leaves
oblong, ovate or ovate-oblong, arachnoid hairy-pubescent on both
sides (densely beneath). Receptacular bristles shorter, up to 2-3
mm long (plants of Tadzhikistan-Zeravshan River Basin) ............
wi sencwtihhn Stee pn een RE Eee 87. S. saxosa Lipsch.
Capitula in terminal, more or less compact corymbs, at apices of
stem and branches; peduncles usually not exceeding capitula......

382

375

+ Capitula in paniculate inflorescence; peduncles usually exceeding
Cappitalawens.ritareet eet ells. vetlel rida AN heath Rebbe ene 128.
127. Stem 4—-20(25) cm long. Leaves with scarcely noticeable arachnoid-
hairy pubescence on upper surface, finely grayish arachnoid-hairy
beneath, and like peduncles sparsely covered with sessile, lustrous
glandular hairs. Outer involucral bracts narrow, oblong-ovate (plants

of Chui Steppe in Altai and Tuva [Autonomous] Republic) ........
cocealey.. hoch tachernen 2 cet ees saesdal. pepe 86. S. ambigua Kryl. ex Serg.

+ Stem 25-70 cm long. Leaves glabrous, green above, more or less
grayish pubescent beneath or glabrous. Peduncles and leaves
without or with isolated scattered glandular hairs. Outer involucral
bracts wide, \OMatGuwsidbes. .enedackcsinids cate 82. S. alpina (L.) DC.
128. Leaves usually glabrous beneath or very sparsely tomentose; lower
leaves narrow-lanceolate in type specimen, 1—2 cm wide, less often
wider; cauline leaves usually more than 10, not decurrent on stem.
Floriferous branches forming paniculate inflorescence, less
numerous; inflorescence of fewer, (3)10—15(25), capitula. Florets
lilac, with strong blue tinge (plants of Estonia and the northwestern
European Part of the USSR)...... 84. S. esthonica Baer ex Rupr.

+ Leaves usually gray- or grayish arachnoid hairy-tomentose beneath;
lower leaves broadly lanceolate in type specimen, 2—3.5(4) cm
wide; cauline leaves usually less than 10, usually weakly decurrent

on stem. Floriferous branches forming paniculate inflorescence,
numerous; inflorescence of larger number of capitula (10)15—25(40).
Florets deep red (plants of Siberia, predominantly eastern)..........
sutton. Teemuteeeee: Moe uactdlve Ra veaseransca ls 85. S. stubendorffii Herd.

Subgenus 1. Stephanodontos Lipsch. in Tr. Mosk. Obshch. Ispr.
Prir 3 (1960) 178.—Achenes spinulose with corona of long, stiff, sharp
teeth. Pappus white, double; outer bristles less numerous, distinctly
unequal, up to 10 mm long, persistent; inner bristles numerous, almost
equal, about 15 mm long, plumose, connate at base into a ring.

1. S. carduicephala (Ijin) Ijin in Lipsch. in Tr. Mosk. Obshch.
Ispr. Prir. 3 (1960) 178.—Jurinea carduicephala \\jin in Izv. Tadzh.
Bazy Akad. Nauk SSSR, Botanika, I, 1 (1933) 64.

Perennial. Caespitose; caudices covered with remains of hardened
bases of previous year’s leaves. Stem (5)10—12 cm high, straight, simple,
like leaves covered with short, scattered hairs, sparsely leafy. Leaves
usually at stem base, entire to more or less crenate-toothed or almost
pinnatifid, green above, grayish-floccose beneath with prominent midrib
and less distinct lateral veins; basal leaves numerous, reaching capitulum
or shorter, elliptical, elliptical-lanceolate, oblong-spatulate or lanceolate,

383

376

gradually attenuate into petiole, sheathed, 4-10 cm long with petiole and
1.5—2 cm wide, petiole (1.5)2—4 cm long; cauline leaves numerous, linear
or lanceolate, 2-5 cm long and 1-5 mm wide. Capitula solitary, large,
about 2—2.5 cm long and about 2.5—3 cm wide. Involucre arachnoid hairy-
pubescent, glabrescent, light colored or anthocyanin-colored; involucral
bracts numerous, many-rowed, linear, aristate, slightly recurved when dry;
outer bracts almost as long as inner. Receptacle densely covered with
linear-subulate, short, about 2—3 mm long scales; florets pink, about 2 cm
long, narrow part of tube about 7 mm long, broadened part together with
lobes of limb about 1.2 cm long, both with occasional lustrous glandular
hairs; appendages of anthers divided, setaceous, 3-4 mm long. Achenes
oblong, about 6 mm long, dark brown, ribbed, spinulose along ribs,
especially in upper part, forming corona; corona of long, almost equal,
stiff, sharp teeth; pappus snow-white, two-rowed; outer bristles less
numerous, distinctly unequal, 3-10 mm long, finely plumose-scabrous,
persistent; inner bristles numerous, almost equal, about 15 mm long,
plumose, connate at base into shedding ring. Flowering VII-VIII.

Stony slopes, clefts in rocks, screes——Soviet Central Asia: Pamiro-
Alai (mountainous Badakhshan). Endemic. Described from Roshan. Type
in Leningrad.

Note. Differs from all other Saussurea species found in the USSR
by the spinulose achenes and a corona of long, stiff, acute teeth. With
respect to these characters, it approaches the Indian species S. ceratocarpa
Decne. It is an intermediate species connecting the genera Saussurea
and Jurinea.

Subgenus 2. Amphilaena (Stschegl.) Lipsch. in Tr. Mosk. Obshch.
Isp. Prir. 3 (1960) 182; Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 369.—Sect. Amphilaena Stschegl. in Bull. Soc.
Nat. Mosc. XXI, 3 (1848) 244.—Sect. Eriocoryne (gen. Aplotaxidis)
Wall. ex DC. Prodr. VI (1837) 541 p. p——Sect. Bracteatae Clarke Comp.
Ind. (1876) 220.—Sect. Obvallatae Maxim. in Bull. Acad. Sc. Pétersb.
XXVII (1881) 489.—Bracteal leaves wide, colored, crowded below
corymb of clustered capitula, and enclosing them. In several species
found outside the USSR, included in this subgenus, the capitula are
solitary or few, with swollen peduncles. Receptacle sparsely covered
with short scales. Pappus biseriate, outer bristles few, small, finely
plumose-scabrous, inner bristles long-plumose.

2. S. involucrata Kar. and Kir. ex Maxim. in Bull. Acad. Sc. Pétersb.
XXVII (1881) 489 in textu; Lipschitz in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVI (1954) 439.—S. involucrata (Kar. and Kir.) B.
Fedtsch. in Perech. Rast. Turk. TV (1911) 235.—Aplotaxis involucrata

384

B17

Kar. and Kir. in Bull. Soc. Nat. Mosc. XV, 2 (1842) 389—Haplotaxis
involucrata (Kar. and Kir.) Ldb. Fl. Ross. I, 2 (1845-1846) 673.—S.
karelini Stschegl. in Bull. Soc. Nat. Mosc. XXI, 3 (1848) 244.—S. lioui
Ling Yong in Contr. Inst. Bot. Nat. Acad. Peiping, III, 4 (1935) 149,
tab. XXIX; Lipschitz, op. cit. (1954) 440.—Amphilaena involucrata
Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954)
439.—Ic.: Iubilaeum semisecularem doctoris...G. Fischer de
Waldheim... Mosquae (1847) tab. 2.

Perennial. Root collar stupeous from fibrous remains of previous
year’s leaves. Stem (10)13—25(35) cm high and 2-3 cm thick, fistular,
densely leafy and, hence, often not visible, succulent, glabrous. Basal
and cauline leaves coriaceous, oblong, or oblong-ovate, up to 14 cm
long and up to 2.5 cm wide, sessile, semiamplexicaul, narrowed toward
base, obtuse or subacute, with distinct, straw-colored midrib broadened
in lower part of leaf, smooth on both sides or somewhat scabrous due
to more or less numerous, scattered, lustrous glandular hairs, strongly
and finely serrate-toothed, teeth numerous, small, terminating in
cartilaginous cusp, with dense, capitate glandular hairs between teeth.
Bracteal leaves 13-17, tow-rowed, light (creamy) scaly, with many veins,
like cauline leaves in shape but usually wider, up to orbicular, 5.5—7
cm wide, with frequent sharp teeth, two times as long as and completely
enclosing inflorescence. Capitula sessile, 10-20, in compact terminal
globule enclosed in bracteal leaves. Involucre three or four-rowed, bracts
lanceolate, acute, with blackish border or entirely black, with stiff hairs,
especially at apex, sometimes glochidiate, as a result pubescence of
bracts often indistinct. Receptacle sparsely covered with short scales;
florets dirty violet, about 1.4 cm long; narrow part of tube about 0.6 cm
long, broad part of tube with about 0.8 cm long lobes of limb, filaments
papillate; appendage of anthers ciliate. Pappus double, outer bristles less
numerous, finely plumose-scabrous, fragile, about 5 mm long, inner
bristles long-plumose, about 1-1.2 cm long; achenes dark brown,
glabrous, ribbed. Flowering VII-—VIII.

Stony screes and deposits in alpine zone-—Soviet Central Asia: Tien
Shan, Dzhungaria-Tarbagatai; Eastern Siberia: Angara-Sayans (Tanu-
Ola). General distribution: Dzhungaria-Kashgaria (Tien Shan). Described
from the alpine zone of the Dzhungarian Alatau. Type in Leningrad;
isotype in Moscow.

Note. This species is close to the related. Himalayan species S.
obvallata Edg. According to Karelin and Kirilov, the entire plant,
especially the capitula, emits a strong odor causing vomiting.

The biological role of the bracteal leaves is not fully understood.
Apparently, they protect the inflorescence from sharp temperature
fluctuations and play the role of floral flags attracting pollinator insects.

38

lo)

378

Subgenus 3. Frolovia (DC.) Lipsch. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XXI (1961) 370.—Genus Frolovia (DC.) Lipsch. in
Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954) 461.—Sect.
Frolovia generis Aplotaxidis DC. Prodr. VI (1837) 538.—Sect. Frolovia

_ generis Haplotaxidis in Endlich. Gen. pl. (1836-1840) 468 and in Ldb.

Fl. Ross. II, 2 (1845-1846) 672.—Sect. Frolovia (DC.) Kitam. generis
Saussureae in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII (1937) 141.—Stem
fistular, tall one-headed, less often, when branched, capitula two to five.
Basal leaves large, lyrate or crenate-lobed in type specimen. Capitula
cupuliform, large, 3-5(6) cm wide, often drooping, on long peduncles.
Involucral bracts lanceolate-linear; outer bracts more or less decurved.
Florets up to 3 cm long. Receptacle densely covered with 0.8—1.5 cm long
scales. Pappus one-rowed. Achenes large, usually with corona.

3. S. frolovii Ldb. Ic. pl. Fl. Ross. IV (1833) 16; Ldb. Fl. alt. 1V
(1833) 15; Kryl. Fl. Alt. II, 711; Kryl. Fl. Zap. Sib. XI, 2928—Aplotaxis
frolovii DC. Prodr. VI (1837) 538—Haplotaxis frolovii Ldb. Fl. Ross.
II, 2 (1845-1846) 672.—Cnicus altaicus Steph. ex DC. Prodr. VI (1837)
538 (nom. nud.).—Frolovia lyrata Ldb. op. cit. (1833) 17 and DC. Prodr.
VI (1837) 598 (pro syn. nom. nud.).—Frolovia ledebouriana Lipsch. in
Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954) 461.—Ie.:
Ldb. Ic. pl. Fl. Ross. IV (1833) tab. 352.

Perennial. Rhizome thick; stems solitary, less often two or few,
straight, 25-100 cm high and up to | cm wide, fistular, finely sulcate,
arachnoid hairy-pubescent, in upper part, mainly below capitulum,
tomentose, sometimes glabrous, usually simple, one-headed, occasionally
with one branch and then two-headed, sparsely leafy or leafless. Leaves
lyrate-pinnatisect, with larger triangular or ovate-triangular, acuminate
(less often somewhat obtuse), almost sagittate, unevenly crenate-toothed
or almost entire terminal lobe, lateral lobes gradually reduced toward
base, usually runcinate; all leaves green above, with distinct midrib,
petiolate, sparsely covered with short stiff hairs, grayish arachnoid hairy-
woolly beneath, almost tomentose; basal leaves 4—11, long-petiolate,
(10)15—50 cm long with petiole and 5—20 cm wide; cauline leaves absent
or few (two or three), smaller, -short-petiolate; uppermost leaf reduced,
oblong-lanceolate or almost linear, entire or weakly toothed. Capitula
solitary terminal, occasionally two, 3-4 cm wide, drooping. Involucre
of numerous bracts; bracts lanceolate-linear, long acuminate, blackish-
green on outerside, pale yellow on inner side, covered with hairs and
short bristles often along margin and apex, usually subglabrous; inner
bracts divergent, connivent after fruiting; outer bracts decurved.
Receptacle densely covered with linear, about 1 cm long, straw-yellow,
lustrous scales; florets dirty reddish-violet, about 1.5 cm long, narrow

385

Plate XXX.

1—Saussurea salemanii Wink|.; 2—S. masarica Lipsky.

379

387

380

part of tube about 0.7 cm long, broad part with lobes of limb (excluding

_ anthers) 0.6 cm long; anthers blackish-violet, basal appendages of anthers

linear, pubescent. Pappus one-rowed, its bristles somewhat stiff,
yellowish, long-plumose, about 1.1 cm long; achenes oblong-obovoid,
mature achenes dark brown, ribbed, 6-8 mm long and 2-3 mm wide,
with corona of obtuse, somewhat thick teeth. Flowering VII—VIII.

Alpine and subalpine meadows, tundra, forest meadows, coniferous
forests, including larch forests.—Soviet Central Asia: Dzhungaria-
Tarbagatai; Western Siberia: Irtysh (eastern part), Altai; Eastern Siberia:
Angara-Sayans (western Sayans, Tuva Region). General distribution:
Mongolia. Described from Altai. Type in Leningrad.

Note. According to the original diagnosis of Ledebour, in S. frolovii
the pappus is 2-seriate and its outer bristles are deciduous; I found only
a one-rowed pappus in the entire material at my disposal. This species,
as also the three species close to it, namely, the two species described
by M.M. Iljin from Soviet Central Asia.—S. sulcata Ijin and S. asbukinii
Iljin—and the third described from Taiwan—S. formosana Hayata (Ic.
Pl. Formos. VUI (1919) 69; Kitam. in Mem. Coll. Sc. Kyoto Univ. ser.
B, XIII (1937) 142, tab. XIX, fig. 2 = Frolovia formosana (Hayata)
Lipsch. comb. nova)—given a more detailed understanding of composite
genera, can be separated with complete justification under the separate
genus Frolovia (DC.) Lipsch.

4. S. asbukinii Iljin in Bot. Zhurn. SSSR, XXVII, 6 (1942) 144.—
S. frolovii var. procera Herd. in O. and B. Fedtsch. Perech. Rast. Turk.
IV (1911) 235.—Frolovia asbukinii (Iljin) Lipsch. in Bot. Mat. Gerb.
Bot. Inst. Akad. Nauk SSSR, XVI (1954) 462.

Perennial. Rhizome thick, woody, many-headed, covered with brown,
more or less split remains of basal leaves; stem up to 70 cm high, with
well-developed whitish pith, strongly sulcate, arachnoid hairy-pubescent,
especially densely in upper part, branched slightly:above base, branches
simple, long, upward spreading, leafless, terminating in solitary
capitulum. Leaves with petiole up to 30 cm long, up to 10 cm wide,
light green, glabrous above, grayish green beneath due to arachnoid
hairy more or less dense coating, lyrate, petiole up to 15 cm long, with
broad midrib and weak lateral veins beneath; basal leaves numerous,
half as long as stem, with large, obtuse or orbicular, terminal lobe,
middle lobes oblong-oval, lower and lateral lobes similar in shape,
smallest and remote; lateral lobes almost entire with occasional teeth;
lower cauline leaves like basal but somewhat smaller; middle leaves on
shorter petioles; upper leaves remote, small, lanceolate or linear, entire.
Capitula cupuliform, 5—6 cm wide. Involucre imbricate, 5-6 cm wide
and 3-4 cm long at fruiting, strongly arachnoid hairy-pubescent;
involucral bracts stiff, green on outer side, lanceolate-linear, long

388

38]

acuminate, cusp somewhat stiff, with one more or less distinct, weakly
carinate vein; outermost bracts smaller, about 1—-1.2 cm long and 1 mm
wide, recurved; middle bracts about 2 cm long and 2 mm wide, erect,
with only slightly divergent cusp; inner bracts straw-yellow, glossy, about
2.5-3 cm long and 2-3 cm wide, with purple, ciliolate, straight apex;
cilia distinct in majority of outer and middle involucral bracts. Receptacle
densely covered with narrow-linear, snow-white, about 1.5 cm long scales
almost half as long as inner bracts; florets pink; numerous, about 3—3.2
cm long (including narrow part of tube and anther); narrow part of tube
white, about 1.6 cm long, broad part about 0.7 cm long, lobes of limb
about 0.5—0.6 cm long; appendages of anthers strongly divided,
pubescent. Pappus snow-white, one-rowed, 2—2.25 cm long, its bristles
somewhat unequal, long-plumose; achenes oblong-obovoid, compressed,
about 7 mm long and 3 mm wide, glabrous, punctate (with blackish-
brown outline) with prominent veins, and short corona above. Flowering
FV-V.

On rocks and in debris.—Soviet Central Asia: Tien Shan (Mogoltau,
Kuramin Range). Endemic. Described from Soviet Central Asia (Okur-
Tau, Tadzhikistan). Type in Leningrad.

Economic Importance. These plants contain an alkaloid, which has
not been studied in detail; the local people consider it poisonous.

Note. This species, closest to S. sulcata Iljin, has been described
from only one specimen. It differs from the latter by the absence of
tuberous rhizome, lyrate leaves, somewhat larger capitula, and,
particularly, by achenes with a distinct corona (fringe). The material at
my disposal was extremely inadequate for a conclusive decision about
the independent status of the two species. It is quite possible that at a
later stage S. asbukini Ijin will be combined with S. sulcata Iljin into
a single species; in that case, according to priority, it should be named
S. sulcata Ijin.

5. S. sulcata Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada, III, 26
(1922) 101.—Frolovia sulcata (Iljin) Lipsch. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XVI (1954) 462.

Perennial. Rhizome tuberous; collar covered with remains of previous
year’s leaves. Stem tall, 40-60 cm high, deeply sulcate, furcate
(occasionally simple and then one-headed) in upper part, weakly
arachnoid-hairy, almost leafless, only with isolated small leaves at base
of branching. Leaves green; basal leaves numerous, ovate-triangular, 10—
20 cm long, less often up to 25 cm including petiole. Petioles 5-8 cm
long, less often longer; lamina with irregular wide lobes or weakly
incised, in some specimens more deeply pinnatifid (var. pinnatifida
Lipsch.), with prominent veins beneath, and midrib merging with petiole;

382

lamina sometimes slightly oblique. at base, more or less arachnoid-hairy
on both sides, predominantly so beneath; cauline leaves reduced, only
at point of branching of stem, sessile, entire. Capitula solitary (two to
five in specimen), terminal on floriferous branches (10—20 cm long),
large, 2—2.5(3) cm wide and up to 3—3.5 cm long including florets.
Involucre imbricate, bracts light green, stiff, with indistinctly carinate
vein, arachnoid-hairy, ciliate, all bracts distinctly acuminate; outer bracts
smaller, 0.5-1 cm long and 2 mm wide, recurved; middle and inner
bracts oblong, middle about 1.5—2 cm, inner 2—2.5 cm long. Receptacle
densely covered with narrow, snow-white, about 1 cm long scales almost
half as long as inner involucral bracts; florets pink, 2—2.5 cm long
including narrow part of tube and limb; basal appendages of anthers
strongly divided, pubescent. Pappus snow-white, one-rowed, pappus
bristles somewhat unequal, about 2 cm long, long-plumose; achenes about
5 mm long, roundish at apex, without corona, glabrous, glossy, olive-
green, with blackish-brown pattern. Flowering V.
Varicolored outcrops (red clays, conglomerates)—Soviet Central
389 Asia: Pamiro-Alai (Margelan District), Tien Shan (former Karavan
District of the Kirgizian SSR). Endemic. Described from the Margelan
Region, vicinity of the village of Kul. Type in Leningrad.

Note. In the original description, M.M. Iljin did not compare this
species with any other. Genetically and morphologically, S. sulcata Iljin
is close to S. frolovii Ldb., but differs by having a tuberous rhizome,
branched stem, ovate-triangular, usually not very deeply divided leaves
(in S. frolovii they are lyrate-pinnatifid), light-colored involucral bracts,
of which the outer are weakly twisted (in S. frolovii they are blackish-
green, the outer ones strongly twisted), pink, longer (2—2.5 cm long)
florets (in S. frolovii they are dirty reddish-violet, about 1.5 cm long),
smaller achene with roundish apex without corona (achenes are larger
in S. frolovii with a distinct corona of obtuse, somewhat thick teeth).
Conceming the relationship between S. sulcata and S. asbukinii Ijin,
see the note to the latter.

Subgenus 4. Eriocoryne Wall. ex DC. Prodr. VI (1837) 541, p. p.;
Ldb. Fl. Ross. II, 2, 673 p. p., pro sectione generis Haplotaxidis; Hook.
f. Fl. Brit. India, III (1881) 373.—Sect. Congestae Clarke Comp. Ind.
(1876) 221 p. max. p—Small, densely woolly-pubescent alpine plants.
Stem usually short, simple, clavate, with densely arranged leaves.
Capitula rather numerous, sessile, in compact capitate inflorescence on
fistular stem broadened at apex, surrounded by woolly bracteal leaves
(woolly cover brownish or white). Achenes terminating with stiff corona
and style base. Pappus brown or white, one- or two-rowed and then its
outer bristles small.

390

383

6. S. glacialis Herder in Bull. Soc. Nat. Mosc. XL, 3 (1867) 144;
Kryl. Fl. Zap. Sib. XI, 2928; Lipschitz in Bot. Mat. Gerb. Inst. Akad.
Nauk SSSR, XVI, 443.—S. pamirica Winkl. in Tr. Peterb. Bot. Sada,
XI, 1 (1889) 171.—S. sorocephala auct.: Kryl. Fl. Alt. Ili (1904) 710
non Schrenk.—S. (Aplotaxis) approximata Schischk. and Serg. in
Sistemat. Zam. Gerb. Tomsk Univ. 1 (1944) 2.—? S. chthonocephala
Bornm. in Fedde, Repert. Beih. CVIII, 1 (1938) 59.

Perennial. Rhizome long, branched, usually many-headed, spreading
between stones and debris, densely covered with brown remains of
previous year’s leaves; caudices issuing short, 1.5—6 cm long, solitary,
erect, flattened, densely leafy floriferdus shoots; some caudices terminating
in rosettes of leaves. Leaves oblong or lobate, but 1.5—-4 cm long and 0.4—
1 cm wide, somewhat obtuse, entire or, especially in upper part, erose-
toothed (teeth somewhat obtuse), variable in color, green, or grayish-green
due to pubescence, often reddish-violet; basal and lower cauline leaves
somewhat narrowed toward base in short petiole; middle leaves like basal;
upper leaves approximate, reduced, subtending capitate inflorescence;
all leaves, especially those surrounding inflorescence, woolly-villous above
due to thick pubescence of long white hairs; lower surface less hairy
(predominantly in the Soviet Central Asian specimens) or even subglabrous
(in the Altai specimens). Capitula 0.7—-1 cm wide, 10-20 or more,
aggregated at stem apex in almost capitate inflorescence, subtended by
upper bracteal leaves. Involucral bracts equal, membranous; outer bracts
oblong-ovate or oblong, subacute, colored (reddish-violet) throughout,
especially strongly in upper half, villous due to dense, white, long,
entangled hairs; inner bracts lanceolate, reddish-violet, lustrous, acuminate,
often toothed in upper half along margin and at apex, appressed-pubescent
(hairs white). Receptacle covered with short scales or almost glabrous,
florets pink, purple when dry, about 1—-1.2 cm long; narrow part of tube
about 0.6 cm long, broad part 0.6 cm long including lobes of limb; anthers
whitish, with linear hairy at apex basal appendages. Outer pappus bristles
isolated, serrate, shedding, about 2 mm long, inner bristles long-plumose,
slightly hard at base, connate into ring; white, only in lower part sometimes
brownish-violet, about 1—1.1 cm long. Achenes oblong-cylindrical, about
0.5 cm long, glabrous, smooth. Flowering VII-VIII.

Stony screes, gravelly slopes, pebbles and sands in river beds in
alpine zone.—Western Siberia: Altai; Soviet Central Asia: Pamiro-Alai,
Tien Shan, Dzhungaria-Tarbagatai (Saur, Mustau Range). General
distribution: Indo-Himalayas (?), Dzhungaria-Kashgaria. S. glacialis was
described from the Saryjas Glacier in Tien Shan, and S. pamirica from
Pamir. Type in Leningrad.

Note. S. chthonocephala Bornm., described by Bornmiiller from
northwestern India (Chitral), as is evident from the diagnosis and diagram,

384

should be included among the synomyms of S. glacialis; | did not see
Bornmiiller’s type specimen of this species.

7. S. gnaphalodes (Royle) Sch. Bip. ex Klatt in Nova Acta Acad.
Leopol.-Carol. Nat. Curios. XLI, pars II, 6 (1880) 406; O. and B. Fedtsch.
Perech. Rast. Turk. IV, 236; Lipschitz in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVI, 442.—S. gnaphalodes (Royle) Ostenfeld in
Hedin Southern Tibet, VI, 3 (1922) 33.—Aplotaxis gnaphalodes Royle,
Ill. Bot. Himal. II (1839) 251 and DC. Prodr. VI (1837) 542.—Aplotaxis
sorocephala Schrenk, Enum. pl. nov. I (1841) 43.—Saussurea
sorocephala Schrenk, Enum. pl. nov. II (1842) 38.—Haplotaxis
sorocephala Schrenk in Ldb. Fl. Ross. II, 2 (1845-1846) 673.—Saussurea
sorocephala Hook. f. and Thoms. in Clarke, Comp. Ind. (1876) 226;
Hook. f. Fl. Brit. India, III, 377.—Ie.: Royle, Ill. Bot. Himal. II (1839)
t. 59, f. 1; Blatter, Beautiful Flowers of Kashmir, I (1927) t. 33;
Pampanini, R. La. Fl. del Caracorum in Spedicione Ital. de Filippi nell
Himalaia, Caracorum etc. (1913), ser. I], v. X (1930), tab. VII, fig. 2
(nigra).

Perennial. Rhizome brown, spreading among stones, many-headed,
caudices densely covered with brownish remains of old leaves; some
caudices terminating in leaf rosettes, most producing single, floriferous,
erect, miniature (1.5—6 cm long), very densely leafy stem terminating
in capitate inflorescence of numerous sessile capitula, enclosed by densely
brownish-woolly bracteal leaves. Leaves white-tomentose on both sides,
oblong or lobate, 2-4 cm long and 0.3—1 cm wide, obtuse or subobtuse,
toothed (mainly in upper part of leaf; teeth roundish) or entire; basal
leaves short-petiolate; middle cauline leaves like basal; bracteal leaves
approximate, sessile, not narrowed at base, enclosing capitate
inflorescence. Involucre 0.7—1 cm wide; bracts equal in length, outer
oblong or oblong-ovate, subacute throughout, more strongly in upper
part, densely villous with reddish-brown or white hairs; inner bracts
lanceolate, membranous, lustrous, acuminate, often weakly toothed,
reddish brown-woolly; receptacle covered with short, linear-subulate,
lustrous scales; florets pink, about 0.9-1 cm long, narrow part of tube
about 0.4 cm long, broad part including limb about 0.5 cm long; anthers
whitish, basal appendages of anthers densely ciliate. Pappus colored
(brownish-red), outer pappus-bristles less numerous or solitary, scarcely
noticeable, shedding, about 3 mm long, inner ones 12—14, long-plumose,
about | cm long, almost as long as corolla; achenes oblong-cylindrical,
about 0.5 cm long, smooth. Flowering VII-—VIII.

Stony screes, gravelly slopes, pebbles, moraines, shale rock in alpine
zone.—Soviet Central Asia: Pamiro-Alai (Alai Range, Pamir, Shugnan,
including Trans-Ili Alatau), Tien Shan, Dzhungaria-Tarbagatai
(Dzhungaria-Alatau). General distribution: Indo-Himalayas, Dzhungaria-

385

Kashgaria, Tibet, China (Szechuan). S. sorocephala Schrenk was
described trom the Dzhungarian Alatau; type specimen in Leningrad; S.
gnaphalodes (Royle) Sch. Bip. was described from the Himalayas; type
in Kew; isotypes in Leningrad and Geneva.

Note. This plant was first described by Schrenk as Aplotaxis, and
later he included it in the genus Saussurea. This combination (S.
sorocephala Schrenk) remained unknown to English authors (Hooker,
Thomson, Clarke), and therefore, they wrongly and illegitimately
proposed the c-mbination S. sorocephala Hook. f. and Thoms. S.
sorocephala Schrenk is identical with the earlier described Aplotaxis
gnaphalodes and, therefore, should be called Saussurea gnaphalodes
(Royle) Sch. Bip. Ostenfeld did not know of the existence of this
combination, which was left out of “/ndex Kewensis’”; therefore, he
proposed the same name (S. gnaphalodes (Royle) Ostenfeld) again.

392 Subgenus 5. Saussurea—Subgenus Eusaussurea Hook. f. Fl. Brit.
India, III] (1881) 365.—Glabrous or pubescent, sometimes villous or
woolly herbs. Stems solitary or several, usually distinctly developed,
simple or branched, less often almost completely reduced, and then plants
without stem. Leaves rather variable in shape and size of lamina and its
base (entire, crenate-toothed, pinnatipartite or pinnatifid, etc.), decurrent
on stem forming wing or nondecurrent; basal leaves often withering by
the time of blooming. Capitula sessile or pedunculate, solitary, less
numerous or numerous; in latter case, capitula arranged in compact or
lax corymbs, in turn often forming paniculate inflorescence. Involucre
cylindrical, ovoid, globose, hemispherical, etc.; bracts obtuse or acute,
with straight or recurved cusp, many-rowed, unequal and then imbricate,
less often equal or almost equal; in some groups of species, outer bracts
herbaceous, in others with apical, colored appendages; appendages
herbaceous, saber-shaped, straight or recurved. Receptacle of various
types: 1) scaly with unequal; 2) papillate; or 3) glabrous, without scales.
Achenes truncate or with short corona, bearing outer pappus bristles, if
present. Pappus one or two-rowed, outer bristles, if present, serrate or
finely plumose, stiffer and shorter, usually fragile, inner bristles long-
plumose, with thickened bases connate into entirely shedding ring.

. Section 1. Pyecnocephala Lipsch. in Bot. Mat. Gerb. Inst. Akad.
Nauk SSSR, XVI (1954) 447; Ej. op. cit. XXI (1961) 370.—
Lagurostemon Cass. Dict. sc. nat. LIII (1828) 466 p. p.; Wydl. in
Linnaea, V, 425; DC. Prodr. VI, 532 p.p. saltem §* excl. S. taraxacifolia;
Less. Synops. Compos. (1832) 11.—Sect. Strictae Clarke, Comp. Ind.
(1876) 222 p. p. saltem §b.—Sect. Caulescentes Hook. f. FI. Brit. India,
III (1881) 371, saltem §*** p. min. p.—Rhizome and root not divided
in fibers along their length. Capitula large 1.5—2.5(3) cm wide, in racemes

393

386

or corymbose-paniculate inflorescence, sometimes solitary. Involucral
bracts wide, almost equal. Receptacle covered with cartilaginous yellow
papillae, often connate almost up to apex, or covered with somewhat
long, linear-subulate, white, lustrous scales. Basal appendages of anthers
woolly. Pappus two-rowed.

Series 1. Baicalenses Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 370.—Stem fistular. Inflorescence racemose,
less often capitula solitary. Leaves lanceolate, oblong or oblong-ovate.
Receptacle covered with cartilaginous yellow papillae, sometimes connate
almost up to apex.

Besides S. baicalensis (Adams) Robinson, which is found in the
USSR, this series includes two more species: S. dorogostaiskii Palibin
(Mongolia) and S. calobotrys Diels (China).

8. S. baicalensis (Adams) Robinson in Proc. Amer. Acad. XLVII
(1911) 216; Lipschitz in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XVI, 446; Popov, Fl. Sr. Sib. II, 764—Liatris baicalensis Adams in
Mem. Soc. Nat. Mosc. V (1811) 115.—Saussurea pycnocephala Ldb.
Ic. pl. Fl. Ross. 1 (1829) 15; Ej. Fl. alt. 1V, 14 p. max. p.; Ej. Fl. Ross.
II, 2, 661 p. max. p.; Herd. in Bull. Soc. Nat. Mosc. XLI, 3, 7, saltem
a. genuina Herd.; Kryl. Fl. Zap. Sib. XI, 2907.—Saussurea pycnocephala
a. papillosa Kryl. Fl. Alt. Ill (1904) 702.—Saussurea liatroides Fisch.
in DC. Prodr. VI (1837) 532; Turcez. Fl. baic.-dahur. II, 1, 104.—
Lagurostemon liatroides Fisch. ex Wydl. in Linnaea, V, 3 (1830) 427.—
Serratula liatroides Adams ex Wydl. op. cit. (1830) 427; DC. op. cit.
(1837) 532.—Carphephorus baicalensis DC. op. cit. (1836) 132.—Ie.:
Ldb. op. cit. (1829) tab. 59; Popov, op. cit. Plate 89, Fig. 2.

Perennial. Root collar densely covered with brownish, often fibrously
split remains of old petioles. Stem fistular, straight, solitary, less often
few, 15-60 cm long and 0.5—1 cm wide, ribbed, often flattened and
colored, usually glabrous below, pubescent in middle and especially upper
part with glandular as well as simple, long, white hairs. Leaves
lanceolate, oblong or oblong-ovate, long acuminate, serrate-toothed, teeth
cartilaginous at apex; all leaves covered, especially above and along
margin, with fine, somewhat stiff, glandular hairs besides sparse white
tomentum, more strongly tomentose along margin and on upper leaves;
lower leaves numerous, 6—25 cm long and I—3 cm wide, short-petiolate,
strongly broadened at base and enclosing stem; veins on lower surface
reaching 5 mm in width and extended in petiole; middle and upper
cauline leaves sessile, not decurrent or weakly decurrent over stem,
reduced and narrower. Capitula 1.5—-3 cm wide, less often solitary, often
numerous (4—15), on short peduncles, exserted from axils of small upper
leaves, forming racemose inflorescence at stem apex; less often capitula

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aggregated at apex in dense, almost corymbose inflorescence; peduncles
and bracts, often subtending lower capitula, densely covered with
somewhat thick glandular hairs. Involucre white-woolly or almost
glabrous; all involucral. bracts almost equal, outer resembling bracteal
leaves, ovate-lanceolate or lanceolate, long acuminate, dark colored
(blackish-violet), with less numerous glandular hairs; inner bracts
narrower, glabrous in lower half, usually with numerous, short glandular
hairs in upper half, often with simple long hairs at apex. Receptacle
covered with short, yellowish, cartilaginous papillae, often connate almost
up to apex; florets dirty violet, about 1.5 cm long; narrow part of tube
about 0.8 cm long; broad part about 0.7 cm long including lobes of
limbs; basal appendages of anthers flocculose-woolly. Pappus double,
outer bristles serrate, small, 0.1—-0.2 cm long, usually persistent, inner
ones long-plumose, about 1—-1.2 cm long; achenes glabrous, weakly
glossy, ribbed; mature achenes black or olive-green, with poorly
developed corona at apex. Flowering VII—VIII.

Moss-lichen covered tundras, gravelly and stony slopes in alpine
belt, open larch forests near the forest limit, less often subalpine
meadows.—Western Siberia: Altai; Eastern Siberia: Angara-Sayans;
Dauria (southwestern). General distribution: Mongolia. Described from
the Baikal Region (Liatris baicalensis) and Altai (S. pynocephala ); types
in Leningrad.

Note. Ledebour (FI. Ross. II, 661) and a series of later authors
wrongly reported S. baicalensis for Soviet Central Asia. Specimens from
Soviet Central Asia distributed as this species belong to S. sordida Kar.
and Kir. It hybridizes with S. schanginiana (Wydl.) Fisch. The hybrids
were described as var. heteromorpha (Turcz.) Lipsch. of the latter species
(see note to S. schanginiana). On the other hand, it undoubtedly is very
close to S. krylovii. Thus, some specimens (Baikal—Berguzin,
Shtubendorf; Altai—Elo, Bunge; Altai—upper reaches of Yustyd,
Vereshchagin) are completely the same as S. baicalensis, in habit, leaf
shape, and inflorescence, but their receptacle is similar to that of S.
krylovii, 1.e., distinctly scaly.

9. S. dorogdstaiskii Palibin in Zhurn. Russk. Bot. Obshch. XIII, 1—
2 (1928) 109; Lipschitz in Bot. Mat. Gerb. Bot. Inst. XVI, 448.—Ic.:
Palibin op. cit. (1928) 110.

Perennial. Plants up to | m high; root collar densely covered. with
fibrous remains of previous year’s leaves. Stem fistular, robust in type .
specimen, about 1.5~2 cm wide in middle and below, ribbed, more or
less densely pubescent with long white hairs; subsequently often almost
turning bare, usually densely leafy. Leaves oblong-ovate, hard, frequently

[sic]; recte pycnocephala—Sci. Ed.

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388

and more or less deeply crenate-toothed, teeth cartilaginous, lamina
scabrous on both sides, especially along margin, due to more or less
numerous spinules; basal and lowermost cauline leaves large, obtusely
acuminate, decurrent in broad-winged petiole; midrib straw-yellow
beneath, usually very broad, up to 1-1.5 cm wide, merging into petiole;
upper cauline leaves sessile, not decurrent or somewhat decurrent,
somewhat amplexicaul. Capitula numerous, large, on thick pubescent
peduncles, in dense racemose inflorescence, reaching 5-8 cm in width.
Involucre densely white-lanate, up to 3.5 cm wide, bracts almost equal,
outer ovate-lanceolate or lanceolate, dark-colored, long acuminate,
glandular-hairy, spinose along margin, inner narrower. Receptacle covered
with short, cartilaginous, yellowish papillae, often connate almost up to
apex; florets dirty-brown, about 1.5—1.6 cm long; narrow part of tube
about 0.8—0.9 cm long, broad part including lobes of limbs about 0.6—
0.7 cm long; basal appendages of anthers woolly. Pappus double, outer
pappus bristles serrate-scabrous, unequal, up to 3 mm long, persistent,
inner ones long-plumose, about I—1.1 cm long; achenes glabrous, ribbed,
somewhat curved, blackish-brown, with poorly developed corona at apex.
Flowering VII—VIII.

Gravelly and stony slopes, screes, alpine meadows. Not yet found
in the USSR; however, it may be found in future as it grows in the
bordering areas of Mongolia. General distribution: Mongolia. Endemic.
Described from Mongolia; type in Leningrad.

Note. In the original diagnosis and description of the species in
Russian, I.V. Palibin made several mistakes: 1) the pappus in S.
dorogostaiskii Palibin is not one-rowed, but two-rowed; 2) the achenes
are blackish when mature, not differing from S. baicalensis; and 3) the
receptacle is not glabrous but covered with cartilaginous papillae. In a
number of specimens, the characteristic features: 1) plant height; 2)
width and density of racemose inflorescence; 3) width of vein in the
petiole; 4) diameter of stem; 5) woolly-tomentose pubescence of
involucre; and 6) blunt taper of leaf apex—are transitional to S.
baicalensis. Possibly, these are hybrids between the two species;
additional observations in nature are needed.

Series 2. Interpositae Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 370.—Capitula (two) three or four, densely
arranged at stem apex, or more (5-11), in racemose, inflorescence, less
often solitary. Leaves narrow-lanceolate, with distinct remote teeth.
Receptacle covered with scales.

Type of series: S. krylovii Schischk. and Serg.

This series connects the series Baicalenesis with Pygmaeae (see
below). The only species of this series (S. Arylovii) and the forms close
to it are undoubtedly of hybrid origin. In my view, the characters of S.

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389

krylovii are quite persistent and stable in typical specimens, and,
therefore, it should be considered as a separate species. It is quite
probable that it is a result of ancient hybridization between S. baicalensis
and S. schanginiana, subsequently fixed for selection. In that case, S.
krylovii inherited the multiple capitula, a tendency for forming racemes,
and the leaf shape and dentation from S. baicalensis, and the scaly
raceptacle from S. schanginiana. In the “spectrum of segregation” S.
krylovii, like any population of hybrid origin, produces forms closer to
one of the parents, i.e., closer to S. schanginiana (see below, especially
var. heteromorpha (Turcz.) Lipsch.), or S. baicalensis.

10. S. krylovii Schischk. and Serg. in Sistemat. Zam. Gerb. Tomsk.
Univ. | (1944) 1; Kryl. Fl. Zap. Sib. XI, 2908.—S. pycnocephala Ldb.
B. paleata Kryl. Fl. Alt. III (1904) 702.—S. pycnocephala Ldb. F1. alt.
IV (1833) 14 ex min. p.; Ej. Fl. Ross. II, 2, 661 ex min. p.—S. pygmaea
Spr. y. heteromorpha quoad pl. Song. a cl. Kar. and Kir. leg.; sched. in
herb. Inst. Bot. Acad. Sc. URSS.

Perennial. Root collar densely covered with brown, somewhat fibrous
remains of petioles. Stems (10)15—50 cm high, straight, green or red,
ribbed, thinly covered with long white hairs together with somewhat
thick glandular hairs, often glabrescent. Leaves narrow-lanceolate to
lanceolate-linear, long acuminate, distinctly serrate-toothed, teeth remote,
straight or weakly falcate; basal and lower cauline leaves narrow and
long, petiole, thickened at base, 10-20 cm long with petiole and (0.6)1-—
1.2(1.5) cm wide; middle and upper cauline leaves equal in width, sessile,
not decurrent or weakly decurrent; all leaves scabrous on both sides, as
well as along margin due to small cilia. Capitula (two) three or four,
densely arranged at stem apex, or more (5—11) in racemose inflorescence,
and then lower capitula remote on 1-4 cm long peduncles, less often
solitary. Involucre 1-2 cm wide; outer bracts shorter than middle and
inner in type specimen, less often almost as long, oblong-ovate or ovate-
lanceolate, dark colored (blackish-violet), sparsely pubescent with long
white hairs and scattered glandular hairs; inner bracts lanceolate, long
acuminate, straw-yellow in lower part, lustrous, dark colored above and
similar to outer bracts. Florets dirty violet, about 1.3—1.4 cm long; narrow
part of tube about 0.7—-0.8 cm long, its broad part including lobes of
limb about 0.5—0.6 cm long; basal appendages of anthers woolly-barbate.
Receptacle densely covered with linear-subulate, white, lustrous, about
0.5 cm long scales; pappus double, outer pappus-bristles serrate-scabrous,
about 0.1—0.2 cm long, persistent, inner ones long-plumose, about 1-1.1
cm long. Achenes 0.5—-0.7 cm long, reddish-brown or dark brown.
Flowering VII—VIII.

Gravelly slopes, mountain plateaus, forest edges, meadows in lower
part of the alpine belt and upper forest zone-——Western Sibera: Altai;

390

Soviet Central Asia: Dzhungaria-Tarbagatai. Described from Altai
(“between the valley of the Kalgutta River and the pass to Ak-kol
River”). Type in Tomsk; isotype in Leningrad.

Note. This species is close to S. baicalensis (Adams) Robinson,
from which it can be distinguished immediately by the presence of linear-
subulate receptacular scales. Ideas on the origin of S. krylovii are
discussed in the note to the description of series /nterpositae. It differs
from S. sordida Kar. and Kir. by having racemose inflorescences,
narrower, apically attenuated, lanceolate or linear-lanceolate leaves, and
more remotely toothed leaf margins. The specimens of Karelin and
Kirilov, which are cited in the literature under the name S. pygmaea y.
heteromorpha (Herd. in Bull. Soc. Nat. Mosc., XLI, 3 (1868) 5 p. p.
saltem quoad pl. Kar. and Kir.) also belong to S. krylovii Schischk. and
Serg. Unquestionably the latter specimens approach S. krylovii, whereas
the Sayan and Baikal specimens approach S. schanginiana (cf. note to
S. schanginiana).

Series 3. Pygmaeae Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 370.—Stem one-headed. Leaves narrow-linear,
entire, less often with isolated teeth along margin. Receptacle with shorter
scales (sometimes absent, for example, in S. Jeucophylla Schrenk). Plants
forming small turf.

Note. Besides the indigenous species S. schanginiana (Wydl.) Fisch.
and S. leucophylla Schrenk, this series also includes species found outside
the Soviet Union: S. pygmaea Spr. (Alps of Western Europe), S.
graminifolia Wall. (Himalayas), S. brunneopilosa Hand.-Mazz. (China,
Kansu), S. eopygmaea Hand.-Mazz. (Tibet). Apparently, the following
species known to me only from their descriptions should also be included
under this series: S. graminea Dunn. (= S. poophylla Diels, according to
Handel-Mazzeti) (Tibet), S. Janicaulis Hand.-Mazz. (Szechuan, Yunnan)
and others.

11. S. schanginiana (Wydl.) Fisch. ex Herd. in Bull. Soc. Nat.
Mosc. XLI, 3 (1868) 5; Kryl. Fl. Zap. Sib. XI, 2906.—Lagurostemon
pygmaeum $*. schanginianum Wydl. in Linnaea, V (1830) 427.—
Heterotrichum schanginii MB. ex DC. Prodr. VI (1837) 532; Herd. op.
cit. (1868) 5—Saussurea pygmaea f. schanginii DC. Prodr. VI (1837)
532.—S. pygmaea auct.: Ldb. Ic. pl. Fl. Ross. I (1829) 15; Ej. Fl. alt.
IV, 14; Ej. Fl. Ross. II, 2, 660 excl. B. leptophylla, non Spr.—S. pygmaea
a. communis Turcz. Fl. baic.-dahur. II, 1 (1856) 105——S. pygmaea a.
typica Kryl. Fl. Alt. III (1904) 701.—S. pygmaea B. sibirica Herd. op.
cit. (1868) 5. .

Perennial. Stem 4-30 cm high, solitary or few, forming small turf
with rosettes of basal leaves; stem usually dark-colored (violet or brown),

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391

more or less villous or pubescent, less often subglabrous, leafy, sometimes
quite densely (var. polyphylla (Schrenk) Lipsch.), basally covered with
shining brownish sheaths of basal leaves of previous year. Leaves linear,
4-20 cm long and 1.5—5(10) cm wide; lower leaves somewhat broader
in middle, entire, less often with occasional, scarcely noticeable, scattered
teeth along margin (f. dentifolia Lipsch.), flat or convolute, subglabrous
or slightly villous; cauline leaves scarcely broadened at base, weakly
amplexicaul. Capitula solitary, occasionally two to five, terminal, up to
3 cm long with florets and (1.5)2—3 cm wide; upper cauline leaves
subtending capitulum usually shorter than capitulum. Involucre more or
less pubescent or villous, less often subglabrous, bracts almost equal,
dark colored (brownish-black), acuminate; outer bracts wider, ovate or
lanceolate; inner bracts narrower, linear-lanceolate or almost linear, straw-
yellow in lower part (covered with outer involucral bracts), glossy.
Receptacle covered with linear-subulate, lustrous, about 0.3—0.5 cm long
scales; florets violet-lilac, about 1.4—-1.5 cm long, narrow part of corolla
tube about 0.7—0.8 cm long, broad part about 0.6—0.7 cm long including
lobes of limb; anthers about 0.6 cm long; basal appendages of anthers
woolly. Pappus smoky, double, outer pappus-bristles serrate, unequal, up
to 24 mm long, inner ones plumose, about 1 cm long; achenes glabrous,
about 0.6 cm long, with short corona at apex. Flowering VII-VIII.

Moss-lichen covered and gravelly tundras, screes and rocks, less
often alpine meadows in the alpine zone, also near the upper limit of
cedar and larch forests——Soviet Central Asia: Tien Shan, Dzhungaria-
Tarbagatai; Western Siberia: Irtysh (eastern high-mountain part), Altai;
Eastern Siberia: Angara-Sayans (Sayans, Tuva [Autonomous] Republic,
southern part of Irkutsk), Yenisei, Dauria (eastern), Lena-Kolyma,
Anadyr. General distribution: Mongolia. Described from Altai (“on shales
in the Kumid Mountains,” Shangin). Type in Leningrad.

Note. The authentic sheet has three labels: the first, written in the
hand of F.B. Fischer with the label “Schanginiana m”, the second, in
Russian, “in Kumid bald peaks on shales,” and the third, in Latin, “in
alpibus, praesertim corgonensibus,” and, in a different hand-writing
below, “D. Schangin.” The third label corresponds to the citation of the
location reported by Wydler (Linnaea, V, 1830, 427). This species varies
in stem length, nature of leafiness, form, and width of leaves, and number
of capitula. These are more clearly demarcated variations.

1. Saussurea schanginiana var. polyphylla (Schrenk) Lipsch. in Bot.
Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XXI (1961) 371.—S. pygmaea
B. polyphylla Schrenk Enum. pl. nov. II (1842) 35.—S. polyphylla
Schrenk ex Herd. in Bull. Soc. Nat. Mosc. XLI, 3 (1868) 5.—S. pygmaea
y. polyphylla Turez. seu Turczaninowiana Herd. op. cit. (1868) 5—The
type specimen originated from the Chocondo Alps. The plant has a

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392

simple, short (5—10 cm long), densely leafy, villous, unicapitulate stem,
linear-filiform, narrow, sparsely villous convolute leaves, upper leaves
more or less enclosing capitulum (shorter than capitulum), and villous
involucral bracts. This variety is found together with typical S.
schanginiana sporadically in the Tuva Republic, on the Vitim Plateau,
Chocondo bald peaks, Gargan Pass of the Turkin Region and in some
other places.

Some specimens deviate from typical S. schanginiana (Wydl.) Fisch.
and are undoubtedly the result of hybridization with S. baicalensis
(Adams) Robinson. These specimens were described as var. heteromorpha
by N.S. Turczaninow.

2. S. schanginiana var. heteromorpha (Turcz.) Lipsch. in Bot. Mat.
Gerb. Bot. Inst. Akad. Nauk SSSR, XXI (1961) 371—S. pygmaea y.
heteromorpha Turcz. Fl. baic.-dahur. II, 1 (1865) 105.—S. schanginiana
(Wydl.) Fisch. var. heteromorpha Turcz. in Popov. Fl. Sr. Sib. II (1959)
764.—Stem 20-30 cm high, usually more or less branched, sparsely
villous, one to five-headed; capitula in somewhat compact paniculate
inflorescence. Basal leaves linear or linear-lanceolate, usually with rather
broad midrib beneath, continued in broad petiole, weakly convolute,
entire or somewhat toothed, teeth often with cartilaginous cusp; cauline
leaves less numerous, usually linear, entire; uppermost leaves not
enclosing capitulum. Involucral bracts dark-colored, villous. This variety
was described from the Oka River (in eastern Sayans). To it besides the
type, belong specimens from the vicinity of the Arashan Health Resort
(Tulkin Region, Buryat Republic), the village of Goloustnyi on the shores
of Baikal, the Munku-Ardyk Alps, the Minusinsk Region (Saksar and
Byshtakh Mountains), and Yakutia. Var. heteromorpha is also quite
close to S. krylovii Schischk. and Serg. (Cf. the description and note of
the latter), a species undoubtedly of hybrid origin, but differing from
typical S. krylovii by having narrower, weakly crenate-toothed or entire
leaves and a more compact inflorescence of fewer capitula. It differs
from typical S. schanginiana by more robust growth, usually wider,
often slightly crenate-toothed leaves, and a mostly branched stem and
few capitula, often in a compact terminal inflorescence. In its multi-
headedness, var. heteromorpha resembles somewhat S. baicalensis,
however, the latter clearly differs by the presence on the receptacle of
cartilaginous yellow papillae and by wider leaves. As in all hybrid forms,
many of these characters are quite unstable in var. heteromorpha and
strongly lean to one or the other parent.

12. S. leucophylla Schrenk in Bull. sc. Acad. Sc. Pétersb. X, 23
(1842) 354; Enum. pl. nov. II (1842) 35; Kryl. Fl. Zap. Sib. XI, 2907.—

393

ae
SSS SSS =

——

Plate XXXI.
Saussurea tomentosa Kom.; 2—S. kitamurana Miyabe and Tatewaki;

401

1

3—S. sajanensis Gudoschn.

400

394

S. pygmaea B. leucophylla Ldb. .Fl. Ross. II, 2 (1845-1846) 660.—S.
pygmaea var. juldusica Winkl. in O. and B. Fedtsch. Consp. Fl. Turkest.
IV (1911) 228.

Perennial. Plants caespitose; root Be tehy many-headed, producing
more or less numerous stems and rosettes of basal leaves. Stem prostrate
or often upright, simple (not at all branched), 2.5-12 cm long, one-
headed, densely white-woolly, covered at base with brown, oblong-
lanceolate, decorative yellow scales—remains of dead leaves. Leaves of
vegetative branches oblong, usually exceeding stem, (4)6-14 cm long
and 1—2(4) mm wide, linear, entire, obtuse, convolute, somewhat
attenuate basally into scaly, brownish, glabrous, persistent sheath; cauline
leaves four to seven, remote, like basal, but shorter, wider at base,
somewhat decurrent, densely white-tomentose up to base on both sides;
uppermost leaves crowded below capitulum, as long as or slightly
exceeding capitulum. Capitula solitary, up to 2.5 cm long including
florets and up to 3 cm wide, densely villous. Involucre, like stem,
somewhat or not at all colored; outer involucral bracts herbaceous,
remote, sometimes squarrose, with broadly ovate-triangular base; inner
bracts narrower, lanceolate or linear, straw-yellow in lower part.
Receptacle with white, linear-subulate, lustrous, about 3-5 mm long
bristles; florets reddish-lilac, about 1.4—1.5 cm long, narrow part of tube
0.7—-0.8 cm long, wider, including lobes of limb about 0.7 cm long;
basal appendages of anthers woolly. Pappus double, outer pappus bristles
serrate, less numerous, unequal, about 3 mm long, inner ones plumose,
about 1.2 cm long; achenes about 0.5 cm long, glabrous, with narrow
margin at apex. Flowering VII-—VIII.

Gravelly tundra, screes, rocks, clayey deposits, alpine meadows in
the alpine zone.—Soviet Central Asia: Pamiro-Alai, Tien Shan,
Dzhungaria-Tarbagatai; Western Siberia: Altai; Eastern Siberia: Angara-
Sayans (Tuva Republic). General distribution: Dzhungaria-Kashgaria,
Mongolia (western). Described from the Dzhabyk Range in the
Dzhungarian Alatau. Type in Leningrad.

Note. By growth habit it can be divided in two forms: almost stem-
less (f. subacaulis Lipsch.) and with distinct stem (f. caulescens Lipsch.),
with a whole range of intermediate forms between them.

Series 4. Sordidae Lipsch.—Inflorescence corymbose-paniculate, .
sometimes capitula solitary. Leaves broadly lanceolate or ovate-
lanceolate, 2.5—7 cm wide, with approximate teeth along margin.
Receptacle covered with scales.

13. S. sordida Kar. and Kir. in Bull. Soc. Nat. Mosc. XV (1842)
389; Ldb. FI. Ross. II, 2, 661; Lipschitz in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVI, 448.—S. pycnocephala Ldb. 8. sordida (Kar.

402

395

and Kir.) Herd. in Bull. Soc. Nat. Mosc.. XLI, 3 (1868) 7.—S.
pycnocephala auct. fl. turk. non Ldb.—S. russowii Winkl. in Catal. sem.
Horti Bot. Petropol. (1883) 41; Ej. in Acta Horti Petropol. IX, 2 (1886)
423.—S. sordida var. russowii (Winkl.) Lipsch. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XVI (1954) 449.—S. serawschanica B. Fedtsch.
in Consp. Fl. Turkest. TV (1911) 229.

Perennial. Root collar covered over a larger part with fibrously split
remains of previous year’s leaves. Stem 20-100 cm high, erect, simple
or branched, weakly or strongly hard-fibrous and then rough, usually
villous in upper and lower parts, sometimes almost glabrous, with sparse,
long, white hairs. Leaves 2-7 cm wide, broadly lanceolate, ovate or
oblong-ovate, acuminate, sparsely pubescent above, scabrous due to
spinules, hispid beneath, especially along veins and margin, less often
almost glabrous; all leaves with prominent midrib, serrate-toothed, teeth
approximate, cartilaginous at apex; basal and lowermost cauline leaves
reaching 20-35 cm in length, petiolate, petioles winged, hispid in large
specimens, sheathing at base; midrib somewhat broadened and attenuate
into petiole; middle cauline leaves sessile, decurrent; upper leaves
reduced. Capitula up to 2—2.5 cm wide, usually numerous, more or less
long-pedunculate in corymbose-paniculate inflorescence, less often
solitary or few (three), almost sessile (var. oligocephala Winkl. ex Lipsch.
in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XXIJ (1961) 371).
Involucre subglabrous or more or less pubescent with white stiff hairs;
bracts appressed, lanceolate or ovate-lanceolate, acuminate, often
spinulose along margin, usually blackish-brown, less often green; outer
bracts almost as long as inner or shorter. Florets many, dirty pink-violet,
about 1.4—1.5 cm long; narrow part of tube about 0.8 cm long, broad
part including lobes of limb about 0.6 cm long; basal appendages of
anthers woolly; receptacle densely covered with linear-subulate, lustrous,
about 0.5—0.7 cm long scales almost half as long as inner bracts. Pappus
double, outer bristles fragile, toothed-scabrous, unequal, about 1-4 mm
long, inner ones long-plumose, about 1-1.1 cm long; achenes glabrous,
mature achenes about 0.6 cm long, dark brown, with narrow corona at
apex. Flowering VIJ—VIII.

Subalpine and alpine meadows, meadow-covered glades in forests,
rocks and screes.—Soviet Central Asia: Pamiro-Alai, Tien Shan,
Dzhungaria-Tarbagatai. General distribution: Dzhungaria-Kashgaria?
(mountainous part). Described from Dzhungarian Alatau; S. russowii
Winkl.—From the Borborogussun River (Xinjiang, toward east of
Sayram-Nor); S. serawschanica B. Fedtsch.—From the Zeravshan River;
type specimens of all these “species” in Leningrad.

Note. An apparently polymorphic species, variable in the following
characters: 1) stem height; 2) shape and width of leaves; 3) degree and
nature of pubescence; 4) roughness of stem surface; 5) number of capitula

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in the inflorescence; 6) presence or absence of peduncles; 7) nature of

pubescence of involucre (almost glabrous or densely pubescent); 8) length
‘of receptacular scales and their ratio to the length of inner involucral

bracts; and 9) ratio of the length of outer and inner involucral bracts.
Consequently, some extreme forms of S. sordida have been incorrectly
described as separate species. These include S. russowii Winkl. and S.
serawschanica B. Fedtsch. K.U. Winkler tried to distinguish S. russowii
Winkl. from the species of Karelin and Kirilov by the following
characters: 1) greater height (up to 1 m) and larger size of all plant
organs: 2) unequal involucral bracts, i.e., outer bracts considerably shorter
than inner ones (for S. sordida Winkler indicated that they are almost
equal); 3) receptacular scales one-third as long as the inner involucral
bracts (according to Winkler, in S. sordida they are half as long as the
inner bracts); and 4) densely hispid stem and petioles. All these characters
are unstable and result in the full range of transitions to S. sordida;
these characters are also not constant in the type material of S. russowii.
Accordingly, I cannot accept the independent status of S. russowii Winkl.
and include it under the robust .forms of S. sordida (S. sordida Kar. and
Kir. var. russowii (Winkl.) Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVI (1954) 449). S. serawschanica B. Fedtsch., described
from Zeravshan by B.A. Fedtschenko, compares not with S. sordida but
only with S. russowii. The specimens from Zeravshan, at first glance,
appear to have thinner, wider, ovate leaves in comparison with the type
specimens of S. sordida, and are characterized by lesser pubescence on
the stem. However, after a careful comparison of them with large
collections of S. sordida the above-mentioned characters do not appear
to hold up absolutely and do not permit us to recognize S. serawschanica
B. Fedtsch separately. The report of Ledebour (Fi. Ross. II, 661) on the
presence of S. baicalensis in Soviet Central Asia, which was based on
the herbarium specimens of Schrenk from the Dzhungarian Alatau, is
incorrect. The specimens of Schrenk, distributed as S. baicalensis, have
a distinctly developed receptacle with scales, corymbose-paniculate (and
not racemose) inflorescence and broad leaves (2-7 cm wide) and should
be referred to the species S. sordida. Thus, true S. baicalensis is not
found in Soviet Central Asia. All reports of authors, who following
Ledebour, have confirmed the discovery of S. baicalensis in Soviet
Central Asia, should be referred to S. sordida.

Section 2. Laguranthera (C.A.M.) Lipsch. in Bot. Mat. Gerb. Inst.
Bot. Akad. Nauk UzbSSR, XV (1959) 17.—Subsect. Laguranthera
C.A.M. ex Endlich. Gen. pl. (1836-1840) 468.—Sect. Lagurostemon
(Cass.) DC. Prodr. VI (1837) 532 p. p. (excl. §*); Kitam. in Mem. Coll.
Sc. Kyoto Univ. ser. B, XIII, 150 and 151 p. p—Rhizomes and roots
fibrously split along length. Capitula narrow campanulate or cylindrical,

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small or up to 1 cm wide, usually numerous, in corymbose or paniculate
inflorescence. Involucral bracts undivided, without appendages, distinctly
imbricate, appressed, ovate, subobtuse or acute, in some species cusp
recurved. Receptacle covered with scales; basal appendages of anthers
fascicular-woolly. Pappus double. Basal leaves mostly withering before
flowering. Perennials or semishrubs.

Series 5. Rigidae Lipsch.—Sect. Lagurostemon, saltem §** in DC.
Prodr. VI (1837) 532.—Stem not woody at base, tall, winged due to
decurrent leaves. Basal leaves withering before flowering. Cusps of
involucral bracts curved.

14. S. rigida Ldb. Ic. pl. Fl. Ross. I (1829) 19; Ej. Fl. alt. IV, 32;

DC. Prodr. VI, 532; Ldb. Fl. Ross. II, 2, 671; Kryl. Fl. Zap. Sib. XI,
2927.—Ic.: Ldb. op. cit. (1829), tab. 79.—Exs.: Kar. and Kir. (1841)
No. 1625.
— Perennial. Root thick, bearing stems up to 80 cm tall and up to 1
cm wide. Stem not woody at base, strong, straight, angular-sulcate, rough,
with scattered glandular hairs, winged, branched. Basal leaves oblong,
pinnatipartite, withering before flowering; lower and middle cauline
leaves undivided, narrow-oblong, decurrent over stem or (according to
Fl. Zap. Sib. [Flora of Western Siberia]) crenate-lobate, sometimes
toothed, with lanceolate-linear acuate lobes, 10-14 cm long and 0.5—1
cm wide; upper cauline leaves and leaves on branches broadly or narrow-
linear, somewhat broadened at base, more or less amplexicaul or slightly
decurrent; all leaves arcuate, scabrous above due to numerous spinules,
glandular-hairy beneath, veins and margin hairy, convolute. Capitula
1.2-1.4 cm long and 0.4—0.7 cm wide, numerous, in dense approximate
corymbs at apices of main stem and branches, aggregated in broadly
corymbose-paniculate inflorescence. Involucre cylindrical, imbricate,
many-rowed; outer involucral bracts ovate, narrowed in subulate,
decurved cusp; inner bracts oblong, colored at apex and with short cusp;
all bracts usually with prominent midrib, more or less arachnoid hairy-
pubescent. Florets pink-lilac, about 1—-1.1 cm long, narrow part of tube
about 0.5 cm long, broad part with lobes of limb about 0.5-0.6 cm
long; appendages of anthers with dense tuft of woolly hairs. Receptacle
covered with dense membranous linear-subulate, lustrous, about 0.8 cm
long scales. Achenes 0.5—0.6 cm long, brownish, finely ribbed; pappus
snow-white, double; outer pappus bristles serrate-scabrous, fragile, 2-3
cm long, inner ones long-plumose, about 1 cm long. Flowering VI-—
VIII.

Shrub and meadow steppe.—Western Siberia: Irtysh. Endemic. .
Described “from meadows of the Dzhungarian-Kirgizian desert”
(Semipalata Region). Type in Leningrad.

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Note. This species differs from the closely related species S. elata
Ldb. and S. elegans Ldb. by having involucral bracts with subulate,
decurved tips and decurrent leaves, because of which the stem appears
winged.

Series 6. Suffruticosae Lipsch—Stems woody at base, wingless,
20-45 cm high. Involucral bracts obtuse or acute; in latter case cusps
not decurved, except in S. ninae Iljin.

15. S. ninae [jin in Bot. Zhurn. SSSR, XXVII, 6 (1942) 145.—S.
dshungarica Iljin in Bot. Zhurn. SSSR, XXVII, 6 (1942) 145 p. p. (f.
canescens \ljin).

Perennial. Root thick, woody, fibrously branched, many-headed, with
more or less numerous, woody, long caudices. Stems up to 10, 15-35
cm high, brownish at base, woody, covered with remains of old petioles
of basal leaves, somewhat ascending, obtusely ribbed, weakly floccose
and arachnoid hairy-pubescent with scattered, sessile, lustrous glandular
hairs, simple or branched in upper half as well as in inflorescence.
Leaves green, weakly arachnoid-hairy (particularly beneath), and with
lustrous glandular hairs; lower and middle cauline leaves oblong or
lanceolate, attenuate toward base and relatively short-petiolate, with
distinct midrib, acute, cuspidate with short, straight or weakly bent
spinulous cusp, crenate-toothed, teeth usually not very frequent,
acuminate-aristate; upper cauline leaves usually reduced, narrowed toward
base, sessile, lanceolate or oblong, sometimes almost entire, with
spinescent cusp. Capitula pedunculate with filiform of lanceolate-linear
bracteal leaves, in corymbose inflorescence; sometimes with branching
of stem, additional peduncles originating from axils of upper leaves,
terminating in one or two capitula. Involucre cylindrical, 1—1.2 cm long
and 0.6—0.8 cm wide, flocculose-arachnoid hairy, especially at apices;
involucral bracts green or violet, imbricate, all terminating in long,
usually decurved, subulate cusp; innermost bracts with shorter and usually
nondecurved cusp; outer bracts ovate or lanceolate, inner oblong-linear.
Receptacle densely covered with linear-subulate, lustrous, unequal scales,
longest up to 0.8 cm long; florets purple, about 1.4—1.5 cm long, narrow
part of tube about 6—7 mm long, broad part (covered with lustrous
glandular hairs) with lobes of limb almost as long as narrow part; anthers
about 0.6—-0.7 cm long, basal appendages of anthers woolly. Pappus
double, snow-white, outer pappus bristles unequal, serrate-scabrous, short
(up to 0.5 cm long), fragile, inner ones long-plumose, about 1—1.1 cm
long. Flowering VII.

Crevices in rocks.—Central Asia: Dzhungaria-Tarbagatai
(Dzhungarian Alatau). Endemic (?). Described from the Koksu River
Basin. Type in Leningrad.

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399

Note. In the original description of S. ninae M.M. Iljin compared it
with S. dshungarica, also from the nearby localities in the Dzhungarian
Alatau and described immediately following S. ninae in the same
publication. An examination of the material used by M.M. Iljin for
describing S. dshungarica showed its heterogeneity, which was also clear
to the author himself, who distinguished two forms—viridis Iljin and
canescens \ljin—in the Herbarium. I do not recognize the separate status
of S. dshungarica but distribute the specimens of this species between
S. ninae \ljin and S. coronata Schrenk. Details of the relationship between
S. dshungarica and S. coronata Schrenk are discussed in the note to the
latter species.

L.E. Rodin’s specimens from Altyn-Emelskii Pass, like the collections
of E.P. Matveeva from the same sites (the latter were annotated by
M.M. Iljin as S. dshungarica f. canescens \ljin) differ from typical S. ninae
Iljin by the absence of a long, subulate, usually decurved cusp of the
involucral bracts, more dense leafy cover on the stem, more deeply
crenate-toothed leaves, a more strongly pubescent lower surface of the
leaf and the capitula more compactly clustered in corymbs at tips of
stems. Evidently, this is a unique race (S. rodinii Lipsch.), which requires
further study in nature. In having of a decurved, subulate cusp on the
involucral bracts, S. ninae Iljin is closer to the series Rigidae, but in the
woody stem base, habit and shape of the capitula and leaves, the texture
of the latter, and other characters, it is undoubtedly closer to the series
Suffruticosae, thus connecting both the series.

16. S. coronata Schrenk in Bull. Acad. Sc. Pétersb. III, 7 (1845)
107; Lipschitz in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI,
443.—S. coronata var. colorata Herd. in Bull. Soc. Nat. Mosc. XL, 3
(1867) 141, No. 583.—S. dshungarica lljin in Bot. Zhurn. SSSR, XX VII,
6 (1942) 145 p. p., f. viridis Iljin in herb.

Perennial. Roots thick, woody, fibrously branched, many-headed,
with more or less numerous, woody, long caudices. Stem up to 40 cm
high, dark brown above, woody, glossy, covered with hard remains of
basal leaves, sulcate, simple or branched in upper part, glabrous. Leaves
coriaceous, green on both sides or more or less grayish-pubescent
beneath, sometimes almost glabrous, oblong or lanceolate-oblong,
acuminate, petiolate, with less prominent midrib beneath, slightly
broadened and merging into petiole; lower and middle cauline leaves
more or less deeply crenate-toothed, teeth unequal, callous-spinescent;
upper leaves lanceolate, reduced, sometimes almost entire; bracteal leaves
on peduncles of capitula rather numerous, small or larger, acuminate or
subobtuse; all leaves with numerous, sessile, glossy glandular’ hairs.
Capitula 3-10, ovoid, in a relatively dense corymb, sometimes peduncles

400

long, and then corymbose inflorescence more spreading. Involucre about
1.5 cm long and up to | cm wide, many-rowed; bracts hard, appressed,
almost glabrous or arachnoid-hairy, glandular-hairy, green; inner bracts
usually violet, acuminate, with straight, not recurved cusp. Florets purple
(?), narrow and broad parts of tube and lobes of limb glandular-hairy;
appendages of anthers with tuft of woolly hairs at base. Receptacle
covered with linear-subulate, lustrous, up to 0.8 cm long, scales. Pappus
double, outer bristles serrate-scabrous, unequal up to 0.5 cm long, inner
ones long-plumose, about I—1.1 cm long; achenes with narrow corona
at apex. Flowering VII-—VIII.

Rocks.—Soviet Central Asia: Dzhungaria-Tarbagatai (Dzhungarian
Alatau). Endemic. Described from the Dzhungarian Alatau, from the
mountains facing the Kizylgach River, north of Kopal. Type in
Leningrad.

Note. In the original description of S. coronata Schrenk, the author
was silent about the affinity of this species and did not compare it with

407 S. blanda Schrenk described from the Dzhungarian Alatau, which, in
my opinion, is closely related. The material of S. coronata is very limited,
and the species is represented by collections of Schrenk as well as some
later collectors (Lipsky, Voroshilov). S. coronata, by comparison with
S. blanda, has less divided and stiffer, more weakly scabrous leaves,
capitula typically more compactly arranged and more numerous
involucral bracts with a distinct cusp, and plants more strongly woody
at base. Concerning the relationship between these two species, additional
collections and observations in nature are needed.

M.M. Iljin described S. ninae Iljin and S. dshungarica \\jin from
the Dzhungarian Alatau. Both are close to S. coronata Schrenk, which
had escaped Iljin’s notice, as he did not compare either of his own
species with the latter, and only considered differences between them.
S. coronata differs from S. ninae by a more strongly reduced stem,
more coriaceous leaves, reduced peduncles of the capitula, and a less
distinct cusp of the involucral bracts, which are not recurved. As regards
S. dshungarica, a study led me to the conclusion that it was described
a second time and must be included among the synonyms of S. coronata.
According to M.M. Iljin, the characteristic features of S. dshungarica
are: 1) strong lignification of the lower part of the stem (which was
also observed in S. coronata); 2) the length of the receptacular scales
reaches only up to half or one-third the length of the involucral bracts
(this character is not stable, showing variations, and the receptacular
scales are similar in length to those in S. coronata); and 3) involucral
bracts with cusps (the latter are present in both S. dshungarica and S.
coronata, and are not recurved).

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401

17. S. blanda Schrenk in Bull. Acad. Sc. Pétersb. X, 23 (1842)
354; Ej. in Enum. pl. nov., II (1842) 37; Ldb. Fl. Ross. II, 671; Lipschitz
in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI, 443.—S. konuroba
Saposhn. in Not. system. Herb. Horti. Bot. USSR, VI, 2 (1926) 32;
Lipschitz. op. cit. 443.

Perennial. Root woody, fibrously branched along length, many-
headed, with more or less numerous caudices, densely covered with
dark brown bases of old leaves. Stems 10-—30(35) cm high, usually
numerous, woody at base, somewhat ascending or upright (less often
weakly flexuous in shady forms), sulcate, glabrous, simple or branched
in inflorescence, sometimes strongly branched from middle in shady
forms (f. umbrosa Ijin), with spreading branches. Leaves hard (thinner
and softer in shady forms), pinnatifid or pinnatisect, oblong, sometimes
divided almost to midrib, sometimes weakly crenate; lateral lobes ovate
to triangular or lanceolate, acute with spinescent cusp or roundish-obtuse;
midrib usually very prominent beneath, especially in lower part of leaves,
merging into petiole; leaves green on both sides, somewhat lighter
beneath; strongly scabrous above and margin in type specimen due to
numerous, stiff, setose hairs and sessile lustrous glandular hairs,
subglabrous beneath or almost glabrous and with numerous lustrous
glandular hairs; uppermost leaves reduced, pinnatifid or entire, with
spinescent cusp. Capitula on long peduncles, solitary or two or three at
stem and branch apices forming broadly corymbose or corymbose-
paniculate inflorescence. Involucre imbricate, many-rowed, about 1.3—
1.5 cm long and 0.8—1 cm wide; involucral bracts appressed, green or
violet, lustrous, almost glabrous or weakly arachnoid-hairy especially at
apex and along margin, besides with less numerous glandular hairs; outer
bracts ovate, inner lanceolate or oblong-linear, somewhat acute but
without recurved cusp. Florets purple, with scattered, lustrous, sessile
glandular hairs, especially distinct on corolla-tube, about 1.3-1.4 cm
long; narrow part of tube about 0.6 cm long, broad part including lobes
of limb about 0.7—0.8 cm long; anthers about 0.6 cm long, basal
appendages white woolly comose. Receptacle covered with linear-
subulate, lustrous, about 0.5—0.6 cm long scales; pappus double, outer
pappus bristles serrate-scabrous, fragile, unequal, about 3-5 mm long,
inner ones long-plumose, about 1—1.2 cm long; achenes mottled-violet,
with small corona. Flowering VII—VIII.

Rocks, cliffs, granite screes.—Soviet Central Asia: Dzhungaria-
Tarbagatai (Dzhungarian Alatau), Tien Shan (Trans-Ili Alatau, western
Talgar). Endemic. Described from the Dzhungarian Alatau (Dzhamantyk
and Dzhabyk). Type in Leningrad.

Note. The specimens from the rocks of Sarkansk Ravine in the
Dzhungarian Alatau (P.P. Polyakov and L.A. Kupriyanova) differ sharply

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402

from the typical S. blanda by having leaves tomentose beneath, strongly
tomentose involucral bracts terminating in a short black cusp and capitula
up to 2 cm long and 1 cm wide. Apparently, this is a new species—S.
pseudoblanda Lipsch. It can also be concluded that these specimens are
of hybrid origin (S. blanda x S. ninae or S. blanda x S. coronata).
This question can be resolved only by further observations in nature. S.
konuroba Saposhn. was not compared with any species in the original
description. A study of the authentic specimen (Dzhungarian Alatau,
Konuroba River—V. Sapozhnikov and Tripolitova) revealed its complete
identity with S. blanda Schrenk.

18. S. mikeschinii I|jin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, VII, II (1937) 248.—Ie.: Iljin, Ibid (1937) 249.

Semishrubs; roots woody, fibrously branched along length, many-
headed, producing many caudices, with stems or sometimes only rosettes
of leaves. Stems 15-30 cm long; numerous, woody at base and densely
covered with brownish, weakly lustrous remains of bases of previous
year’s basal leaves, angular-sulcate, weakly pubescent, sparsely branched
(branches less numerous) or less often simple. Leaves somewhat hard,
arachnoid-hairy on both sides and with more or less numerous sessile
glandular hairs, oblong-lanceolate or lanceolate, 3-7 cm long and 0.3—
0.6 cm wide, acuminate, with light colored midrib beneath, gradually
narrowed below, and again broadened at base in short sheath, crenate-
toothed, teeth remote, terminating in short corniculate cusp, less often
almost undivided, weakly convolute, reduced toward stem apex;
uppermost leaves small, lanceolate-linear or linear, entire, acuminate,
often subtending involucre. Capitula one to three, terminal on stem and
lateral, usually short, branches. Involucre almost ovoid, about 1.2 cm
long and 0.7—0.8 cm wide, imbricate, five-rowed, weakly arachnoid-
hairy and with sparse lustrous glandular hairs; involucral bracts appressed,
with somewhat prominent midrib, dark-colored in upper part, subacute;
outer bracts ovate, middle oblong-ovate, inner lanceolate. Florets pink-
purple, about 1.3—1.4 cm long, narrow part of tube about 0.6—0.7 cm
long, broad part including lobes of limb sparsely covered with lustrous
glandular hairs, about 0.7 cm long; basal appendages of anthers woolly.
Receptacle densely covered with linear-subulate, lustrous scales, largest
scale about 5—6 mm long. Pappus white, double, outer bristles serrate-
scabrous, unequal, readily shedding, up to 4 mm long, inner ones long-
plumose, equal, about 1.1 cm long; immature achenes glabrous.
Flowering VII—VIII.

Gravelly slopes.—Soviet Central Asia: Tien Shan (Karatau Range).
Endemic. Described from the Syr-Darya Karatau. Type in Leningrad.

Note. In the original description, the plant was wrongly included

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403

under the S. alpina complex. S. mikeschinii is undoubtedly a semishrub
and belongs to the series Suffruticosae Lipsch.

19. S. ispajensis [Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, VII, 11 (1937) 250.—Ie.: Ijin, Ibid, 251.

Perennial. Root woody, fibrously branched along length, developing
more or less numerous caudices, densely covered with hardened remains
of old leaves; some caudices producing rosettes of leaves. Stems 15-30
cm long, erect, angular-suicate, glabrous or subglabrous, simple, sparsely
leafy. Leaves green on both sides, more or less scabrous due to short
scattered spines, with more prominent midrib and lateral arcuate veins
beneath; basal and lowermost cauline leaves oval or oblong, crenate-
toothed, especially in lower half of lamina, or almost entire, slightly
cuneate at base and narrowed in long narrow petioles; cauline leaves
oblong, acuminate, narrowed toward base, sessile, weakly decurrent over
stem, slightly convolute; uppermost leaves reduced, lanceolate or linear,
entire, often with rolled margin. Capitula 3—10(12), sessile, in terminal
compact corymbs. Involucre 1—1.4 cm long and 0.6—0.7 cm wide,
imbricate, four- or five-rowed, weakly arachnoid hairy-pubescent; bracts
appressed, subacute, purple-violet in upper part, outer ovate or oblong-
ovate, inner lanceolate. Receptacle covered with linear-subulate, lustrous,
unequal, up to 0.7 cm long scales; florets purple, about 1.2—1.3 cm
long, narrow part of tube about 0.5—0.6 cm long, broad part including
lobes of limb about 0.6 cm long; basal appendages of anthers comose-
woolly. Pappus dirty-smoky, double, outer pappus bristles serrate-
scabrous, unequal, fragile, up to 3 mm long, inner ones long-plumose,
about 1—1.2 cm long; achenes oblong, almost tetraquetrous, about 5
mm long and 2 mm wide, glabrous, smooth, straw-yellow, with purple
striations, and apical corona. Flowering VIII.

Soviet Central Asia: Tien Shan (west, Talass Alatau Range).
Endemic. Described from Ispai in western Tien Shan. Type in Leningrad.

Note. M.M. Iljin wrongly compared this species with 5S. mikeschinii
whereas it is much closer to S. /arionovii Winkl., from which it differs
by lacking aristate cusps on the involucral bracts and having a simple
stem, weak sinuate leaves, smoky-dirty colored pappus, etc. By
comparison with other members of series Suffruticosae Lipsch. the stems
of this species are weakly lignified at the base; however, phylogenetically
this species undoubtedly should be included here. Maybe it has been
confused with S. blanda Schrenk, from which it differs by more weakly
sinuate, less scabrous leaves, capitula densely arranged in a corymb,
and a smoky colored pappus.

Series 7. Salicifoliae Lipsch. in Bot. Mat. Gerb. Inst. Bot. Akad.
Nauk UzbSSR, XV (1959) 17.—Stems not woody at base, wingless or

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404

weakly winged. More or less numerous, short, vegetative branches
axillary (sometimes almost entirely absent). Leaves undivided, crenate-
toothed and pinnatifid, white-tomentose or green beneath; involucral
bracts obtuse or acute, but in latter case cusp straight, not recurved.

20. S. salicifolia (L.) DC. in Ann. Mus. Paris, XVI (1810) 200;
db Fi. alt. TV; 29; excl. "y. incisa Lab.; DC: Prod. Vi, 553. p: pasado:
Fl. Ross. II, 2, 670 p. p. excl. 8. ramosissima Ldb. and &. incisa Ldb.;
Turcz. Fl. baic.-dahur. II, 1, 106 p. p.; Herd. in Bull. Soc. Nat. Mosc.
XCD 3, 8 p. p.. Kryl. Fl. Zap. Sib) xl. 2923: Popov, bl. St. sstoe te
764.—Saussurea multiflora DC. in Ann. Mus. Paris, XVI (1810) 199.—
Serratula salicifolia L. Sp. pl. ed. | (1753) 817.—Heterotrichum
leptophyllum MB. ex DC. Prodr. VI (1837) 533 nomen.

Perennial. Root thick, with fibrously branched bark. Stems numerous,
up to 40(60) cm long, simple or sometimes branched from base, sulcate,
arachnoid-hairy or tomentose. Leaves linear, lanceolate-linear or oblong,
2-10 cm long (less often longer) and 2—10(15) mm wide, entire, rarely
with occasional teeth, convolute, less often flat, green, glabrous or weakly
pubescent above, white-tomentose beneath; basal and lower cauline leaves
short-petiolate, upper sessile. Capitula more or less numerous, 0.4—0.7
cm wide or larger, up to 1 cm wide (var. macrocephala Turcz.), terminal
on stem and branches in dense or slightly lax corymbs. Involucre
imbricate, arachnoid hairy-pubescent; bracts pigmented at apex; outer
bracts ovate, subobtuse, inner oblong, obtuse. Receptacle densely covered
with up to 0.8 cm long scales; florets violet pink, (13)14—23(28) in
each capitulum, narrow part of tube about 0.6—0.7 cm, broad part about
0.6 cm long including lobes of limb; appendages of anthers woolly.
Pappus white, double, outer pappus bristles persistent, serrate, unequal,
up to 4.5 mm long, inner ones plumose, about 1—1.1 cm long; achenes
brownish or olive colored, usually mottled, compressed, with indistinct
corona, mature achenes 44.5 mm long. Flowering VII—VIII.

Cryophilous steppes, steppe meadows, stony-gravelly slopes, rocks —
Western Siberia: Altai; Eastern Siberia: Angara-Sayans, Dahuria, Lena-
Kolyma (vicinity of Olekminsk). General distribution: Mongolia, ?China.
Described from Siberia. Type in Leningrad.

Note. This specimen, preserved in the herbarium of the Botanical
Institute, Academy of Sciences of the USSR, has two labels: the one
written in the hand of academician J.P. Borodin, “S. salicifolia DC. y.
major Ledeb. Gmelin,” and the other, written on an old piece of paper,
“Cirsium inerme erectum, foliis ex lineari lanceolatis infra candidis,” is
very close to Gmelin’s illustration (F/. Sibirica, II, tab. 27), which 1s
cited by De Candolle (1.c.) and, apparently, can be considered the type
or isotype of Saussurea salicifolia DC. In the work mentioned, De
Candolle gives Siberia as the habitat: “Hab. in Sibiriae apricis (v.s.).”

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The species is variable in the shape and width of the leaves, size of the
capitula and the number of florets. The infraspecific division of S.
salicifolia was first proposed by Ledebour in /cones plantarum (1829),
and subsequently in Flora Altaica (1833), later by De Candolle in the
Prodromus (1837) and by Turczanin in Flora baicalensi-dahurica (1856).
Some varieties described by these authors under different names are
identical and do not have separate ranges, while others deserve to be
segregated as separate species. Among the latter is var. incisa Ldb. (Ic.
pl. Fl. Ross. 1 (1829) 19, tab. 76; Ej. Fl. Alt. 1V (1833) 30), with more
or less deeply pinnately cut leaves, which was subsequently split off by
L.P. Sergievskaja as a separate species, Saussurea sumneviczii Serg.
The list of synonyms of the varieties of S. salicifolia is given below.

1. var. macrocephala Turcz. in DC. Prodr. VI (1837) 533; Turcz.
Fl. baic.-dahur. II, 1 (1856) 106.—S. salicifolia “e regione
transbaicalensi” Ldb. Fl. alt. IV (1833) 30.—S. salicifolia var.
transbaicalensis Ldb. ex Herd. in Bull. Soc. Nat. Mosc. XLI, 3 (1868)
8.—S. salicifolia brevifolia Turcz. in DC. Prodr. VI (1837) 533.—
Saussurea multiflora DC. in Ann. Mus. Paris. XVI (1810) 199.—
Heterotrichum leptophyllum MB. ex DC. Prodr. VI (1837) 533 nomen.—
Differs by having larger, often solitary, capitula at the apex of long
peduncles, larger number of florets, and oblong-linear leaves. It is found
scattered in the Trans-Baikal Region, mixed with the type form.

2. var. major Ldb. pl. Fl. Ross. I (1829) 19; Ej. Fl. alt. IV, 29; Ej.
Fl. Ross. II, 2, 670.—S. salicifolia var. angustifolia DC. Prodr. V1 (1837)
533; Turcz. Fl. baic.-dahur. II, 1 (1856) 107.—Characterized by entire
linear leaves, smaller capitula in numerous corymbs, and taller stems.

3. var. intermedia Ldb. Ic. pl. Fl. Ross. I (1829) 19; Ej. FI. alt. [V
(1833) 30; Ej. Fl. Ross. II, 2, 670.—Differs by having oblong-lanceolate
leaves, and shorter stems (10-15 cm high). A variety transitional to S.
sumneviczii Serg.

The specimens from Soviet Central Asia identified by many authors
as S. salicifolia mostly belong to S. elegans Ldb. var. nivea Lipsch., S.
cana Ldb., and other species. ;

The inclusion of S. salicifolia (L.) DC. in the section Lagurostemon
Cass. by M.G. Popov (Fl. Sr. Sib. Il (1959) 764) is undoubtedly wrong.

21. S. lomatolepis Lipsch. in Byull. Mosc. Obshch. Isp. Prir. Otd.
Biol. LIX, 6 (1954) 80.—S. salicifolia var. macrocephala Winkl. in
sched. Herb. Inst. Bot. Acad. Sc. USSR (non Turcz.)

Perennial. Root woody, thick, fibrously branched along length,
producing numerous stems. Stems woody at base and covered with

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406

hardened remains of old leaves,-15—35 cm high, erect, sulcate-angular,
simple or less often branched right from base, arachnoid hairy-pubescent,
glabrescent. Leaves broadly lanceolate or oblong, up to 8-10 cm long
and 1.5—2 cm wide, toothed-crenate, almost glabrous, scabrous above,
grayish arachnoid hairy-pubescent beneath with distinct midrib; basal
and lower cauline leaves short-petiolate, upper reduced, narrower, sessile;
vegetative shoots arising from axils of some leaves. Capitula 1.5—-2 cm
long (at fruiting) and about 0.8—1 cm wide, in groups of two to five in
dense corymbs at stem and branch apices. Involucre imbricate, four- or
five-rowed, pubescent; outer bracts ovate, inner oblong, all bracts
subobtuse, with distinct, narrow, dark border along margin. Receptacle
densely covered with linear-subulate, lustrous, up to 6 mm long scales;
florets dark pink, about 1.2 cm long, narrow part of tube about 0.6 cm
long, broad part including lobes of limbs covered with lustrous glandular
hairs, about 0.6 cm long; appendages of anthers flocculose-woolly.
Pappus double, outer bristles serrate, about 2 mm long, shedding, inner
ones about 1—1.1 cm long, long-plumose; achenes cylindrical, mottled,
glabrous, about 0.6 cm long. Flowering VIJ—VIII.

Described from China from the Kunges River in the Kuldzha Region.
Type in Leningrad. Probably it occurs in the USSR. It is closest to S.
salicifolia var. macrocephala Turcz., from which it differs by broader,
distinctly toothed-crenate leaves that are gray-tomentose beneath, obtuse,
narrowly dark-bordered involucral bracts and larger capitula.

22. S. cana Ldb. Ic. pl. Fl. Ross. I (1829) 18; Ej. Fl. alt. IV, 28;
DC. Prodr. VI, 533; Ldb. Fl. Ross. II, 2, 670; Lipschitz in Bot. Mat.
Gerb. Bot. Inst. Akad. Nauk SSSR, XVI, 444.—Saussurea microphylla
C.A.M. (nomen) in Ldb. et al. Reise, II (1830) 413 and herb.—Saussurea
fruticulosa Kar. and Kir. in Bull. Soc. Nat. Mosc. XIV, 3 (1841) 448.—
S. cana B. angustifolia Ldb. Fl. Ross. II, 2 (1845-1846) 670.—Ic.: Ldb.
op. cit. (1829) tab. 75.

Perennial. Root woody, brown, with fibrously branched bark, many-
headed at apex; caudices sometimes covered with loose turf and bearing
floriferous shoots or terminating in rosettes of leaves. Stems 10—15(25)
cm high, erect, weakly ascending, simple, with somewhat numerous,
erect floriferous branches only above, slightly sulcate-angular, gray-
pubescent. Leaves grayish tomentose on both sides; basal leaves usually
withering before flowering; leaves of sterile rosettes oblong, more or
less deeply pinnatipartite or almost entire, acuminate, weakly convolute,
gradually attenuate toward base, short-petiolate, sheathing; cauline leaves
linear, entire, narrow (var. angustifolia Ldb.) or more or less pinnatipartite
in type form; all leaves gradually reduced toward stem apex, usually
sessile, somewhat semiamplexicaul; upper leaves smail, entire. Capitula
cylindrical, 4-20, erect, short-pedunculate, in simple or usually compound

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407.

terminal corymb. Involucre imbricate, five-rowed, grayish arachnoid-
hairy, 1—1.3 cm long, bracts appressed, red-colored, usually entirely or
at apex and along margin, with one distinct vein, sometimes covered by
pubescence; outer bracts ovate, roundish at apex, with indistinct short
teeth; inner bracts oblong, subacute. Receptacle densely covered by
linear-subulate, white, lustrous, narrow, up to 0.8 cm long scales; florets
pink, about 1.2-1.3 cm long; narrow part of tube about 0.6 cm long,
broad part with occasional lustrous glandular hairs, with lobes of limb
about 0.6-0.7 cm long; basal appendages of anthers flocculose-woolly.
Pappus double, white, outer pappus bristles serrate, fragile, about 2-3
mm long, inner ones long-plumose, about 1—1.1 cm long; achenes narrow
cylindrical, mature achenes up to 0.7 cm long, glabrous. Flowering VII—
IX.

White clays, marl, limestones, shales, rock outcrops, stony steppes.—
Western Siberia: Upper Tobol (southern part), Irtysh, Altai; Eastern
Siberia: Angara-Sayan (?); Soviet Central Asia: Aralo-Caspian Region,
Balkhash Region, Dzhungaria-Tarbagatai. General distribution:
Dzhungaria-Kashgaria (Kunges River Valley in Xinjinang), Japan, China
(Shangshi and Kangsu provinces). S. cana Ldb. is described from
Kazakhstan (vicinity of Karakalinsk). Type in Leningrad; S. fruticulosa
Kar. and Kir. was described from Uzbekistan (Tarbagatai). Type -in
Leningrad.

Note. Keller’s specimens from the Kaldzhir Valley (Aine-Bulak,
1908) differ by having (up to 0.6 cm wide) lanceolate or oblong-linear
leaves and deserve to be segregated as a separate variety (var. Jatifolia
Lipsch.). Strangely, S. cana Ldb. was omitted from the Flora of Western
Siberia. Typical specimens of S. cana are characterized by narrowly
cylindrical capitula, gracefully erect, broadly ovate-roundish, obtuse
involucral bracts terminating in a short barely noticeable cusp
(submucronulate), and leaves that are grayish-tomentose on both sides
(in S. salicifolia, with which this species is often confused, they are
green and scabrous above, and white-tometnose beneath), usually oblong-
linear or oblong, entire, and more or less convolute (leaves slightly
pinnately incised, dentate). S. cana is phylogenetically very close to S.
sumneviczii Serg., from which it differs by having leaves that are entire
or slightly divided and grayish-tomentose on both sides (in S. sumneviczii
Serg., they are green and scabrous above, and white-tomentose beneath,
deeply pinnately cut), and capitula that are erect and usually densely
arranged. For the relationship between S. cana and S. caespitans lin,
cf. the note to the latter.

23. S. sumneviczii Serg. in Sistemat. Zam. Gerb. Tomsk. Univ. 1—
2 (73-74) (1949) 12; Kryl. Fl. Zap. Sib. XI, 2924:—S. salicifolia y.
incisa Ldb. Ic. pl. Fl. Ross. I (1829) 19; Ej. Fl. alt. 1V, 30; DC. Prodz.

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408

VI, 533; Ldb. Fl. Ross. II, 2, 671; Kryl. Fl. Alt. III, 709.—S. squarrosa
Fisch. (non Turcz.) ined. ex Ldb. Fl. Ross. II, 2 (1845-1846) 671.—
Saussurea Salicifolia var. squarrosa Bge. ex Herd. in Bull. Soc. Nat.
Mosc. XLI, 3 (1868) 9 nomen.—Ie.: Ldb. op. cit. (1829) tab. 76.

Perennial. Root many-headed, fibrously branched, usually producing
many stems; root collar covered with hardened remains of basal leaves.
Stems numerous, less often solitary, 1.5—-10 (up to 45) cm high, simple
or branched in inflorescence, grayish or white-tomentose. Leaves variable
in shape and degree of incision, in typical form (1)2—5 cm or longer, 6—
14 cm, oblong, glaucous-green, weakly stellate-hairy, and with arachnoid
hairy-floccose, lustrous glandular hairs, or almost glabrous and green
above, white-tomentose beneath; all leaves in type form more or less
deeply pinnatilobate or crenate-dentate, 0.1—-1 cm wide in undivided
part; some of them sometimes undivided or weakly toothed; lower cauline
leaves short-petiolate, base of petiole of basal leaves slightly sheathing;
upper leaves sessile, not decurrent. Capitula two to nine, compactly
arranged at stem and branch apices, less often sparse. Involucre imbricate,
3-5(7) mm wide; bracts reddish or violet, weakly arachnoid hairy-
pubescent, especially along margin and apex, often slightly glandular
dorsally; outer bracts smaller, ovate, often with prominent midrib,
subobtuse or short-acuminate, inner bracts oblong, subobtuse. Florets
violet-pink, about 1.2—1.3 cm long; narrow part 0.5 cm long, broad part
sometimes with lustrous glandular hairs, about 0.7 cm long including
lobes of limbs; appendages of anthers comose-lanate. Receptacle densely
covered with linear-subulate, lustrous, up to 0.7 cm long scales; pappus
snow-white, outer pappus bristles serrate-scabrous, fragile, about 3 mm
long, inner bristles long-plumose, about 1 cm long; achenes 3-4 mm
long, mottled, mature achenes dark violet. Flowering VIJ-—VIII.

Stony and gravelly slopes and plains, rocks, clayey outcrops.—
Western Siberia: Altai; Eastern Siberia: Angara Sayans (southern part
of Tuva Republic). General distribution: Mongolia. Described from Altai.
Type in Tomsk; isotype in Leningrad. S. salicifolia y. incisa Ldb. was
described from Chuya steppes. Type in Leningrad.

Note. This is a polymorphic species with transitions to S.
serratuloides Turcz. and S. salicifolia (L.) DC. L.P. Sergievskaja
recognized several varieties: var. communis Serg.: “... Stem 10-20 cm
long; all leaves deeply pinnately incised over a larger part up to half,
white-tomentose beneath, 6-10 cm long, in undivided part 2-4 mm broad;
lobes or their teeth 1-9 mm long and 1-3 mm wide remote”; var.
mutabilis Serg.: “... Stem 13-45 cm long; leaves grayish to grayish-
white beneath, pinnatilobate or crenate-toothed; upper leaves often entire,
lobes 2.5—-4 mm long and 4—7 mm wide”; this variant is a transition to
S. serratuloides Turcz. To the varieties proposed by L.P. Sergievskaja
in S. sumneviczii must be added one more new variety, var. sergievskajae

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409

Lipsch.: stem up to 25 cm long; basal leaves up to 14 cm long, cauline
leaves 5—7 cm long, up to 1 cm wide, with numerous remote small
teeth along margin, some leaves almost entire; veins, especially in basal
leaves, prominent beneath; vegetative shoots arising from axils of cauline
leaves (Chuya steppe). S. sumneviczii is closest to S. serratuloides Turcz.,
but the leaves of the latter species are pinnately cut almost up to the
middle into linear or lanceolate lobes, 1.5—4 cm long and 2—5 mm wide;
stem strongly branched above; involucral bracts almost glabrous. On
the other hand, S. sumneviczii is very similar to S. cana Ldb., but the
latter is characterized by leaves grayish-tomentose on both sides, which
in the type form are entire and narrower. Moreover according to Ledebour
(FI. alt. IV, 30), the florets of S. cana Ldb. are larger, and the involucral
bracts grayish-tomentose, not villous. Sergievskaja did not compare her
species with S. cana Ldb.; apparently, it was unknown to her.

24. S. serratuloides Turcz. in Bull. Soc. Nat. Mosc. XX, 2 (1847)
35.—S. gebleriana C.A.M. in Mém. Acad. Pétersb. sér. VI, VII (1855)
6; Kryl. Fl. Zap. Sib. XI, 2925.

Perennial. Roots fibrously and transversely branched, producing few
stems; root collar covered with dark brown, weakly glossy sheaths of
basal leaves. Stems more or less flexuous, in type form straight, 20—60
cm long, angular-ribbed, glabrous or grayish-tomentose, branched,
branches more or less bent. Leaves numerous, up to 13 cm long and up
to 4 cm wide; basal leaves petiolate, upper sessile, adnate with stem by
narrow base, but not decurrent; all leaves green, glabrous, weakly
scabrous above due to spinules, white-tomentose beneath, oblong, with
prominent midrib, deeply pinnatipartite; middle undivided part 0.2—0.4
cm wide, somewhat toothed between lobes, lobes incurved, almost linear
or lanceolate, 0.2-0.5 cm wide, acute, convolute; lateral lobes longer,
almost entire or more or less pinnatilobate; lower and upper lobes one-
third to half as long as middle, entire or toothed; uppermost and branch
leaves smaller, toothed or pinnatipartite in middle. Capitula cylindrical,
5—10, on short peduncles terminal on stem and branches, in almost
corymbose inflorescence; corymbs, in turn, forming compound, almost
paniculate inflorescence in several specimens. Involucre imbricate, 1—
1.4 cm long and 0.4—0.5 cm wide; bracts unequal, green or violet, almost
glabrous or less often very weakly pubescent, with weakly noticeable
veins; outer bracts small, ovate, with small cusp, middle oblong, longer,
subobtuse or subacute; inner bracts almost linear, obtuse. Receptacle
covered with linear-subulate up to 0.8 cm long, white, lustrous scales;
florets pink, about 1.3 cm long, narrow part of tube 0.5—0.6 cm long,
broad part about 0.7 cm long including lobes of limb; appendages of
anthers comose-woolly. Pappus snow-white, double, outer pappus-bristles
serrate-scabrous, fragile, unequal, about 4-5 mm long, inner ones long-

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410

plumose, about 1—t.1 cm long; achenes about 0.4 cm long, mottled,
ribbed, with a crown. Flowering VII-—VIII.

Western Siberia: Altai. Endemic. Described from western Altai, from
Topolovka. Type in Leningrad.

Note. This species is closest to S. sumneviczii Serg., from which it
differs by larger leaves which are deeply divided into lanceolate or linear
lobes and taller, strongly branched stem.

During a study of S. sumneviczii Serg. it was found that some
specimens from Altai (Oirotiya, Chuya River Valley) should be referred
to S. serratuloides; others are .intermediate between these species.

25. S. caespitans Iljin in Bot. Mat. Gerb. Bot. Sada, IH, 26 (1922)
102; Lipschitz in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI,
445.

Perennial. Caespitose; roots long, strongly fibrously branched, many-
headed. Stems numerous, reduced, slender, angular-ribbed, arachnoid,
usually densely leafy, covered at base with numerous, brown, hard
remains of previous year’s leaves. Leaves 2.5—3.5 cm long, 0.5—0.8 cm
wide, lanceolate, pinnatifid, teeth remote, with short callose cusp, leaves
gradually narrowed toward base in flattened petiole, white-tomentose
beneath, grayish-green above, more or less pubescent; uppermost leaves
reduced, entire. Capitula cupuliform, 1.5—2 cm long and 0.8—1 cm wide,
numerous, pedunculate on stem apices forming (two)three to five-headed,
weakly spreading corymbs. Involucre strongly arachnoid-hairy,
glabrescent at places, imbricate, five-rowed; bracts dark-colored, subacute,
with distinct midrib, usually covered by pubescence; outer and middle
bracts ovate or ovate-oblong, innermost linear, narrower. Receptacle
covered with linear-subulate, white, lustrous, unequal scales, longest about
5 mm long; florets .1.4—1.5 cm long, narrow part of tube about 0.6—0.7
cm, broad part 0.7—0.8 cm long including lobes of limb; appendages of
anthers comose-woolly. Pappus double, outer pappus-bristles serrate, less
numerous, unequal, up to 3 mm long, fragile, inner ones plumose, 1—
1.1 cm long; achenes 0.5—0.6 cm long, narrow, glabrous, smooth.
Flowering VII-VIII.

Outcrops of red clays, stony and gravelly slopes—Soviet Central
Asia: Tien Shan (region of Sarydzhas in Tekesa Valley). Endemic.
Described from Sarydzhas. Type in Leningrad.

Note. M.M. Ijin did not compare S. caespitans with any species
of the S. salcifolia group in which it was included. The following features
are characteristic for C. caespitans: formation of dense tufts, pinnatifid
leaves, usually pubescent on both sides (more strongly beneath);
cupuliform, rather large capitula; dark-colored, strongly arachnoid-hairy,
somewhat acuminate involucral bracts.

411

From S; cana Ldb., especially the forms with incised leaves, S.
caespitans differs by broader, more deeply incised, lanceolate (not
apically roundish), dark-colored, strongly pubescent leaves, the
pubescence often covering the prominent midrib; capitula larger,
somewhat scattered, cup-shaped (in S. cana they are narrowly cylindrical,
gracefully erect); involucral bracts (even the outermost), somewhat
acuminate. From S. sumneviczii Serg. it differs by the leaves that are
strongly pubescent on both sides in typical specimens (in S. sumneviczii,
they are usually green, almost glabrous above), larger cupuliform capitula,
strongly pubescent involucral bracts with a distinct cusp, and also by
forming dense tufts. After the description of S. caespitans M.M. Ijin
cites specimens: “Semireche Province, Dzharkent district, Sary-Dzhas,
27.V1.1910, No. 1650, A.I. Michaelson”; this is the type of the species.
The second ones cited by Ijin, “Tarbagatai, Chingiz Mountains,
10. VIII.1909, No. 1094, R. Yu. Rozhevits,” in my opinion, do not belong
to S. caespitans, but are S. cana Ldb. (their leaves are undivided, linear-
oblong; the capitula smaller, gracefully erect; the involucral bracts
roundish at the apex, subobtuse).

26. S. canescens Winkl. in Tr. Peterb. Bot. Sada, XI, 1 (1889). 168.

Perennial. Roots woody, fibrously branched along length, with more
or less numerous caudices, covered with remains of previous year’s leaves
and producing solitary floriferous shoots; entire plant grayish. Stems
ascending 1.5—4 cm long (about 10-12 cm high in var. major Wink.
(op. cit. 168)), canescent, sometimes glabrescent, simple, one-headed or
three-headed when stem weakly branched above, leafy. Leaves
coriaceous, about 1.5—3 cm long (3—7 cm long in var. major Winkl.),
pinnatifid or crenate-lobate, lobes subacute, with callose spine at apex
(under lens); basal and lower cauline leaves petiolate; upper leaves
reduced, narrowed below, almost sessile, scarcely amplexicaul, not
decurrent, all leaves canescent on both sides. Capitula one to three on
each stem, in latter case corymbs compact since capitula short-
pedunculate and weakly drooping, 1.5 cm long and 0.7—1 cm wide.
Involucre imbricate, five-rowed, grayish arachnoid hairy-pubescent,
glabrescent; bracts light colored or dark in upper part, distinctly
acuminate, especially inner (cusp often with tuft); outer bracts ovate,
inner oblong or oblong-lanceolate; receptacle covered with scales. Florets
pink, about 1.2 cm long; narrow part of tube about 0.5 cm long, broad
part with lobes of limb about 0.7 cm long; appendages of anthers comose-
woolly; pappus double, white, outer pappus bristles serrate, fragile, about
3 mm long, inner ones long-plumose, about 1.1 cm long; achenes
(immature) about 0.3 cm long, with an indistinct corona. Flowering VII—
VU.

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412

So far not found in the USSR. Described from Kuldzha Region
(from the alpine zone of Kokkamyr and Lake Sayram-Nor). Type in
Leningrad. Its occurrence in the border areas of the USSR is quite
probable.

Note. K. Winkler groundlessly compared S. canescens with S.
salicifolia, with which it has no connection. Nevertheless, in the same
publication, Winkler described another new species S. salemannii Winkl.
also from the Kuldzha Region, to which he should have compared his
S. canescens, especially its var. major Winkl. These two species (S.
canescens and S. salemannii) are geographically and phylogenetically
extremely close. Until more material is available for these species, they
can be distinguished as separate species by the following characters: S.
canescens is a shorter, grayish-pubescent plant, one- to three-headed,
the capitula being 0.7—1 cm wide (in S. salemannii they usually are
wider); the leaves are grayish-pubescent on both sides and barely scabrous
(whereas the leaves of S. salemannii are not grayish-pubescent and are
quite strongly scabrous on both sides). For the relationship between S.
canescens and S. vvedenskyi, cf. the note to the latter species.

27. S. vvedenskyi Lipsch. in Bot. Mat. Gerb. Inst. Bot. Akad. Nauk
UzbSSR, XV (1959) 16.

Semishrub. Roots woody, fibrously branched, many-headed, bearing
numerous stems and rosettes of leaves, forming turf. Stems 7-11 cm
high, 1-1.5 mm wide, angular, upright or slightly arcuate, pubescent,
strongly leafy, terminating in solitary or less often two capitula; old
leaves remain at base for long along with imbricate remains of previous
year’s leaf sheaths. Leaves narrow-lanceolate or linear-lanceolate, entire
or indistinctly toothed, canescent on both sides, with prominent midrib;
basal leaves 4—5.5 cm long and 0.5—0.7 cm wide, middle cauline and
upper leaves reduced, sessile, capitula campanulate, one, less often two
or three at stem apex, about 1.5-1.7 cm long and 1-1.1 cm wide.
Involucre imbricate; bracts three- or four-rowed, almost carinate, more
or less pubescent; outer and middle bracts ovate, inner oblong; all bracts
violet, with short cusp, sometimes weakly recurved; receptacle covered
with linear-subulate scales. Florets pink, about 1.5 cm long; corolla tube:
about 0.8 cm long, campanulate, limb five-fid, about 0.7 cm long, with
sessile, scattered, lustrous glandular hairs; appendages of anthers comose-
woolly. Achenes cylindrical, with short corona at apex, glabrous, about
5 mm long, with four whitish ribs; pappus double, outer pappus bristles
about 2 mm long, fragile, scabrous, inner ones about 1.1 cm long, soft,
silky, plumose. Flowering VII—VIII.

Stony and gravelly slopes.—Soviet Central Asia: Tien Shan
(western). Endemic. Described from Akbashtau Mountain. Type in
Tashkent.

413

\
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Plate XXXII.
Saussurea kurilensis Takewaki; 2—S. riederi Herd.

419

1

42]

414

Note. This species is close to S. canescens Winkl. but differs by
the distinctly semishrub habit, form of the leaves which are narrowly
lanceolate or linear-lanceolate, and entire or indistinctly toothed (and
not crenate-lobate or almost pinnatifid), taller, more strongly leafy stem,
and so on.

28. S. salemannii Winkl. in Tr. Peterb. Bot. Sada, XI, 1 (1889)
166; Lipschitz in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XXI,
371.—S. salicifolia var. viridifolia Winkl. in sched. and ex O. and B.
Fedtsch. Perech. Rast. Turk. IV (1911) 230.

Perennial. Entire plant strongly scabrous; roots woody, fibrously
branched along length, many-headed. Stems 13-25 cm long, covered
with hardened remains of leaves at base, scabrous due to scattered soft
bristles, sometimes almost glabrous, sulcate, usually branched. Leaves
green on both sides, in type form strongly scabrous due to numerous
bristles all over leaf, especially along margin; basal and lower cauline
leaves petiolate, up to 7-8 cm long and 1.5—2 cm wide; lamina obovate
or broadly lanceolate, more or less crenate-lobate, lobes roundish or
subacute, or less often sinuate-pinnatifid; cauline leaves like basal sessile,
weakly amplexicaul, not decurrent on stem or short-petiolate; uppermost
cauline leaves reduced, toothed or entire. Capitula about 2 cm long and
1 cm wide, compactly borne on short peduncles, forming corymbs,
sometimes solitary on lateral, rather long, curved floriferous shoot.
Involucre imbricate, four- or five-rowed, appressed, pubescent; outer
bracts ovate, inner oblong or oblong-lanceolate; all bracts with more or
less distinct midrib, attenuate into short cusp; receptacle covered with
stiff scales, as long as inner bracts. Florets pale pink, about 1.4—1.5 cm
long; narrow part of tube almost as long as broad part including lobes
of limb; lower appendages of anthers comose-lanate. Pappus double,
outer bristles fragile, almost as long as achene, scabrous, inner white,
about 1.3 cm long, plumose, two or more times as long as outer bristles;
achenes 0.4—0.5 cm long, glabrous, mottled, finely ribbed, smooth along
ribs. Flowering VII—VIII. (Plate XXX, Fig. 1.)

Soviet Central Asia: Dzhungaria-Tarbagatai (found only once in the
Chulak area). Probably this species also occurs in other parts of the
USSR adjoining China.

Described from China (vicinity of Lake Sayram, Talki River and
Talkibash Mountain). Type in Leningrad.

Note. K. Winkler, absolutely without justification, compares this
species with S. elegans Ldb., with which, in my opinion, it has no
connection, except for scabrous leaves, either in the habit of the plant,
size of the capitula, or form of the leaves. I also include under S.

422 salemannii Winkl. the specimens of Regel from Upper Borotal (1878), —

423

415

identified by Winkler as S. salicifolia var. viridifolia Winkl. The
specimens from the Dzhungarian Alatau (Chulak area, S. Lipschitz) differ
slightly from the type specimen by having thinner and weakly scabrous
(almost smooth) leaves. In all other respects, they are identical with the
type of S. salemannii Winkl., to which I refer them.

29. S. elata Ldb. Ic. pl. Fl. Ross. I (1829) 20; Ej. Fl. alt. IV, 33;
DE ProdreVi532)idbs Fly Ross. Ws 23672; Kayli Fly Alt Il: 709;
Kryl. Fl. Zap. Sib. XI, 2925.—Ic.: Ldb. op. cit. (1829) tab. 80.—Exs:.:
Kar. and Kir. (1840) No. 835.

Perennial. Stem fistular, erect, tall, 1-2 m high and 1 cm thick,
ribbed-angular, ribs often reddish, especially in upper part, stem weakly
pubescent and covered with sessile lustrous glandular hairs, glabrescent,
simple below, branched from middle or above, branches upward
spreading. Basal leaves usually withering before blooming, oblong-
elliptical, acuminate, narrowed into petiole, almost entire or unequally
large-toothed; middle cauline leaves hard, lanceolate, acute, entire,
narrowed toward base, sessile or almost sessile, not decurrent on stem,
6—15 cm long and 1-4 cm wide, on both sides covered with short, stiff,
cilia, besides with lustrous glandular hairs beneath; upper cauline leaves
reduced, linear lanceolate. Capitula numerous in dense corymbs at stem
and branch apices, forming long and spreading corymbose-paniculate
inflorescence. Involucre imbricate, 1—1.2(1.5) cm long and 0.5—0.7 cm
wide, mouldy pubescent, with or without sessile glandular hairs; outer
involucral bracts ovate or oblong-ovate, subobtuse; inner bracts oblong,
obtuse; all bracts greenish or pigmented in upper half as well as along
margin, with more or less distinct midrib, covered with pubescence over
larger part of inner bracts; receptacle densely covered with linear-
subulate, small, about 3 mm long scales. Florets violet-lilac, about 1.4
cm long; narrow part of tube about 0.7 cm long; broad part with lobes
of limb about 0.7 cm long; appendages of anthers comose-woolly. Pappus
double, outer pappus-bristles serrate-scabrous, fragile, about 3 mm long,
inner ones long-plumose, about 0.8—1 cm log; achenes smooth,
compressed, about 0.7—0.8 cm long, almost equaling pappus. Flowering
VII-VIII.

Thickets of steppe shrubs, steppe meadow, sands, steppes with Stipa
lessingiana.—Western Siberia: Irtysh from foothills of the Altai from
the Bukhtarma River (between Ust-Kamenogorsk and Bukhtarminsk).
Endemic. Type in Leningrad.

Note. Albino specimens (f. albiflora Lipsch.) are found, for example,
in the former Zaisan district.

30. S. elegans Ldb. Ic. pl. Fl. Ross. I (1829) 19; Ej. Fl. alt. IV, 31;
DC. Prodr. VI, 533; Ldb. Fl. Ross. IJ, 2, 671; Kryl. Fl. Zap. Sib. XI,

416

2926.—S. elegans var. latifolia Kar. and Kir. in Bull. Soc. Nat. Mosc.
XV, 2 (1842) 388.—S. salicifolia var. elegans Trautv. Ibid, XXXIX, 2
(1866) 370.—S. tenuis Ldb. Ic. pl. Fl. Ross. I (1829) 19; Ej. FI. alt. IV,
31; DC. Prodr. VI, 533; Ldb. Fl. Ross. II, 2, 671—S. elegans var.
tenuis (Ldb.) Serg. in Kryl. Fl. Zap. Sib. XI (1949) 2927.—S. amoena
Kar. and Kir. in Bull. Soc. Nat. Mosc. XIV, 3 (1841) 447.—S. elegans
var. amoena (Kar. and Kir). Serg. op. cit. (1949) 2927.—S. salicifolia
var. ramosissima Ldb. Fl. Ross. II, 2 (1845-1846) 670.—S. salicifolia
var. stenocephala Trautv. in Bull. Soc. Nat. Mosc. XXXIX, 2 (1866)
369 p. p. excl. syn.—sS. salicifolia var. scabra Rgl. and Schmalh. in Tr.
Peterb. Bot. Sada, VI, 2 (1880) 313.—S. salicifolia auct. fl. Turkest. p.
max. p.; O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 230 p. p.,
non DC.—Iec.: Ldb. Ic. pl. Fl. Ross. I, tab. 77 and 78.—Exs.: Kar. and
Kir. (1840) No. 834; Ej. (1841) No. 1629.

Perennial. Roots fibrously branched along length. Stems several,
30-60(80) cm high, erect, angular-ribbed, often glabrous below, lustrous,
usually slightly arachnoid-hairy with scattered lustrous glandular hairs,
densely leafy (in ssp. turkestanica Lipsch. weakly leafy); more or less
numerous short shoots in leaf axils (ssp. turkestanica Lipsch. without
vegetative shoots in leaf axils). Leaves scabrous above and along margin
due to short spiny bristles, grayish arachnoid hairy-floccose beneath,
sometimes distinctly white-tomentose (var. nivea Lipsch. in Bot. Mat.
Gerb. Bot. Inst. Akad. Nauk SSSR, XXI (1961) 371) or almost glabrous,
green, usually covered with sessile lustrous glandular hairs; basal leaves
oblong or oblong-ovate, pinnatilobate or almost lyrate-pinnatipartite, up
to 15 cm long, 1.5 to 3(4) cm wide in middle, petiolate, petioles two-
fifths to half as long as lamina; usually plants withering before flowering;
cauline leaves numerous (in ssp. turkestanica Lipsch. less numerous and
sparse), ovate or broadly lanceolate, sessile, entire, often some, usually
in lower half of lamina, crenate-lobate or toothed, all leaves acuminate,
narrowed toward base, sometimes weakly decurrent (in ssp. turkestanica
Lipsch.), 3—10 cm long, 0.5—2.5(3) cm wide; uppermost leaves reduced,
narrower, linear-lanceolate, 0.3—0.8 cm wide. Capitula cylindrical, 1.5—
2 cm long and 0.5—0.8 cm wide, usually numerous, in broad corymbose-
paniculate inflorescence (in ssp. turkestanica Lipsch. inflorescence poorly
developed, usually terminal compact corymb; less often, besides terminal
many lateral corymbs also present). Involucre imbricate; bracts somewhat
arachnoid hairy-pubescent, with scattered lustrous glandular hairs, green,
sometimes entirely or partially dark-colored; bracts of two or three outer
rows ovate, subobtuse or cuspidate (in ssp. turkestanica Lipsch., outer
as well as inner bracts more acuminate, often with a short blackened
cusp); receptacle covered with linear-subulate, lustrous, up to 0.7 cm
long scales. Florets purple-pink, about 1.3—-1.4 cm long, narrow part of

417

tube about 0.8 cm long, broad part with lobes of limb covered with
glandular hairs, about 0.6 cm long; anthers exserted, about 0.5—0.6 cm
long; their appendages comose-woolly. Pappus white, double, outer
pappus-bristles serrate, fragile, about 3-4 mm long, inner ones plumose,
about 1 cm long; achenes about 0.5 cm long, cylindrical, mottled due
to longitudinal violet stripes, glabrous. Flowering VII—VIII.

Meadow, shrub and sheep fescue steppes, stony slopes.—Soviet
Central Asia: Aralo-Caspian Region, Syr-Darya, Balkhash Region, Tien
Shan (predominantly ssp. turkestanica Lipsch.), Dzhungaria-Tarbagatai,
Pamiro-Alai (predominantly ssp. turkestanica Lipsch.); Western Siberia:
Irtysh (eastern part).

S. elegans Ldb. is described from the Chingistau Mountain. Type
in Leningrad. S. tenuis Ldb. is described from the Arkat Mountains.
Type in Leningrad. S. amoena Kar. and Kir. is described from Chorga
Creek, draining into Lake Nor-Zaisan. Type in Leningrad; isotype in
Moscow.

Note. A quite polymorphic species with extremely unstable
characters. The original diagnosis of the species by Ledebour says: “Folia
supra scabra subtus tomento arachnoideo incana.” Specimens with snow-
white tomentum on the lower suface of the leaves (this is var. nivea
Lipsch.) are found throughout the range of this species. Apparently, this
type of pubescence, running the gamut of transitions to green, almost
glabrous or grayish lower leaf surface due to arachnoid-floccose
pubescence, is a consequence of ancient hybridization between S. elegans
and other species (S. salicifolia, S. cana) with a white-tomentose lower
leaf surface. I do not see a basis for segregating S. amoena Kar. and
Kir. as a separate species. The distinctive features of this species are
not constant. These are: 1) stems branched from base to top (that is
explained by the fact that the specimens used by Karelin and Kirlov to
describe their species were cattle-grazed and then grew again with profuse
branching; 2) broadly lanceolate leaves, 2.5-5.5 cm long and 0.8—1 cm
wide (they are also characteristic for several forms of S. elegans); and
3) presence of more or less numerous vegetative shoots in the leaf axils
(which is also observed in typical S. elegans).

Typical S. tenuis Ldb. is distinguished by greater leafiness and more
numerous vegetative shoots in the leaf axils, leaves that are glabrous on
the lower suface and are usually narrrower, with slightly involute margins,
and more strongly crenate-lobate (their lobes sometimes elongate, acute),
and by compact corymbs of smaller capitula. None of these characters
is constant, since in S. elegans specimens also are found that are
characterized by a more or less strongly leafy stem, numerous, short,
vegetative shoots in the leaf axils, pubescence lacking on the lower leaf
surface (contrasting with the distinct snow-white tomentum here in var.

425

418

nivea Lipsch.), variable cutting and size of the leaves, compact and
poorly developed corymbs, as well as by the size of the capitula. For
this reason, I am combining S. tenuis Ldb. with S. elegans Ldb. Albino
specimens of S. elegans f. albiflora Lipsch. occur.

31. S. larionovii Winkl. in Tr. Peterb. Bot. Sada, XI, 2 (1891)
376.—S. takhtadjanii Lipsch. in Byull. Mosk. Obshch. Isp. Prir., Otd.
Biol. LIX, 6 (1954) 83.

Perennial. Root woody, fibrously branched. Stems solitary or few,
(10)15—25(30) cm long, upright or slightly ascending, angular-sulcate,
canescent, sometimes almost glabrous, simple or branched, usually
weakly winged. Leaves quite variable in shape and width, lanceolate,
oblong-lanceolate, linear-lanceolate, in type form distinctly crenate-
toothed or pinnatifid, often entire, strongly scabrous above and along
margin due to numerous spiny bristles, more or less grayish beneath
due to arachnoid-hairy pubescence or white-tomentose (var. nivea
Lipsch.), less often almost glabrous with distinct midrib, merging in a
weakly wingled petiole in basal leaves; basal leaves sometimes early
withering; lower cauline leaves 5—10 (less often more) cm long including
petiole, (0.5)1—1.5 cm wide, petiole (1)2—3(5) cm long; middle and upper
cauline leaves reduced, sessile, semiamplexicaul and more or less
decurrent, as a result stems sometimes somewhat winged. Capitula 1.2—
1.4(1.6) cm long, (0.3)0.5(0.7) cm wide, narrow cylindrical, in compact,
erect corymbs; sometimes, when stem branched, few lateral corymbs
besides central corymb borne in corymbose-paniculate inflorescence.
Involucre withered, four- or five-rowed, reddish, sometimes green,
arachnoid-hairy, glabrescent; outer bracts ovate, inner oblong-lanceolate;
all bracts, especially inner acuminate, with distinct midrib, attenuate in
aristate straight or weakly curved cusp, sometimes dark-colored, readily
breaking, with hyaline hairs along margin and apex on outer as well as
inner sides. Receptacle covered with long scales, scales subulate, three-
fourths as long as inner involucral bracts; lustrous; florets pink, up to
1.4 cm long, narrow part of tube about 0.7 cm long, as long as broad
part of tube including lobes of limb, limb with sessile, lustrous glandular
hairs; appendages of anthers floccose-lanate. Pappus double, outer pappus
bristles short, up to 2—2.5 mm long, fragile, scabrous-serrate, inner bristles
plumose, about 1.1—1.2 cm long; achenes glabrous, with short corona.
Flowering VII-—VIII.

Stony slopes, outcrops of red clays and shales, shore eels saline
meadows, Lasiagrostis complex.—Soviet Central Asia: Tien Shan (valleys
of Kegen, Sarydzhas, Tekes, and other rivers). General distribution:
Dzhungaria-Kashgaria. Endemic. Described from China, from the Great
Yulduz Range and from the Kharsal River. Type in Leningrad.

419

Note. The plant was described by K. Winkler from specimens
without fully developed capitula. The main character given by Winkler
as distinguishing this species from other “narrow-leaf forms of S.
salicifolia” is the presence of distinct appendages on the involucral bracts.
Winkler wrote: “All Saussurea with involucral bracts having apical
appendages are distinguished by more or less roundish, and not linear-
lanceolate, appendages.” In this Saussurea by “appendage” Winkler
meant a sinuate or straight cusp, which is the continuation of the midrib
of the involucral bract. This cusp is aristate, usually without pigmentation,
with hyaline cilia along the margin and at the apex; it breaks off easily.
Without any hesitation, I include under S. larionovii the Saussurea
specimens from the central Tien Shan (Sarydzhas and Kegen River
valleys). These specimens differ somewhat from the authentic specimens
of S. Jarionovii originating from the Kharsal River in the region of the
Great Yulduz, where they were collected by Larionov. Our specimens
differ from the authentic ones by a longer stem, distinctly dentate-crenate
or pinnatifid leaves (this tendency is also readily observed in the type
specimens), a wider and distinct branching of the stem and, consequently,
the formation of paniculate inflorescence in some specimens. However,
according to my broad interpretation of the species, they undoubtedly
should be referred to S. larionovii, given the presence of the aristate
cusp on the involucral bracts, the more or less winged stems, and the
shape of capitula. In 1954, I described S. takhtadjanii, from material
collected outside the USSR (the former Chinese Turkestan—Muzart,
Maral Bashi; type in Leningrad). I pointed out that this species is closest
to S. larionovii Winkl., but differs from the latter by having a wingless,
strongly branched stem with numerous, erect corymbs of capitula, and
broader leaves, crowded at the base of the stem, where they form rosette-
like structures. The revision showed that S. takhtadjanii should be
relegated to synonymy under S. larionovii. S. larionovii apparently is
closest to S. alberti Rgl. and Winkl. It differs from the latter by leaves
that are grayish-tomentose beneath, erect corymbs of capitula, number
of florets in a capitulum (12-13, and not 19-20), and more acute
involucral bracts.

32. S. alberti Rgl. and Winkl. in Tr. Peterb. Bot. Sada, VI, 2 (1880)
298.

Perennial. Stem sulcate, weakly winged. Basal leaves petiolate,
elliptical-lanceolate or oblong-lanceolate, acute, crenate-dentate, weakly
cuneate toward base, glabrous on both sides, scabrous along margin due
to small bristles that are visible under a lens, up to 7 cm long, excluding
petiole, and 2—2.75 cm wide; cauline leaves narrower, usually entire,
glabrous, decurrent on stem as wing. Capitula with 19—20-florets,

427

420

numerous, pedunculate in corymbs at the apices of bent floriferous
branches. Involucre almost cylindrical; bracts imbricate, appressed, acute,
arachnoid-hairy, glabrescent; outer bracts ovate, middle oblong-lanceolate;
inner bracts almost linear; receptacle covered with unequal, up to 0.8
cm long scales. Florets pink, about 1.3 cm long, narrow part of tube
0.6—-0.7 cm long, as long as its broad part including lobes of limb;
appendages of anthers comose-woolly. Pappus double, outer pappus-
bristles scabrous, small, inner ones plumose, four times as long as outer
ones; achenes oblong, angular, with short apical corona.

Soviet Central Asia: Tien Shan. General distribution: China (the
former Kuldzha District). Type in Leningrad.

Note. The species was described from specimens grown in the
Petersburg Botanical Garden from seeds sent by Dr. A. Regel from
“Chinese Turkestan.” The authors of the species compared it with S.
salicifolia, distinguishing it from the latter and all other closely related
species by the glabrous leaves, of which the cauline leaves are decurrent
as a rather broad wing, and also the setose-scaly receptacle. The latter
character, introduced by Regel and Winkler, is characteristic for all
members of series Salicifoliae Lipsch. In my view, S. alberti is closest
to S. larionovii (cf. the note to the latter species).

Series 8. Pulviniformes Lipsch.—Plants forming pulvinate turf. Stem
short, 1.5—2 cm long, one-headed. Leaves pinnatifid, small. Pappus
bristles, according to Winkler, broader at apex.

33. A. pulviniformis Winkl. in Tr. Peterb. Bot. Sada, XI, 2 (1891)
BT:

Perennial. Plants forming dense pulvinate turf, almost glabrous. Root
thick, woody, with fibrously split bark, many-headed. Stems 1.5—2 cm
long, numerous, at densely covered base with hard brown remains of
old leaves. Leaves small, pinnate; basal leaves petiolate, sheathing at
basally, pinnatilobate, lobes small, almost roundish, obtuse, entire or
toothed, and then teeth subobtuse; terminal lobe larger, orbicular,
subobtuse or less often elongate-subobtuse; cauline leaves few, like basal
in shape, amplexicaul. Capitula up to 1 cm long and 0.5 cm wide,
solitary, terminal on stem apex, glabrous. Involucre five- or six-rowed,
all involucral bracts membranous; outer bracts ovate, subobtuse; inner
bracts lanceolate or ovate-lanceolate, with short cusp or subobtuse;
receptacle with numerous small scales; appendages of anthers woolly
(?). Achenes obpyramidal, four-ribbed; pappus bristles long-plumose,
broader at apex. Flowering V.

Found outside the USSR, in the Kuldzha District (Iren-khabirga and
Taldy ranges). Possibly may be found in the USSR. Type in Leningrad.

421

Note. It has not been collected by anyone since A. Regel. The
428 authentic specimen, which has been-in the hands of Winkler and me, is
incomplete and does not permit one to give a more detailed description
of the floret and pappus (apparently, the latter is two-rowed). The species
is tentatively included in section Laguranthera.

Section 3. Benedictia DC. Prodr. VI (1837) 533; Endlich. -Gen. PI.
(1836-1840) 468.—Eusaussurea C.A.M. ex DC. op. cit. (1837) 522
(ined.).—Sect. Lagurostemon (Cass.) DC. ex Kitam. in Acta Phyt. et
Geo-bot. IV (1935) 4 p. p.; Kitam. in Mem. Coll. Sc. Kyoto Univ. ser.
B, XIII (1937) 150 p. p—Sect. Chaetocline Franch. in Journ. de Bot. Il,
18 (1888) 311.—Root and rhizome not fibrously split. Involucral bracts
with or without herbaceous saber-shaped appendages. Receptacle with
scales. Appendages of anthers ciliate, not comose-woolly. Pappus double.

_— Subsection 1. Odontolepis Kitam. in Acta Phyt. et Geobot. IV (1935)
4; Ej. in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII (1937) 150.—Involucral
bracts cristate-pinnatifid.

34. S. odontolepis Sch. Bip. ex Herd. in Bull. Soc. Nat. Mosc.
XLI, 3 (1868) 131 p. syn.; Maxim. in Mel. Biol. XI (1883) 803 (descr.
emend.) and Bull. Acad. Sc. Pétersb. XXIX (1883) 176; Kom. FI.
Manchzh. III, 732; Nakai, Fl. Korean. II, 42; Ej. in Bot. Mag. Tokyo,
XXIX, 206; Ej. Fl. sylv. Korean. XIV, 119; Kom. and Alis. Opred.
Rast. Dalnevost. Kr. II, 1071; Kitam. in Mem. Coll. Sc. Kyoto Univ.
ser. B, XIII, 150.—Saussurea pectinata B. amurensis Maxim. Prim. FI.
Amur. (1859) 171.—S. ussuriensis Maxim. 5. odontolepis Herd. op. cit.
(1868) 13.—Ic.: Kom. and Allis. op. cit. Plate 320.

Perennial. Rhizome strong, inclined, with fibrous roots. Stem erect,
40-100 cm high, densely leafy, sulcate-angular, sparsely pubescent or
almost glabrous, branched or less often simple in upper part. Leaves
sharply scabrous, especially above and along margin due to more or
less numerous short spines, pubescent beneath and usually with numerous
lustrous glandular hairs, less often subglabrous and not glandular,
petiolate, deeply pinnatipartite almost up to midrib, with numerous (10—
16, sometimes less) lateral pairs of lobes; lateral lobes remote, in type
form long, almost linear, acute, entire or in turn toothed with many
pairs of teeth, their tips like primary lobe with cartilaginous spiny cusp;
middle and upper cauline leaves not decurrent on stem; uppermost leaves
reduced, sessile, cristate-pinnate. Capitula pedunculate, 1.2—-2 cm long
and 0.5—0.8 cm wide, in dense compound corymb. Involucre tubular, .
arachnoid-hairy. Involucral bracts with lustrous glandular hairs; outer
bracts lanceolate, with straight or weakly deflexed saber-shaped

430

422

appendages, cristate-toothed, inner bracts longer, linear, dark red or violet
at apex, usually entire. Receptacle with linear-subulate, lustrous, unequal,
up to 0.4—0.5 cm, long scales. Florets pink, about 1—1.1 cm long, narrow
part of tube 0.5—0.6 cm long, almost as long as its broad part with
deeply divided lobes of limb; appendages of anthers ciliate. Pappus
double, outer pappus-bristles less numerous, fragile, toothed, unequal,
up to 2 mm long, inner ones about 6-8 mm long, plumose; achenes
glabrous 3—5 mm long, about 2 mm wide, with short-toothed corona,
narrowed toward base, dark mottly. Flowering VII—VIII.

Meadows, sandy-gravelly deposits, thickets of shrubs, oak forests,
turf-covered rocks.—Far East: Zeya-Bureya, Ussuri. General distribution:
Korea, northwestern China. Described from southern Amur and from
the Burei Range. Type in Leningrad.

Note. The Chinese species S. pectinata Bge., which is not found in
the USSR, also belongs to subsection Odontolepis Kitam. S. odontolepis
Sch. Bip. differs from S. pectinata by having capitula only about half
as large, narrower and more weakly pubescent involucral bracts, and
also more numerous and narrower lateral lobes of the leaves.

Subsection 2. Hololepis Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 372.—Involucral bracts entire.

Series 1. Monocephalae Kitam. in Mem. Coll. Sc. Kyoto Univ.
ser. B, XIII (1937) 152, emend. Lipsch. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XXI (1961) 372.—Capitula solitary, less often two.
Involucre ovoid or campanulate, bracts imbricate, acute or subacute.
Leaves white-tomentose beneath, more or less pubescent or almost
glabrous above, glabrous on both sides only in S. sajanensis, but
arachnoid-hairy along margin, midrib, and petiole.

35. S. jadrinzevii (~jadrinzevi) Kryl. in Tr. Bot. Muz. XIV (1915)
143; Kryl. Fl. Zap. Sib. XI, 2922.—Ie.: Kryl. op. cit. (1915) 145, Plate
4 (colored).

Perennial. Rhizome inclined or ascending, dark brown, with fibrous
roots in lower part. Stems 7-20 cm high and 1-2 mm thick, solitary,
erect, simple, obtusely ribbed, somewhat tomentose; particularly densely
in upper part, weakly leafy. Leaves, except uppermost, petiolate,
coriaceous; basal leaves two to five, petiolate, petioles almost as long
as lamina, 2—7 cm long and 1-—2.5 cm wide, oblong-elliptical, obtuse or
cuspidate, with roundish or less often somewhat cuneate, sometimes
weakly cordate base, toothed throughout, teeth infrequent, unequal, with
very short callose, erect or decurved cusp, teeth sometimes indistinct,
very small; lamina bicolored, snow white-tomentose beneath excluding
midrib, green above, covered with white-tomentum only along veins,

423

forming distinct white network; cauline leaves like basal, three or four,
short-petiolate, narrower and long acuminate, lanceolate or elliptical-
lanceolate; uppermost leaves small, sessile, linear or lanceolate-linear.
Capitula solitary, terminal on stem, about 2 cm long at fruiting, up to
1.5 cm wide. Involucre ovoid, imbricate, four- or five-rowed, bracts
unequal; outer bracts ovate, cuspidate, more or less tomentose and with
sparse lustrous glandular hairs, 5-7 mm long and 3—4 mm wide; inner
bracts oblong, subobtuse, 10-14 mm long and 2-3 mm wide, densely
covered with straight, long, silky, stiff, appressed hairs and isolated
lustrous glandular hairs. Receptacle densely covered with linear-subulate,
lustrous, 0.5—0.8 cm long scales; florets lilac-pink, about 1.4 cm long,
narrow part of tube about 0.6 cm long, almost as long as broad part
along with lobes of limb; latter divided slightly deeper than half in
linear, obtuse lobes; appendages of anthers ciliate. Pappus double, outer
pappus-bristles serrate-scabrous, unequal, about 3-4 mm long, persistent,
inner ones long-plumose about 1.2—1.3 cm long; achenes dark brown,
smooth, 0.5—-0.6 cm long. Flowering VII.

Limestone rocks and stony slopes.—Western Siberia: Altai (Kuldzha
River Valley). Endemic. Described from the Chuya River. Type in
Leningrad; isotype in Tomsk.

Note. Sergievskaja (Kryl. Fl. Zap. Sib. (Flora of Western Siberia),
XI, 2923) reported a hybrid between S. jadrincevii Kryl. and S.
controversa DC., which she named var. intermedia Serg. and
characterized as having taller (up to 40 cm high) stems, two to five
compactly arranged capitula, and a more weakly developed tomentum
on the upper leaf surface.

36. S. tomentosa Kom. in Bot. Mat. Gerb. Glavn. Bot. Sada, II,
33-34 (1921) 135.

Perennial. Stem (4)15—20(25) cm high (occasional specimens with
2.5—3 cm high stem), pubescent, leafy, simple, one-headed, less often
weakly branched at apex and then two-headed. Leaves lanceolate-oblong
or linear, acuminate, sometimes subobtuse; basal and lower cauline leaves
usually broader, (3)5.5-12 cm long and 1-2 cm wide, long-petiolate;
upper cauline leaves short-petiolate, weakly toothed or entire, grayish
or whitish-tomentose beneath, weakly arachnoid-hairy or almost glabrous
above. Capitula solitary, less often two, in type form up to 2—2.5 cm
wide, usually subtended by one to three linear bracteal leaves. Involucre
four-rowed; involucral bracts almost glabrous, white-tomentose only along
margin and apex, ovate-triangular, acuminate, recurved. Receptacle with
about 0.6—-0.7 cm long scales; florets pink-violet, about 1.2 cm long;
narrow part of tube about 0.6 cm long, almost as long as broad part

431 including lobes of limb; anthers lustrous, grayish-blue; appendages of

424

anthers ciliate. Pappus double, smoke-colored, outer pappus-bristles less
numerous, serrate, fragile, small, about 2-3 mm long, inner ones long-
plumose, about 1—1.1 cm long; achenes about 5 mm long, dark brown,
glabrous, ribbed, with toothed corona. Flowering VIJ-—VIII.

Rocks; stony-lichen tundra. Far East: Ussuri (Sikhote-Alin, ‘Dusse-
Alin’), ?Uda region Endemic. Described from the left bank of the Amur
River in the Khabarovsk District. Type in Leningrad.

Note. In the original description it was compared with representatives
of the group of S. alpina DC., from which it can be differentiated at
once by the one-headed stem, broad receptacle and involucral bracts
twisted in the upper part. Undoubtedly, it belongs to the series
Monocephalae Kitam. of one-headed species, which mostly have
bicolored leaves, white-tomentose beneath and green, glabrous or more
or less pubescent above. It is particularly close to S. porcellanea Lipsch.;
S. tomentosa Kom. differs from this species by larger capitula, up to 2—
2.5 cm wide, distinctly acuminate, deflexed involucral bracts, and lower
leaves lacking a cordate base and with weaker pubescence. (Plate XXXI,
Fig. 1.)

37. S. porcellanea Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVI (1954) 455 (nomen).—S. nivea Kom. and I. Schischk. in
Bot. Mat. Gerb. Sada, VI, I (1926) 17, non Turez. 1837.—S. nakaiana
Kom. in Izv. Bot. Sada Akad. Nauk SSSR, XXX, 1-2 (1932) 220, saltem
ex descriptone, non e typo.

Perennial. Stem erect, (6)8—15(20) cm high, densely white-tomentose,
glabrescent, simple, one-headed, less often slightly branched at apex
and then two-headed. Leaves cordate, petiolate, acuminate, toothed, teeth
with blackish cusps, densely white-tomentose beneath, tomentose-
arachnoid hairy above, glabrescent, and then green, basal leaves with
petiole 512 cm long, 2-3(6) cm wide, petiole 2-4(5.5) cm long; cauline
leaves gradually reduced toward stem apex, uppermost leaves linear-
lanceolate or linear, narrow. Capitula solitary or less often two, usually
subtended by one or two, linear white-tomentose bracts, about 2—2.5 cm
long and 0.8—1.1 cm wide. Involucre four-rowed, outer involucral bracts
ovate or ovate-triangular, pubescent or almost glabrous, white-tomentose
only above; middle and inner bracts oblong, usually dark-colored,
glabrous, white-tomentose only along margin, less often entirely
pubescent; all bracts subobtuse. Receptacle with linear, about 0.5—0.7
cm long scales; florets pink-violet, about 1.3 cm long, narrow part of
tube 0.5—0.6 cm long, as long as broad part together with lobes of limb;
anthers brown-violet, appendages of anthers ciliate. Pappus dirty, double,

432 outer pappus-bristles serrate, fragile, less numerous, about 4 mm long,

433

425

inner ones long-plumose, about 1—1.1 cm long; achenes about 4-5 mm
long, glabrous, with crown. Flowering VII—VIII.

Gravelly, heath-lichen alpine tundra, stony glades among the thickets
of Japanese stone pine.—Far East: Ussuri Region (Sikhote-Alin Reserve).
Endemic. Described from Sikhote-Alin. Type in Leningrad.

Note. Saussurea nivea Kom. and I. Schischk. was found to be
identical with S. nakaiana Kom. Since the name S. nivea Kom. and
Schischk. is a later homonym, it cannot be retained. True S. sovietica
has been wrongly cited as the type specimen in the description of S.
nakaiana Kom., although S. nakaiana Kom. itself was described correctly
from a different specimen. In view of the confusion about this species,
I was compelled to give it a new name. In the nature of pubescence of
stems, leaves and involucral bracts, S. porcellanea is closest to S.
sovietica, possibly its derivative; it differs by weaker growth and the
presence of one or two capitula and connects the two series
Monocephalae Kitam. and Sovieticae Lipsch.

38. S. kitamurana Miyabe and Tatewaki in Trans. Sapporo Nat.
Hist. Soc. XIV, 4 (1936) 267; Kitam. in Mem. Coll. Sc. Kyoto Univ.
ser. B, XIII, 152; Sugaw. Ill. Fl. Saghal, IV, 1883.—S. komarovii
Soczawa in Sov. Bot. XIII, 1 (1945) 29 (nomen); Lipschitz in Bot. Mat.
Gerb. Bot. Inst. Akad. Nauk SSSR, XVI, 455.—Ic.: Miyabe and
Tatewaki, op. cit. (1936) fig. 16; Sugaw. op. cit. (1940) tab. 865.

Perennial. Rhizome short-creeping, sparsely covered with bases of
previous year’s leaves; stem short, 4.5—7 cm high, erect, simple, sulcate,
arachnoid-hairy, glabrescent, one-headed. Leaves succulent, ovate, or
ovate-oblong, somewhat acuminate, with cartilaginous cusp, cuneate or
almost roundish at base, lamina about 6 cm long and 1—1.5(2) cm wide,
crenate-toothed, teeth cartilaginous; basal leaves persisting up to
flowering; lower cauline leaves petiolate, petioles 1.5—2 cm long; upper
leaves lanceolate-oblong or lanceolate, smaller and narrower, 0.2—0.7(1.2)
cm wide, short-petiolate or sessile. Capitula solitary, (0.5)1—2(2.5) cm
wide, often with less numerous bracts near base, similar to involucral
bracts. Involucre campanulate, imbricate; involucral bracts four-rowed,
dark, outer and middle ovate, obtuse; inner bracts oblong or oblong-
linear, far exceeding outer, acuminate, often flocculose bracts. Receptacle
with linear-subulate, lustrous about 0.5—0.6 cm long scales; florets about
1.1—1.2 cm long, narrow part of tube about 0.6 cm long, almost as long
as broad part including lobes of limb; appendages of anthers ciliate.
Pappus double, outer pappus-bristles serrate-scabrous, fragile, about 1—3
mm long, inner ones long-plumose, about 0.9-1.1 cm long; achenes
brown, 4—5 mm long, cylindrical, glabrous. Flowering VII—VIII. (Plate
XXXI, Fig. 2.)

434

426

Alpine lichen tundra; stony areas in the high-mountain region.—
Far East: Ussuri (northern Sikhote-Alin), Sakhalin. Endemic. S.
kitamurana was described from Sakhalin; type in Japan. The name S.
komarovii Soczava was proposed for the plant described from northern
Sakhalin; type in Leningrad.

Note. A comparison of specimens of S. kitamurana and S. komarovii
revealed their complete identity. Thus, S. kitamurana, earlier believed
to be a special Sakhalin endemic, is not endemic to this area but is also
distributed on the continent. S. kitamurana is quite close to S.
porcellanea, from which it differs by leaves that are grayish arachnoid
beneath (and not densely white-tomentose) and have a cuneate or almost
roundish base (distinctly cordate in S. porcellanea).

39. S. sajanensis Gudoschnikov in Sistemat. Zam. Gerb. Tomsk.
Univ. 77-78 (1954) 7; Lipschitz in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI, 372.—Ic.: Gudosch—nikov, op. cit. 8.

Perennial. A miniature alpine plant. Stem solitary, 2-7 cm high,
dark-colored, arachnoid-hairy, leafy throughout, wingless. Leaves
somewhat hard, mainly below, green, glabrous on both sides, arachnoid-
hairy along prominent midrib and margin; basal leaves petiolate, petioles
one-third to half long less often as long as lamina, lamina lanceolate or
almost ovate, basally cuneate or almost roundish, (1)2—5 cm long and
0.4-1 cm wide, shallow crenate-toothed, teeth remote, with short callose
cusp or somewhat convolute; cauline leaves few, narrower and smaller,
linear or lanceolate. Capitula solitary or in groups of two. Involucre
cylindrical, 1.1—-1.2 cm long and 6—10 mm wide; outer involucral bracts
ovate, inner lanceolate, both obtuse, usually dark-colored, pubescent.
Receptacle densely covered with linear-subulate, lustrous, up to 7 mm
long scales; florets pink-lilac, about 1.2 cm long, narrow part of tube
almost as long as broad part including lobes of limb; appendages of
anthers hairy. Pappus double, outer pappus-bristles fragile, small, inner
ones plumose, longer; achenes not mature, 3-4 mm long, brownish,
ribbed, glabrous. Flowering VII. (Plate XXXI, Fig. 3.)

Rocks.—Eastern Siberia: Angara-Sayan (western Sayans). Endemic.
Described from the Abakan River System (Lake Antyuzher-Kul). Type
in Tomsk; isotype in Leningrad.

Note. The author considers his species to be close to Saussurea
parviflora var. contracra Ldb. In my opinion, S. sajanensis belongs to
the series Monocephalae Kitam., which includes the unique unicapitulate
species found on rocks and stony outcrops, with a narrow range confined
to the local mountain systems from Altai to Sikhote-Alin.

Series 2. Sovieticae Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 372.—Capitula numerous, in dense compact

435

Plate XXXIII.

1—Saussurea sachalinensis Fr. Schm.; 2—S. acuminata Turcz.

NACA oN

Seq

Ud

436

428

corymb. Involucre ovoid or campanulate; involucral bracts imbricate,
with short blackish cusp. Leaves cordate. Entire plant, including leaves
on both sides, densely whité-tomentose.

40. C. sovietica Kom. in Izv. Bot. Sada Akad. Nauk SSSR, XXX,
1—2 (1932) 220; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1064.—
S. nakaiana Kom. in Izv. Bot. Sada Akad. Nauk SSSR, XXX, 1-2 (1932)
221 p. p. saltem quoad pl. No. 943.—Ic.: Kom. and Allis. op. cit. (1932)
Plate 319.

Perennial. Entire plant white-tomentose. Stem 25—60 cm high, erect,
woody, simple, densely white-tomentose. Leaves densely white-tomentose
on both sides (weakly above), acuminate, distinctly irregularly toothed,
teeth terminating in short, callose black cusp; basal leaves hastate,
acuminate, cordate or less often cuneate, long-petiolate, 10-15(18) cm
long and 4—7 cm wide, petiole weakly sulcate, 2.5-8 cm long and about
3 mm wide; middle cauline leaves short-petiolate, broadly cuneate; upper
leaves much smaller, lanceolate, narrowly cuneate. Capitula up to 13 in
compact corymb; individual corymbs on hard, erect, short, whitish-
pubescent peduncle; young capitula ovoid, white-tomentose, ovoid-
campanulate when mature, somewhat swollen below, about 2 cm long.
Involucre imbricate, four- to six-rowed, initially densely white-tomentose
(bracts tomentose), later sometimes subglabrous in upper part; outer bracts
ovate, inner oblong, longer than outer, erect; all bracts with short dark-
colored cusp. Receptacle densely covered with linear, lustrous, about
0.9 cm long scales, almost as long as inner involucral bracts; florets
pink; about 1.2 cm long, narrow part of tube about 0.6 cm long, as long
as broad part including lobes of limb; appendages of anthers ciliate.
Pappus double, outer pappus bristles serrate, fragile, about 3-4 mm long,
inner ones plumose, about 1 cm long; achenes (immature) glabrous,
ribbed, about 4 mm long. Flowering VIII-IX.

Rocks and stony heaths——Far East: Ussuri (southern Sikhote-Alin).
Endemic. Described from southern Sikhote-Alin. Type in Leningrad.

Note. Komarov compared his species with S. eriolepis Bge., from
which it differs by leaves which are densely white-tomentose above,
white-tomentose involucral bracts, and by hastate and not cordate lower
leaves in the type specimen. It appears to me that S. sovietica is closer
to S. porcellanea Lipsch., from which it can be distinguished instantly
by the presence of numerous white-tomentose capitula (capitula are
solitary in S. porcellanea, rarely 2; involucral bracts glabrous, only at
apex white-lanate), and taller plants. In the list of typical specimens of
S. sovietica, Komarov included the herbarium sheet No. 935 (Tazovskaya
Mountain in southern Sikhote-Alin, B.K. Schischkin), which totally agrees
with the diagnosis; I accept this specimen as the lectotype of S. sovietica.

429

Along with this specimen, Komarov wrongly mentioned No. 943; this
specimen already belongs to S. porcellanea Lipsch. (compare the note
to the latter). S. sovietica Kom. is one of the most ornamental species
of Saussurea, which should be introduced into cultivation.

Series 3. Squarrosae Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 373.—Capitula usually numerous, in dense
compact corymb. Involucral bracts imbricate, with long squarrose cusp.
Leaves white-tomentose on both sides, more densely beneath, cuneate
or cordate at base. |

41. S. squarrosa Turcz. in Bull. Soc. Nat. Mosc. XX, 2 (1847) 44;
Ej. Fl. baic.-dahur. II, 1, 117; Popov, Fl. Sr. Sib. I, 762; Lipschitz in
Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XXI, 373.—Ic.: Popov,
op: citai(1959), Plate).895 Fig: 1:

Perennial. Root collar covered with blackish-brown remains of old
leaves. Stem arachnoid, glabrescent, reddish, simple, sometimes weakly
branched only above, 10—20(30) cm long. Leaves sparsely flocculose-
arachnoid hairy to villous above, white-tomentose beneath, variable in
form and size, more or less serrate-toothed, teeth terminating in black,
sometimes weakly curved cusp; basal and lower cauline leaves petiolate,
lanceolate or oblong-lanceolate, 4-10 cm long including petiole, 1—2(2.5)
cm wide, petiole arachnoid-white tomentose, 1.5—3 cm long; upper
cauline leaves like lower, short-petiolate or sessile, not decurrent; all
leaves distinctly cuneate. Capitula less numerous (three to seven), 1.2—
1.4 cm long and 0.8—-1 cm wide, pedunculate, in compact corymbs,
usually subtended by bracts. Involucre four-rowed; involucral bracts
usually dark-colored, ovate or oblong, acute; outer bracts in type form
squarrose, white-tomentose, inner appressed, erect, pubescent. Receptacle
with linear, lustrous, about 4-5 mm long scales; florets dark violet,
about 0.8—1 cm long; narrow part of tube about 0.4-0.5 cm long, almost
as long as broad part including lobes of limb; appendages of anthers
ciliate-hairy. Pappus double, outer pappus-bristles quite numerous,
unequal, toothed-scabrous, about 2-3 mm long, inner bristles long-
plumose, about 0.8—0.9 cm long. Flowering VII—VIII.

Limestone rocks in the alpine zone.—Eastern Siberia: Angara-Sayans
(Tunkin alpine zone). Endemic. Described “from alpine zone of Nukhu-
Daban.” Type in Leningrad.

Note. A forgotten species, described by N.S. Turczaninow and
subsequently not cited by anyone except M.G. Popov. I saw the authentic
specimen of Turczaninow, and without any doubt I include the specimen
from the Tunkin alpine zone under this unique, elegant species segregated
by Turczaninow. While describing the new species, Turczaninow had a

438

430

few specimens, apparently dwarfs; Turczaninow’s indication that the
receptacle of S. squarrosa is glabrous is incorrect; it is distinctly scaly.
This species differs from S. jadrincevii Kryl. by more or less numerous
capitula in corymbs and unique squarrose involucral bracts; it differs
from S. porcellanea Lipsch., with which it was identified by some
authors, by capitula in a corymbose inflorescence, cuneate (and not
cordate) leaves, and squarrose, distinctly pointed involucral bracts; from
S. lenensis M. Pop. it differs by the shape of the leaves and pubescence
on the upper leaf surface, as well as by smaller capitula on short
peduncles. Popov collected a form very close to S. squarrosa Turcz.
(eastern Sayans, Arshan health resort, alpine zone, cirque, 21, VIII, 1953,
M.G. Popov), which has to be considered as a new species—S.
pseudosquarrosa M. Pop. and Lipsch.

42. S. pseudosquarrosa M. Pop. and Lipsch. in Popov, FI. Sr. Sib.
II (1959) 762 (descr. ross); Lipschitz in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961), 373 (descr. lat.)—-S. foliosa var. dentosa Turcz.
in DC. Prodr. VI (1837) 536 p. p.; Turcez. Fl. baic.-dahur. II, 1, 116 p.
p.; Herd. in Bull. Soc. Nat. Mosc. XLI, 3, 38 p. p.

Perennial. Root short, vertical, many-headed, root collar covered
with hardened remains of previous year’s leaves. Stem (10)!5—20 cm
long, cylindrical or often flattened, predominantly in upper part. Leaves
of vegetative shoots on long glabrous petioles, with oblong, acute, lobed
at base or almost cordate lamina, light green and glabrous above, white
tomentose beneath, crenate-sharply toothed, 6-10 cm long, 2-4 cm wide;
cauline leaves on shorter petioles, lanceolate-oblong or lanceolate, like
basal leaves but narrower. Capitula 7-10, on short peduncles, with linear
bracteal leaves, aggregated in dense, almost capitate corymb. Involucre
imbricate, more or less arachnoid hairy-tomentose, about 10 mm long,
outer involucral bracts ovate, terminating in long, deflexed or erect
cusp. Florets purple, about 1 cm long, narrow part of tube almost as
long as broad part including lobes of limb. Receptacle scaly; scales
linear-subulate, lustrous, unequal. Pappus somewhat dark, outer pappus
bristles unequal, small, fragile, scabrous, inner ones plumose, about
0.9—1 cm long; achenes about 5 mm long, glabrous, lustrous. Flowering
VII-VIII.

Alpine and subalpine zone.—Grass plots, stony screes, gravelly
slopes.—Eastern Siberia: Angara-Sayans (Tunkin alpine peaks, Sokhodno
and other mountains). Endemic. Described from the alpine peaks of

‘Arashan. Type in Leningrad.

Note. Evidently this species is of hybrid origin, involving S.
squarrosa, S. controversa and, possibly, S. pricei, as well as S. sukaczevii.
It differs from S. squarrosa Turcz. by its leaves that are glabrous above,

439

431

more strongly crenate-sharply toothed, and almost cordate or lobed at
base. Earlier I wrongly included some specimens of this species under
S. sukaczevii Lipsch.

Series 4. Bicolores Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 374.—Capitula more or less numerous, in simple
corymb or paniculate-corymbose inflorescence. Involucral bracts
imbricate, subobtuse or acute. Leaves distinctly cordate or less often
truncate at base, bicolorous—densely white-tomentose beneath, and
glabrous and green above.

43. S. controversa DC. in Ann. Mus. Paris, XVI (1810) 199; Kryl.
Fl. Zap. Sib. XI, 2921.—S. discolor DC. Prodr. VI (1837) 534 p. p. and
B. elatior DC. op. cit. (1837) 535 (excl. syn. S. eriolepis Bge.); Ldb. Ic.
pl. Fl. Ross. I, 18; Ej. Fl. alt. 1V, 27; Ej. Fl. Ross. II, 2, 668; Turcz. FI.
baic.-dahur. I], 1, 113; Kryl. Fi. Alt. III, 707; Popov. Fl. Sr. Sib. II,
758.—S. discolor B. elatior s. sibirica (Fisch. herb.) Herd. in Bull. Soc.
Nat. Mosc., XLI, 3 (1868) 30.—S. propinqua I\jin in Pavl. in Byull.
Mosc. Obdshch. Isp. Prir. Ser. Biol. XX XVIII, 1-2 (1929) 146; Fl. Yugo-
Vost. VI, 385.—Ic.: Fl. Yugo-Vost. VI, 387.

Perennial. Root collar loosely covered with remains of petioles of
previous year’s leaves. Stems 25—100 cm high, erect, solitary (less often
two), simple, often branched only in inflorescence, more or less
arachnoid-hairy, usually dark, violet, or dark-brown. Leaves bicolored,
green, glabrous or at places weakly flocculose-arachnoid hairy above,
white-tomentose beneath, toothed, teeth broad, unequal, callosely-pointed;
leaf in apical part usually entire; lower leaves long-petiolate, with cordate,
ovate-triangular or ovate-oblong lamina, attenuate-acuminate, deeply
cordate at base, 5-17 cm long and 2.5—9 cm wide, almost as long as
pulvinate petiole; middle cauline leaves short-petiolate, like lower leaves
but usually narrower, with obtuse, sometimes weakly not equilateral
(but not cordate) base; upper leaves reduced, narrow-lanceolate, sessile.
Capitula up to 1.5 cm long (including florets) and about 0.8—1 cm wide,
sometimes aggregated in more dense solitary terminal corymb, sometimes
at the apices of more or less numerous, lateral branches of lax panicle.
Involucre arachnoid-hairy, imbricate; its bracts usually black-violet or
rarely light colored; outer bracts broadly ovate, short-acuminate; inner.
bracts oblong, narrowed above and obtuse, densely woolly-pubescent.

Receptacle densely squamose, ‘scales linear-subulate, lustrous, up to 0.5

cm long; florets lilac-violet, about 1.2 cm long; narrow part of tube
about 0.6 cm long, almost as long as broad part including lobes of
limb; appendages of anthers pubescent. Pappus double, outer pappus-
bristles 2—3(4) mm long, serrate, fragile, inner ones plumose, about

432

0.9-1 cm long; achenes about 4 mm long, glabrous, with a narrow
corona. Flowering VII—VIII.

Mountain forest-meadows, open forests (larch, birch and pine), forest-
covered stony slopes of mountains, limestone deposits, chernozem steppe-
meadows.—European Part: Dvina-Pechora (eastern part), Volga-Kama;
Western Siberia: Ob’, Upper Tobol, Irtysh (eastern part), Altai; Eastern
Siberia: Yenisei, Angara-Sayans, Dauria, Lena-Kolyma. General
distribution: Mongolia (northern). S. controversa DC. is described from
Siberia; type in Geneva. S. propinqua Iljin is described from Mongolia;
type in Leningrad.

Note. It hybridizes with: 1) S. jadrinzevii Kryl. (cf. note to this
species); 2) S. alpina (L.) DC. (cf. below, S. uralensis Lipsch.); and 3)
S. parviflora (Poir.) DC. The last hybrid is known from the Irkutsk
District, and has been found in the materials of N. Malzev for the issue
of the Herbarium of the Russian Flora. Malzev collected S. parviflora
in 1903 from the former Balagansky District, near the village of
Bazhevskoe, and this material included an “admixture” of several sheets
of S. controversa. While studying a large number of specimens of S.
parviflora (Poir.) DC., I found several sheets with specimens intermediate
between S. parviflora and S. controversa, which undoubtedly should be
considered as hybrids of the two species. These specimens are
characterized by somewhat larger capitula than in S. parviflora (deviation
towards S. controversa), much broader ovate-oblong leaves than in
S. parviflora, and some of them with a distinct weakly cordate base
(in others with an unequal, truncate base), leaves slightly arachnoid
beneath (deviation towards S. controversa), as well as leaf petioles
weakly or not at all decurrent. These hybrids were not mentioned in the
literature.

44. S. discolor (Willd.) DC. in Ann. Mus. Paris, XVI (1810) 199;
Ey. Prodr Viv 534 p. p.; jeer, I Pe VI, 2, 836; Vizn. Rosl. URSR,
562.—Serratula discolor Willd. Sp. pl. II (1800) 1641.—Saussurea
discolor a. genuina s. europaea Herd. in Bull. Soc. Nat. Mosc. XLI, 3
(1868) 30.—Iec.: Rchb. Ic. Fl. Germ. XV, tab. 818; Cusin. Herb. FI.
Franc. XIII, tab. 381; Hartinger, Atlas Alpenpfl. V, tab. 474; Bonnier,
Fl. Compl. France, Suisse and Belg. VI, tab. 333; Hegi, Ill. Fl. VI, 2
lig S936, 554. 555:

Perennial. Quite close to previous species from which it differs by
having a shorter (5-25 cm high, less often taller), usually slightly
ascending stem, a more compact inflorescence, fewer (two or three, less
often five) capitula, and a more weakly developed cordate bases of the
basal and lower cauline leaves. Flowering VII-IX.

440

433

Crevices in rocks.—European Part: Upper Dniestr. General
distribution: Central Europe (from France to Austria). Described from
Central Europe. Type was in Berlin.

Note. I did not see the herbarium specimens from the Soviet
Carpathians. Judging from the numerous specimens of this species from
Central Europe examined by me, S. discolor (Willd.) DC. is extremely
close to S. controversa DC. The distinguishing characters of both species
listed above are quite unstable and poorly developed. But since the
ranges occupied by these species are sharply isolated, in the present
treatment of Saussurea | consider the two species as morphologically
weakly differentiated geographic races of a previously single species.

45. S. hypargyrea Lipsch. and Vved. in Byull. Mosc. Obshch. Isp.
Prir., Otd. Biol. LIX, 6 (1954) 74.—S. discolor auct. fl. Jacutiae.

Perennial. Root collar loosely covered with remains of petioles and
sheaths of previous year’s leaves. Stem single, 25-45 cm high, erect,
simple or more or less branched, usually dark violet, less often green,
tomentose, glabrescent, weakly leafy. Leaves oblong-lanceolate or linear-
lanceolate, bicolorous, green above, white-tomentose beneath, weakly
toothed, teeth with dark callose, sometimes slightly curved cusp, more
or less cuneate or, less often roundish-truncate at base but never cordate;
basal and lower cauline leaves petiolate, 4—12(17.5) cm long (including
1-5(9) cm long petiole), 0.5—1.7(2) cm wide; middle and upper cauline
leaves reduced, linear-lanceolate to linear, short-petiolate or almost
sessile. Capitula 1.8—2 cm long and 0.7—0.8 cm wide, in terminal corymbs
of two to five capitula, besides often solitary or in groups of three on
long lateral branches arising at different heights on stem. Involucre
imbricate, four-rowed, villous; outer involucral bracts ovate, inner oblong,
all subobtuse, weakly lilac colored or concolorous, especially along
margin. Receptacle densely covered with whitish, linear-subulate,
lustrous, about 0.8 cm long scales; florets pink-violet, 1.1—1.2 cm long,
narrow part of corolla tube about 6 mm long, almost as long as broad
part including lobes of limb; anthers violet, their appendages hairy.
Pappus double, outer pappus bristles serrate, less conspicuous, less
numerous, short, fragile, inner ones plumose, about 1 cm long. Flowering
Vi-VIl.

Limestone rocks, pine forests——Eastern Siberia: Lena-Kolyma
(Aldan River and its tributary). Endemic. Described from Yakutia. Type
in Leningrad.

Note. It is an elegant Yakutian race of S. controversa DC., forming
hybrids with S. pseudoangustifolia Lipsch., which are distinguished by
grayish (and not white-tomentose) pubescence on the lower leaf surface,
and narrower basal and lower cauline leaves.

441

434

46. S. ajanensis (Rgl.) Lipsch. in Byull. Mosk. Obshch. Isp. Prir.
Otd. Biol., LIX, 6 (1954) 73.—S. discolor 8. ajanensis Rgl. in Nouv.
Mém. Soc. Nat. Mosc. XI (1859) 107; Herd. in Bull. Soc. Nat. Mosc.
M153, "30:

Perennial. Stem 15-20 cm high, erect, simple, tomentose,
glabrescent, weakly leafy. Leaves bicolored, glabrous above, white-
tomentose beneath; basal leaves elliptical-oblong or oblong-lanceolate,
subobtuse or short acuminate, petiolate, 5.5—11 cm long (including 2.5—
4.5 cm long petiole), 1-1.5(2) cm wide, cuneate (not cordate) at base,
somewhat toothed, teeth terminating in black, scarcely curved cusps, or
leaves almost entire; lower cauline leaves like basal, almost sessile or
with extremely short petiole; uppermost leaves reduced, linear or linear-
lanceolate, sessile. Capitula in compact corymbs, three or four in one
corymb, almost sessile, less often solitary, about 1.5—-1.7 cm long and
0.7-0.8 cm wide, sometimes subtended by one or two bracteal leaves.
Involucre imbricate, four-rowed, hirsute; outer involucral bracts ovate,
inner oblong, all obtuse; florets pink-violet, 1.2 cm long; narrow part of
tube about 0.5 cm long, as long as broad part including lobes of limb;
appendages of anthers pubescent. Pappus double, outer bristles poorly
developed, less numerous, small, fragile, serrate, inner ones about 1 cm
long, plumose. Flowering VII.

Stony slopes.—Eastern Siberia: Okhotsk (vicinity of Ayan).
Endemic. Described from the vicinity of Ayan. Type in Leningrad.

Note. S. ajanensis (Rgl.) Lipsch. is closest to S. lenensis M. Pop.,
from which it differs by a shorter stem, almost sessile capitula in compact
corymbs, obtuse involucral bracts, short acuminate and weakly toothed
leaves (in S. Jenensis M. Pop. the involucral bracts are distinctly
acuminate, the leaves are acute and deeply toothed). From S. hypargyrea
Lipsch. and Vved. it differs by a shorter stem, sessile capitula, three or
four in a corymb, or less often solitary (S. hypargyrea has pedunculate
capitula in terminal corymbs; besides, there are lateral capitula on long
branches).

47. S. lenensis M. Pop. ex Lipsch. in Byull. Mosk. Obshch. Isp.
Prir. Otd. Biol. LIX, 6 (1954) 75; Popov, Fl. Sr. Sib. II, 763.

Perennial. Stem 30-35 cm long, erect or weakly flexuous, sometimes
dark red, tomentose, glabrescent, branched above, weakly leafy. Leaves
lanceolate or oblong-lanceolate, attenuate-acuminate, almost coriaceous,
bicolored, glabrous and green above, white-tomentose beneath, with
numerous teeth terminating in blackish somewhat curved cusps, weakly
cuneate at base but not cordate, basal and lower cauline leaves petiolate
up to 13 cm long and about 1—-1.3 cm wide; middle and upper cauline
leaves reduced, almost sessile or short-petiolate. Capitula two to four,

435

442 about 2 cm long, on long peduncles. Involucre imbricate, four-rowed,
villous; outer bracts ovate or ovately triangular, inner oblong; all bracts
distinctly acuminate, florets about 1.2 cm long; narrow part of tube 6
mm long, as long as broad part including lobes or limbs; appendages of
anthers hairy. Pappus double, outer pappus bristles numerous, serrate,
fragile about 3-4 mm long; inner ones long-plumose, about 1 cm long;
achenes 3-4 mm long, glabrous. Flowering VII.

Stony slopes.—Eastern Siberia: Yenisei (Kirensk District, along
Lena River). Endemic. Described from the Kirenk District. Type in
Leningrad.

Note. It is closest to S. ajanensis (Rgl.) Lipsch., from which it
differs by a taller stem, attenuate-acuminate, strongly toothed leaves,
long-pedunculate capitula, distinctly acuminate involucral bracts, and
more strongly developed outer pappus-bristles.. Undoubtedly, S.
hypargyrea Lipsch. and Vved. is also related phylogenetically, from
which it differs by more attenuate-acuminate and distinctly toothed
leaves, as well as acute involucral bracts.

48. x S. uralensis Lipsch. in Byull. Mosk. Obshch. Prir. Otd. Biol.
LIX, 6 (1954) 75.—S. controversa DC. x S. alpina DC. in Lipsch. op.
cit. (1954) 75.

Perennial. Root collar loosely covered with brown remains of leaves.
Stem solitary (less often two), (3.5)5—20 cm long, often violet, erect,
simple, more or less arachnoid-hairy. Leaves bicolored, green and
glabrous above, white-tomentose beneath, flat along margin, toothed,
teeth callose, unequal; lower leaves long-petiolate, with oblong-
lanceolate, roundish-truncate at base (not cordate) or weakly cuneate
lamina, attenuate-acuminate, (2.5)4.5-10 cm long and (0.8)1.5—2.5(3.5)
cm wide, petiole sheathing; middle leaves like lower, short-petiolate;
upper leaves reduced, sessile, linear or lanceolate. Capitula cupuliform,
up to 1.5—2 cm long and about 1 cm wide, in dense corymb of less
numerous (two to five) capitula, less often solitary. Involucre 0.8—1(1.2)
cm wide, imbricate, villous; bracts usually dark colored; outer bracts
broadly ovate, short acuminate, sometimes glabrescent, inner oblong,
subobtuse, densely woolly. Receptacle with linear-subulate, lustrous,
unequal, up to 5 mm long scales; florets lilac-violet, about 1.1 cm long,
narrow part of corolla tube about 6 mm long, almost as long as broad
part including lobes of limb; appendages of anthers slightly hairy. Pappus
double, outer pappus bristles 2-3 mm long, serrate, fragile, inner ones
about 0.9—1 cm long, plumose; achenes (immature) 3 mm long, brownish.
Flowering VII-VIII.

Mountain lichen-shrub tundra, birch-cedar dwarf forests, stony screes

443 of alpine zone. Also found in upper part of subalpine zone at tree-line

436

(larch-birch open forests) and sometimes descending on “tongues” of
stony screes in the lower part of this belt—European Part: Volga-Kama
(middle and southern Urals). Endemic. Described from Konzhakovskii
Kamen. Type in Leningrad.

Note. This species is the hybrid S. controversa DC. x S. alpina DC.
It is widely distributed over the entire Ural Range. It has inherited the
white-tomentose pubescence of the lower leaf surface and the shape and
size-of the capitulum from S. controversa and the plant height, shape of
the leaf base, nature of the involucral bracts, and compactness of the.
terminal corymb from S. alpina. It can immeditately be distinguished
from S. controversa by the shorter height, shorter and narrower, basally
roundish or weakly cuneate (and not cordate) lower cauline leaves, and
compact, few-headed corymb. It is also easily distinguished from the
type specimen of S. alpina DC. by the white-tomentose pubescence of
lower leaf surface and larger capitula.

Section 5. Splendidae Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 374.—Stem, lateral branches of inflorescence,
and outer involucral bracts glandular-hairy. Leaves large, bicolored, green
above, white-tomentose beneath. Involucral bracts with saber-shaped,
erect or curved appendage.

49. S. splendida Kom. in Tr. Peterb. Bot. Sada, XVIII, 3 (1901)
424; Kom. Fl. Manchzh. III, 736; Kom. and Alis. Opred. Rast. Dalnevost.
Kr. II, 1063.—Ic.: Kom. op. cit. (1907) Plate 15.

Perennial. Stem strong, erect, 90-120 cm high, sulcate, pubescent
with grayish-yellow or dark brown, articulate sticky hairs. Leaves
bicolored, green above, white-tomentose beneath; basal leaves long-
petiolate (petioles winged at base, semiamplexicaul, pubescent), up to
16 cm long excluding petiole, 9-11 cm wide, ovate, with almost cordate
base, acute, toothed (teeth with fine cusp) and ciliate under lens, short-
ciliate above, densely white arachnoid hairy-tomentose below; middle
cauline leaves short-petiolate, 10-11 cm long and 9 cm wide; upper
leaves weakly toothed, sometimes almost entire, ovate-lanceolate or
lanceolate, sessile, like middle leaves white-tomentose below.
Inflorescence paniculate, many-headed, comprising corymbs; corymbs
three to five-headed, terminal on lateral ferruginous, glandular-hairy
branches with sticky hairs; bracts lanceolate-linear, acute, ferruginous,
glandular-hairy. Capitula 1.5—2 cm long and 0.7—1 cm wide, on short
peduncles. Involucre imbricate, five-rowed; outer involucral bracts
ferruginous at base, ovate, with small, saber-shaped, erect or curved
appendage, glandular-hairy along margin and apex; inner bracts linear-
lanceolate, acuminate, densely lanate, sometimes glabrescent.

437

444 Receptacle almost conical, densely covered with linear-subulate, white-
lustrous, about 6 mm long scales, florets light pink, up to 1.2 cm long,
narrow part of tube about 6 mm long, as long as broad part including
lobes of limb; appendages of anthers hairy. Pappus double, outer pappus
bristles less numerous, serrate-scabrous, fragile, about 1 mm long, inner
ones long-plumose, ferruginous at base, whitish above, about 9 mm long.
Flowering VIIJ-IX.

Maple forests, meadow-covered slopes(?).—Far East: Ussuri
(Birobidzhan). Endemic. Described from the Sutar River Valley. Type
in Leningrad.

Note. Komarov did not compare his species with any other species.
In my view, S. splendida Kom. is a part of the group of Saussurea with
bicolored leaves, and should be placed in a separate series; it is similar
to S. controversa DC., from which it differs by larger leaves, the presence
of a strong paniculate inflorescence, comprising numerous individual
corymbs, pink (and not lilac-violet) florets, a dark brown pubescent stem,
sticky articulate hairs, glandular hairy outer involucral bracts, terminating
in a small saber-shaped appendage, and glandular-hairy lateral branches
of the inflorescence, bearing individual corymbs.

Series 6. Intermediae Kitam. in Acta Phyt. et Geo-bot. IV (1935)
8; Ej. in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII (1937) 151, 152.—
Capitula in compact corymbs, sessile in type specimen, less often
pedunculate. Involucre globose-campanulate; outer bracts attenuate or
acuminate, erect or recurved. Leaves usually pinnatisect, sometimes entire
or shallow large-toothed. Meadow and forest herbs.

50. S. elongata DC. in Ann. Mus. Paris, XVI (1810) 201; DC.
Prodr. VI, 534; Ldb. Fl. Ross. I, 2, 666; Turcz. Fl. baic.-dahur. II, 1,
109; Herd. in Bull. Soc. Nat. Mosc. XLI, 3, 11 saltem a. genuina and
B. polypodiifolia DC.; Kom. Fl. Manchzh. Il, 722 p. p.; Grubov, Konsp.
Fl. Mong. 275; Popov, Fl. Sr. Sib. II, 756.—S. polypodiifolia DC. in
Ann. Mus. Paris, XVI (1810) 201.—S. elongata 8. polypodiifolia DC.
Prodr. VI (1837) 534; Turcz. op. cit. 109.—S. scabiosaefolia Turcz. ex
Ldb. FI. Ross. II, 2 (1845-1846) 666, nomen.—S. elongata scabiosaefolia
Turcez. ex DC. Prodr. VI (1837) 534 nomen; Turcz. Fl. baic.-dah. IJ, 1,
110.—Serratula stelleri Steph. ex DC. Prodr. VI (1837) 534 nomen.—
Heterotrichum stelleri MB. ex DC. Prodr. VI (1837) 534 nomen.—Ic.:
DC. op. cit. (1810) tab. VI—Exs.: GRF No. 3991.

Perennial. Rhizome short, strong, with numerous fibrous roots; stem
25-80 cm (up to 1 m) high, solitary, erect, glabrous, smooth, sulcate,
simple or branched in inflorescence. Leaves, except upper, long-petiolate
(petioles narrow-winged); lamina irregularly divided, oblong-ovate or

445

438

oval, deeply pinnatisect in type specimen (var. polypodiifolia DC.), less
often almost entire or shallow large-toothed, primary lobes of deeply
pinnatisect leaves straight or sometimes weakly decurved, linear or
oblong, cuspidate, cusp cartilaginous, usually slightly convolute and often
crenate-toothed (teeth with cartilaginous cusp) or entire; all leaves green
and scabrous above, slightly grayish beneath, with more or less glandular
and curved hairs, particularly along prominent midrib and primary lobes
of leaf, as well as in sinuses between primary lobes; upper leaves sessile
or short-petiolate, linear-lanceolate or linear, predominantly entire, less
often pinnatifid. Capitula 1.5—1.8(2) cm long with florets, about 0.8 cm
wide, in compact corymb at stem apex or on lateral peduncles in
individual corymbs (mostly three in each), and then forming more or
less lax, paniculate-corymbose inflorescence. Involucre oblong, imbricate,
six- or seven-rowed; involucral bracts not recurved, usually blackish-
violet, less often light colored; outer bracts ovate, inner oblong; all bracts,
especially along margin and at apex, weakly pubescent, often glabrescent.
Receptacle densely covered with 0.7—0.8 cm long scales; florets pink,
about 1.2 cm long, narrow part of tube about 0.7 cm long, broad part of
tube including lobes of limb about 0.5 cm long; anthers violet, strongly
exserted, about 0.6 cm long, their appendages ciliate. Pappus dirty white,
double, outer pappus bristles fragile, about 0.5 cm long, inner ones about
1 cm long, long-plumose; achenes cylindrical, glabrous, about 0.4—0.5

-cm long. Flowering VII-—VIII.

Forest meadows, forests and their edges, stony screes and slopes,
limestone deposits.—Eastern Siberia: Angara-Sayans (Baikal Shore),
Dauria, Lena-Kolyma. General distribution: Northern Mongolia.
Described from Siberia. Type in Geneva.

Note. The form with distinctly pinnatisect leaves (var. polypodiifolia
DC.) is predominant in the Trans-Baikal Region. The leaves are
sometimes thicker and coriaceous, sometimes thinner. The capitula are
also variable in size and shape, sometimes being smaller and narrower.
A whole series of transitional forms for these characters is found. M.G.
Popov (op. cit. 1959) erroneously referred S. elongata DC., lacking
membranous appendages on the involucral bracts, to section Theodorea
DC.

51. S. recurvata (Maxim.) Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 374.—S. elongata B. recurvata Maxim. Prim.
Fl. Amur. (1859) 167; Herd. in Bull. Soc. Nat. Mosc. XLI, 3, 12; Nakai,
Fl. sylv. Korean: XIV, 117; Kitam. in Mem. Coll. Sc. Kyoto Univ. ser.
B, XIII (1937) 153.—S. elongata 6. pectinata Herd. op. cit. (1868) 12.—
S. elongata 5. glehniana Maxim. in Meél. Biol. IX, 3 (1874) 338.—S.
elongata auct. Fl. Ross.: Kom. Fl. Manzh. III (1907) 723 p. max. p.;

446

439

Kom. and Alis. Opred. Rast. Dalnevost. Kr. I, 1072.—S. polypodiifolia
auct. Fl. Ross.: Kom. and Alis. op. cit. (1932) 1071.—S. asperifolia Sch.
Bip. ex Herd. op. cit. 12, nomen.—S. pectinata 3. amurensis Maxim. op.
cit. 171.—Iec.: Kom. and Allis. op. cit. Plates 321, 323.

Perennial. Species extremely close to the previous one. It differs by
having involucral bracts that are recurved, often arachnoid-hairy or
subglabrous, sometimes narrowed from ovate or triangular base to more ~
or less long herbaceous saber-shaped, recurved cusp (cusp sometimes
toothed along margin), sometimes involucral bracts short-acuminate,
without saber-shaped cusp but with recurved apex. Leaves hard, also
strongly variable in serration, as in previous species from predominantly
undivided, toothed to medium and deeply pinnatisect; lamina triangular,
oval or oblong-ovate. Florets about 1.3 cm long, narrow part of tube
about 0.7 cm long, almost as long as broad part including lobes of
limb. Pappus double, outer pappus bristles short-plumose, fragile, about
2 mm long, inner bristles 1—-1.1 cm long, long-plumose. Achenes 0.5
cm long. Flowering VII—VIII.

Forests and flooded meadows, forests and their edges, shrubby
areas.—Far East: Zeya-Bureya, Uda, Ussuri. General distribution: Korea,
China. Described from Amur. Type in Leningrad.

Note. It is a Far Eastern race of S. elongata DC. Some specimens
characterized by having herbaceous, saber-shaped tips of the involucral
bracts are possibly of hybrid origin (hybrids with S. odontolepis Sch.
Bip. or S. umbrosa Kom.) or progeny of hybrids. At one time, they
were separated as S. elongata 5. pectinata Herd. or S. elongata 5.
glehniana Maxim. (S. recurvata (Maxim.) Lipsch. var. glehniana Lipsch.
comb. nova = S. elongata 5. glehniana Maxim. in Mel. biol. IX, 3
(1874) 338).

Series 7. Ussurienses Lipsch.—Ser. Acuminatae Kitam. in Acta Phyt.
et Geobot. IV (1935) 5 p. min. p.; Ej. in Mém. Coll. Sc. Kyoto Univ.
ser. B, XIII (1937) 159 p. min. p.—Ser. Imbricatae Kitam. in Acta
Phyt. et Geobot. IV (1935) 11 p. min. p.; Ej. in Mem. Coll. Sc. Kyoto
Univ. ser. B, XIII (1937) 198 p. min. p—Rhizome fibrous along length,
and branched (in respect of this character, it is closer to the section
Laguranthera (C.A.M.) Lipsch.). Capitula numerous, relatively smaller,
on short peduncles, in compact, corymbs on more or less long peduncles,
forming lax panicle. Involucre cylindrical, imbricate, arachnoid-hairy;
outer involucral bracts reduced, ovate, with very short cusp; inner bracts
subobtuse or obtuse. Pappus double, outer pappus bristles unequal,
persistent. Leaves coriaceous, pinnatifid, pinnatisect, or less often shallow
incised, lobes of lamina toothed. Usually tall herbs of meadows and
forests.

447

440

52. S. ussuriensis Maxim. in Mém. Acad. Sc. Pétersb. par. div.
sav. IX (1859) 167; Ej. in Mél. Biol. IX, 3, 340; Kom. Fl. Manchzh.
III, 739; Nakai, Fl. Korean. II, 43; Ej. in Bot. Mag. Tokyo, XXIX, 205;
Ej. Fl. sylv. Korean. XIV, 120; Ling. in Contr. Inst Bot. Nat. Acad.
Peip. II, 171; Kitam. in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII, 166
(excl. var. nivea Kitamura).—S. ussuriensis a. genuina Maxim. op. cit.
(1859) 168; Herd. in Bull. Soc. Nat. Mosc. XLI, 3, 13; Kom. op. cit.
740; Nakai in Bot. Mag. Tokyo, XXIX, 205; Kom. and Alis. Opred.
Rast. Dalnevost. Kr. II, 1072.—S. ussuriensis B. incisa Maxim. op. cit.
(1859) 168; Ej. in Mél. Biol. IX, 3, 340; Herd. op. cit. 13; Kom. FI.
Manchzh. III, 740; Nakai, op. cit. (1915) 205; Kom. and Alis. op. cit.
1071; Ling. op. cit. (1935) 171.—S. ussuriensis y. pinnatifida Maxim.
op. cit. (1859) 168; Ej. op. cit. (1874) 340; Kom. Fl. Manchzh. II, 741;
Ling. op. cit. (1935) 171.—Ie.: Kom. and Alis. op. cit. Plate 325.

Perennial. Rhizome prostrate, with numerous roots, fibrous, yellow.
Stems 30-100 cm high, densely leafy, erect, sulcate-angular, wingless,
sparsely pubescent or almost glabrous, usually branched above, branches
ascending. Leaves predominantly hard, sometimes softer, quite variable
in shape of lamina, especially in its incision, scabrous above, weakly
scabrous beneath due to short cilia, especially along veins, and with
numerous, sessile, lustrous glandular hairs; sometimes leaves tomentose
beneath (var. firmifolia Lipsch.); basal leaves persistent or withering
before flowering, like long-petiolate lower cauline leaves, petioles 4.5—
15 cm long, not winged; lamina oval or oblong, 7-18 cm long, long- or
short-acuminate, cordate at base, truncate or less often weakly hastate,
gradually reduced toward stem apex; all leaves gradually reduced upward;
middle cauline leaves short-petiolate, uppermost leaves sessile or almost
sessile, lanceolate, oblong-ovate, lanceolate-linear, acuminate, at base
truncate or cuneate, almost entire or toothed, sometimes serrate-toothed
with unequal spinescent teeth, or sometimes crenate-lobate, somewhat
convolute, lobes unequal, sometimes broader and shorter, or sometimes
narrower and long (var. incisa Maxim.), pinnatifid, with a gamut of
transitional forms between them, segments sometimes reaching up to
midrib, sometimes less deep, often segments at apex broader and more
or less deeply redivided, toothed (var. pinnatifida Maxim.). Capitula
numerous, relatively small, 4-14 mm long, on short peduncles in compact
corymbs, terminal on stem and branches, forming corymbose-panicle.
Involucre tubular, about 1 cm long and 4—6 mm wide; involucral bracts
numerous, imbricate, five-rowed, appressed, arachnoid-hairy, often
colored; outer bracts about 2 mm long, ovate, with short cusp; inner
bracts linear, obtuse. Florets dark red, about 1—1.2 cm long, narrow part
of tube about 5—6 mm long, as long as broad part including linear lobes
of limbs, latter about 3.5-4 mm long; appendages of anthers ciliate.

441

448 Receptacle densely covered with 5—6 mm long scales. Achenes about 5
mm long, 1—1.2 mm wide, with short corona at apex, weakly narrowed
toward base, slightly compressed, lustrous, densely black-purple punctate;
pappus double, outer pappus bristles up to 2 mm long, unequal, serrate,
fragile, but persistent, inner ones 7—8 mm long, long-plumose. Flowering
VII-IX.

Shrub thickets, oak forests, dry mountain slopes, flooded meadows.—
Far East: Ussuri. General distribution: Korea, Japan, China. Described
from Ussuri. Type in Leningrad.

Note. This species is close to S. elongata DC., from which it differs
by having many smaller capitula and a more densely arachnoid-hairy
involucre and by the nature of the involucral bracts (smaller, numerous,
cuspidate), the darker leaves, divided into broader lobes in the pinnatifid
varieties, and the teeth terminating in long cusps.

Apparently, it hybridizes with S. odontolepis Sch. Bip. I place here
the specimens from the floodplains of the Bei-Chi-Khe River, in the
vicinity of the village of Dukhovskoe (1913, No. 617, N.V. Shipchinski1).
The involucral bracts have a saber-shaped cristate-toothed appendage,
partly recurved.

The specimens from the limestone hill “Zmeinaya” in the Shkotovo
Region (1932, No. 26, D.P. Vorobev) are distinguished by leaves that
are more firm and strongly scabrous and arachnoid hairy-tomentose
beneath (var. firmifolia Lipsch; probably, this variety is identical with
S. firma (Kitag.) Kitam. in Acta Phyt. et Geobot. IX, 3 (1940) 112.
Specimens of which I have not seen). Additional field observations of
this variety are needed in order to make a final judgment about it as a
separate species. Cf. also note to S. maximoviczii Herd.

53. S. maximoviczii Herd. in Bull. Soc. Nat. Mosc. XLI, 3 (1868)
14; Maxim. in Mel. Biol. [X, 3, 337; Kom. Fl. Manchzh. III, 731;
Nakai in Bot. Mag. Tokyo, XXIX, 202; Ej. Fl. sylv. Korean. XIV, 119;
Kom. and Alis. Opred. Rast. Dalnevost. Kr. I], 1071; Kitam. in Acta
Phyt. et Geobot. IV, 11; Ej. in Mem. Coll. Sc. Kyoto Univ. ser. B,
XIII, 198 (excl. f. serrata (Nakai) Kitam. and var. triceps (Level. and
Vaniot) Kitam.).

Perennial. Rhizome woody, prostrate, hairy; stem 40-100 cm high,
erect, sulcate-angular, wingless, almost glabrous or weakly hispid,
scatteredly punctate, simple more or less branched at apex, branches
somewhat divergent, angular. Leaves pinnatifid, green above, more pale
beneath, scabrous on both sides, weakly hispid, their veins, especially
midrib, prominent; basal and lower cauline leaves persistent up to
blooming, up to 70 cm long and 15 cm wide, on 5—20 cm long petioles,
lyrate-pinnatisect, with four to six pairs of lateral, oblanceolate, cuspidate,

449

450

442

finely toothed lobes, teeth spinescent; terminal lobes usually large and
broad; leaf base narrow-cuneately decurrent; middle cauline leaves
smalier, with narrower lateral lobes and shorter petioles, some weakly
crenate or entire, sessile, semiamplexicaul; uppermost leaves smaller
and narrower, oblong-lanceolate or lanceolate, almost entire, cuspidate.
Capitula numerous, erect at anthesis, 5-7 mm wide, in corymbose-
paniculate inflorescence, individual corymbs on more or less long lateral
peduncles, with three to six compactly arranged capitula, sometimes
peduncles two-headed. Involucre narrow tubular, imbricate, about 1—1.4
cm long, about 6 mm wide; bracts coriaceous and hard when dry,
arachnoid-hairy especially along margin, usually colored, five-rowed;
outer involucral bracts reduced, ovate, with short cusp, obtuse-callose;
inner bracts obtuse. Receptacle with unequal, up to 7 mm long scales;
florets violet-purple, 1.1-1.3 cm long, narrow part of tube about 6—7
mm long, broad part including lobes of limb about 5—6 mm long, lobes
of limb 3—3.5 mm long; anthers about 5 mm long, appendages of anthers
woolly. Pappus slightly exserted from involucre, brownish, double, outer
pappus bristles short, unequal, up to 5 mm long, serrate, persistent,
inner ones about 1 cm long, long-plumose; achenes about 5 mm long,
1.5—2 mm wide, truncate at apex, weakly narrowed at base, achene
entirely black, glabrous, smooth. Flowering VII-—VIII.

Meadows (especially marshy meadows with clayey soils), dry
mountain herb slopes, oak forests, shrub thickets——Far East: Ussuri.
General distribution: China, Korea, Japan. Described from the Novik
Inlet on the Russkii Island. Type in Leningrad.

Note. One of the tallest and most typcial species of the genus in
the Soviet flora. It varies in the shape, width and length of the leaf
segments, and also of their terminal lobes, which are sometimes large
and broad, crowning the leaf, giving it a typical lyrate shape and
sometimes less differentiated from the lateral lobes, and then the habit
is similar to S. ussuriensis var. pinnatifida. In a number of forms S.
maximoviczii Herd. is closest to S. ussuriensis var. pinnatifida, from
which it differs by larger capitula, less densely arachnoid involucral
bracts (they are broader and.less acute), as well as by the woolly
appendages of anthers. Cf. above about the nature of leaves of both
species. It may be added that the base of the basal and lower cauline
leaves in S. maximoviczii is narrowly cuneate, tapering toward the petiole.

Series 8. Nupuripoensae Kitam. in Acta Phyt. et Geobot. IV, (1935)
8 emend. Lipsch., hoc loco; Ej. in Mem. Coll. Sc. Kyoto Univ. ser. B,
XIII (1937) 180.—Capitula relatively large, in compact corymbs,
subtended by approximate bracteal leaves. Lower and middle cauline
leaves coriaceous, ovate or deltoid, more or less oblong-cuspidate,

aD

wg re

we a.

Pe —— —

ZA

Lees

Plate XXXIV.
o-serrata Nakai; 2—S. parviflora

45]

(Poir.) DC.

452

444

cordate, sagittate, or truncate at base, lamina not crenate but fine or
large-toothed; petiole winged in type specimen. Stem more or less
winged; receptacle with short and sparse scales.

Note. In the interpretation of Kitamura (1937), this series includes
only one species—S. nupuripoensis. | find two more closely related
species in this series: S. kurilensis and S. riederi, and, therefore, change
the characters of the series. Kitamura considered the presence of villous
pubescence on the involucral bracts, which is actually present in S.
nupuripoensis but absent in two other species included in this series,
typical for it. However, based on the sum total of other characters and
the general appearance of the plants, these three species undoubtedly
form a single phylogenetic group.

54. S. nupuripoensis Miyabe and Miyake, Fl. Saghal. (1915) 277;
Kitam. in Acta Phyt. et Geobot. IV, 8; Nemoto, Fl. Jap. Suppl. 800;
Sugaw. Pl. of. Saghal. (1937) 312; Kitam. in Mem. Coll. Sc. Kyoto
Univ. Ser. B, XIII, 180; Honda, Nom. Pl. Jap. 368; Sugaw. III. Fl.
Saghal., IV, 1871.—Ic.: Sugaw op. cit. (1940) tab. 859.

Perennial. Rhizome thick, woody, oblique, developing numerous
fibrous roots; stem 20-30 cm long, erect, strong, sulcate, puberulent,
leafy, corymbosely branched above, branches ascending, densely hirsute,
with mixture of capitate glandular hairs. Basal leaves large, persistent
up to anthesis, like lower cauline leaves, long-petiolate, more or less
broadly decurrent; petioles 4-11 cm long, usually broad-winged; lamina
ovate or deltoid, 4-12 cm long, 4.5—8(10) cm wide, more or less oblong-
cuspidate, cordate, sagittate, or truncate at base, not crenate, with short-
cuspidate teeth, sinuses finely ciliate; all leaves coriaceous, green above,
lighter colored beneath, weakly pubescent on both sides; middle and
upper cauline leaves broadly ovate or oblong, gradually reduced upward,
short-petiolate or sessile, broadly decurrent. Capitula three to seven,
relatively large, on short and thick peduncles, in corymbs, often enclosed
in uppermost lanceolate bracteal leaves, as long as or longer than
capitulum. Involucre campanulate, roundish at base, about 1.5 cm long,
1.2—1.8 cm wide; involucral bracts five-rowed, outer and middle bracts
lanceolate, as long or almost as long as inner, bracts, acuminate, densely
hirsute along dorsal surface and margins. Receptacle with short scattered
scales; florets purple, about 1.1-1.2 cm long; narrow part of corolla
tube about 0.5 cm long, almost as long as broad part including lobes of
limb, lobes of limb unequal, almost half as long as broad part of corolla
tube; appendages of anthers ciliate. Pappus double, outer pappus bristles
unequal, short, up to 2-3 mm long, fragile, scabrous, finely sulcate or
short-plumose, inner ones about 1—1.1 cm long, dirty brown, long-
plumose, connate at base in ring; achenes narrow, 7 mm long and about

453

445

2 mm wide, glabrous, brownish, with indistinct purple stripes, slightly
compressed, truncate at apex, narrowed at base. Flowering VII—VIII.

Far East: Sakhalin. Endemic. Described from Nupuripo Mountain.
The type specimen was in the Herbarium of the Hokkaido University.

Note. Only one herbarium sheet of this species is preserved in the
Herbarium of the Botanical Institute, Academy of Sciences of the USSR.
The plant was described from the sheet with additions from Kitamura
(1937).

55. S. kurilensis Tatewaki in Journ. Fac. Agr. Hokkaido Univ.
XXIX, 5 (1933) 214; Kitam. in Acta Phyt. et Geobot. IV, 8; Ej.-in
Mem. Coll. Sc. Kyoto Univ. ser. B, XIII, 154.

Perennial. Rhizome thick, densely covered with brown bases of old
leaves; stem 10—20(40) cm high, fistular, erect, simple, sometimes
narrow-winged, densely leafy, sparsely arachnoid-hairy, more strongly
above, below inflorescence. Basal and lower cauline leaves long-petiolate,
“persistent until flowering, petioles narrow, 7-10 cm long, lamina hard,
deltoid or ovate, with long cusp, cordate or truncate at base, 4-7.5 cm
long, toothed, teeth terminating in callose obtuse cusp, green above,
pale beneath, flocculose-arachnoid hairy pubescent on both sides (var.
arachnoidea Tatevaki, op. cit. 238); middle and upper cauline leaves
similar in shape, gradually reduced, short-petiolate, sessile above;
uppermost leaves approximate with inflorescence. Capitula 1.2—-1.5 cm
wide, numerous, on short peduncles or sessile, terminal on stem in
compact leafy globules. Involucre globose-campanulate, 1.5—1.6 cm long,
1.1—1.2 cm wide, arachnoid-hairy; involucral bracts four-rowed; outer
bracts almost as long as middle, ovate-oblong, 3-4 mm wide, like middle,
attenuate-cuspidate, deflexed; middle bracts oblong, inner oblong-
lanceolate, acuminate. Florets pink, 1.1—-1.2 cm long, narrow part of
tube as long as broad part including lobes of limb, limb two and one-
half times as long as broad part of corolla tube. Pappus dirty brown,
0.9-1 cm long, double, outer pappus bristles fragile, short, inner ones
long-plumose; achenes cylindrical, blackish, 5.5—-6 mm long, with many
small ribs, slightly narrowed at base, glabrous. Flowering VIII. (Plate
XXXII, Fig. 1.)

Herb meadows.—Far East: Sakhalin (Kuril Islands). Endemic.
Described from Urup Island. Type preserved in the Hokkaido Herbarium.

Note. A small number of specimens of this species is preserved in
the Herbarium of Botanical Institute, Academy of Sciences of the USSR.
The above description of the species is based on these specimens,
supplemented with the data of Kitamura (1937).

56. S. riederi Herd. in Bull. Soc. Nat. Mosc. XLI, 3 (1868) 35;
Maxim. Prim. Fl. Amur. 167 in nota; Ej. in Mel. Biol. IX, 2, 343;

454

446

Kom. Fl. Kamch. III, 178; Hult..Fl. Kamtsch. IV, 209; Kitam. in Acta
Phyt. et Geobot. IV, 7; Ej. in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII,
170 (excl. subsp. yezoensis (Maxim.) Kitam. and variet.); Sugaw. III, fl.
Saghal. IV (1940) 1879.—Ic.: Hult. op. cit. fig. 9.

Perennial. Rhizome creeping or inclined, with numerous lateral roots,
covered at apex with hard bases of previous year’s leaves; stem (10)15-
30 cm high, erect, densely leafy, simple, terminating with dense corymbs
of capitula, sometimes winged due to decurrent leaves or wingless,
sulcate, puberulent with crisped hairs in upper part, especially in
inflorescence, almost glabrous below. Leaves succulent, ovate to almost
triangular, long-cuspidate, cordate or truncate at base; basal leaves usually
withering before flowering; lower cauline leaves long-petiolate, petioles
2.5-7 cm long, usually broad-winged, lamina 4-10 cm long and 2-6
cm wide, toothed, teeth short-cuspidate, green above, pale beneath,
puberulent on both sides; middle cauline leaves with short, winged
petiole, upper leaves sessile, decurrent with lamina, gradually reduced
and narrower. Capitula numerous, about 1—1.2 cm long and 0.7—0.8 cm
wide, in dense terminal compound corymb, individual corymbs,
subtended by less numerous bracteal leaves, usually longer or as long
as it. Involucre three- or four-rowed, turbinate-globose, arachnoid-hairy;
involucral bracts dark purple in upper part; outer bracts somewhat shorter
than inner, narrowed from ovate base to cusp, or lanceolate; inner bracts
oblong or broadly linear, apiculate, receptacle with scattered, linear-
subulate, up to 4.5 mm long scales; florets purple, about 1 cm long,
narrow part of tube as long as broad part including lobes, almost three
times as long as broad part of corolla tube. Pappus strongly exserted
from involucre, brownish, double, outer pappus bristles numerous, short,
unequal, up to 3 mm long, scabrous, short-plumose, shedding, inner
bristles long-plumose, about 8-9 mm long. Flowering VII—VIII. (Plate
XXXII, Fig. 2.)

Dryland and subalpine meadows, lichen tundra, sea coasts.—Far
East: Kamchatka (southern), Sakhalin (Kuril Islands). Endemic. Described
from Kamchatka. Type in Leningrad.

Note. A closely related race, subsp. yezoensis (Maxim.) Kitam. is
distributed on the Kuril Islands (from Urup to northern Honda). The
illustration given by Sugawara (op. cit., 1940, tab. 863) pertains to the
latter subspecies. According to Kitamura, S. riederi subsp. yezoensis
(Maxim.) Kitam. [Mem. Coll. Sc. Kyoto Univ. ser. B, XIII (1937) 169;
S. riederi var. yezoensis Maxim. in Mél. Biol. IX (1874) 343] is
characterized by a more narrowly winged stem and on involucre 8-11
mm long, always with many rows of bracts (four to six rows). This
subspecies is practically unrepresented in the herbaria of the Soviet Union
and is, therefore, obscure to me.

455

447

Series 9. Acuminatae Kitam. in Acta Phyt. et Geobot. IV (1935) 5
p. p.; Ej. in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII (1937) 159 p. p—
Capitula on short peduncles or sessile, in compact corymbs. Involucre
tubular; outer and middle involucral bracts imbricate, acute or cuspidate
with long herbaceous, saber-shaped cusp, sometimes acuminate.

Note. Series Acuminatae Kitam., in my view, is an unnatural
aggregate and should be divided into several independent series, as has
been done here.

57. S. acuminata Turcz. ex Fisch. and Mey. Ind. sem. hort. Petrop.
1 (1835) 37; Linnaea, X, Litt. Ber. 101; DC. Prodr. VI, 536; Ldb. FI.
Ross. II, 2, 667; Turcz. Fl. baic.-dahur. IJ, 1, 117; Herd. in Bull. Soc.
Nat. Mosc. XLI, 3, 23 (excl. B. sachalinensis and y. gracilis); Freyn. in
Oest. bot. Zeitschr. LII, 7, 281; Kom. Fl. Manchzh. III, 741; Kitam. in
Acta Phyt. et Geobot. IV, 7 (excl. subsp. sachalinensis Kitam.); in Mem.
Coll. Sc. Kyoto Univ. ser. B, XIII, 174 (p. p. excl. subsp. sachalinensis
Kitam.); Lipschitz in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XVI, 453; Popov. Fl. Sr. Sib. I], 763.—S. compacta Fisch. ex Herd. op.
cit. (1868) 23 (nomen).

Perennial. Rhizome inclined, developing numerous fibrous roots,
covered with hard remains of old leaves at apex; stem 25-60 cm (up to
1 m) high, erect, strongly leafy, sulcate, more or less narrow-winged
due to decurrent leaves, slighlty crisped-pubescent or glabrous; simple,
with one (rarely several) compact corymbs at apex. Leaves aeruginous,
glossy above, lighter colored beneath, glabrous, strongly scabrous due
to spiny appendages only along margin, weakly convolute, undivided or
distinctly toothed, then teeth not frequent, small; lamina oblong-lanceolate
or lanceolate, acuminate; basal leaves usually withering before flowering;
lower cauline leaves petiolate (with petiole 12—25(30) cm long) and up
to 2—2.5 cm wide; petioles 5—20 cm long; middle cauline leaves 10—20
cm long, short-petiolate or almost sessile, petioles or leaf bases decurrent;
all leaves gradually reduced upward; apical leaves, linear or linear-
lanceolate, narrowed. Capitula numerous, about 1.5-1.6 cm long,
including florets, about 0.5—0.6 cm wide, in compact terminal corymbs,
latter 4-6 cm wide when dry; less often, lateral corymbs also observed
besides central ones. Involucre cylindrical, four-rowed, arachnoid-hairy;
involucral bracts unequal, outer bracts green, ovate, narrowed from
middle in long, herbaceous, saber-shaped cusp; middle bracts narrowly
oblong, acuminate; inner bracts'linear or linear-lanceolate, dark colored,
distinctly acuminate. Receptacle densely covered with linear-subulate,
about 6 mm long scales; florets purple, 1.3-1.4 cm long, narrow part of
corolla tube about 8 mm long, broad part weakly developed, about 0.5—
1 mm long, lobes of limb 4—5 mm long, narrowly linear; appendages of

456

448

anthers ciliate. Pappus double, outer pappus bristles short, fragile, }
scabrous, short-plumose, inner up to 1.4 cm long. Flowering VII—VIII.
(Plate XXXII, Fig. 2.)

Note. According to Turczaninow, outer pappus bristles are almost
absent in the ray florets but evident in the disk florets; I did not observe
this.

Meadows, thickets of shrubs, birch forests.—Eastern Siberia: Dauria.
General distribution: Mongolia (northern), northeastern China. Described
from Dauria. Type in Leningrad.

58. S. sachalinensis Fr. Schm. in Mém. Acad. Sc. Pétersb. sér. VII,
XII, 2 (1868) 152; Miyabe and Miyake Fl. Saghal. 280; Lipschitz in
Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI, 453.—S. acuminata
6. sachalinensis Herd. in Bull. Soc. Nat. Mosc. XLI, 3 (1868) 23.—S.
acuminata subsp. sachalinensis Kitam. in Acta Phyt. et Geobot. IV (1935)
7, excl. S. yezoensis; Ej. in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII
(1937) 175; Honda, Nom. Pl. Japon. 366; Sugaw. Ill. fl. Saghal. IV,
(1940) 1877.—S. miyagii Miyabe and Kudo in Miyabe and Miyake, FI.
Saghal. (1915) 279.—Ie.: Sugaw. op. cit. (1940) tab. 862.

Perennial. Quite similar to preceding species. Stem densely
leafy up to apex. Distinguished by nonglossy leaves, glabrous above,
sparsely arachnoid-hairy beneath, more broadly lanceolate in type
specimen, distinctly serrate-crenate, teeth more or less cuspidate;
basal leaves large, about 34.5 cm wide, lanceolate, sometimes weakly
cordate at base, and with more or less long petiole. Outer and middle
involucral bracts usually (but not always!) with short, herbaceous,
saber-shaped apical appendages. Flowering VII—VIII. (Plate XXXIII,
Fig. 1.)

Montane, lowland and coastal meadows.—Far East: Sakhalin.
General distribution: Japan (Hokkaido Island). Endemic. Described from
Sakhalin (slopes near Dui). Type in Leningrad.

Note. It is a Sakhalin-northern Japanese race, close to S. acuminata
Turez., and morphologically is relatively weakly isolated. The characters
listed above in the description of S. sachalinensis, nevertheless, permit
us to consider this as a separate race. In my opinion, true S. acuminata
Turcz. is not found in Sakhalin, and specimens mentioned under this
name by Japanese authors, evidently not familiar with typical material
of S. acuminata Turcz., should be referred to S. sachalinensis Fr. Schm.
(compare papers of Kitamura, Sugawara, and others). S. miyagii Miyabe
and Kudo is known to me only from the figure in the publication of
Sugawara (op. cit. 1940); the plant shown in this plate leaves no doubt
about its affinity to typical S. sachalinensis Fr. Schm. Kitamura included
S. miyagii in the synonymy of S. acuminata Turcz., interpreting ssp.

457

449

sachalinensis Kitam. as a subspecies of the latter. Compare also the
note to S. duiensis Fr. Schm.

Series 10. Amurenses Lipsch.—Ser. Acuminatae Kitam. in Acta
Phyt. et Geobot. IV (1935) 7 p. p. and in Mem. Coll. Sc. Kyoto Univ.
ser. B, XIII (1937) 159 p. p—Capitula terminal on stem, usually in
corymbose panicles, less often corymbs solitary, dense, few-headed. Basal
leaves long-petiolate, large, broadly lanceolate, elliptical or ovate.
Involucre tubular, outer and middle bracts acuminate or subobtuse,
without recurved, saber-shaped appendages; inner bracts (also outer and
middle bracts in type specimens) sericeous-villous. Meadow and forest
herbs.

59. S. amurensis Turcz. in DC. Prodr. VI (1837) 534; Ldb. FI.
Ross. II, 2, 668; Turcz. Fl. baic.-dahur. II, 1, 112; Maxim. Prim. FI.
Amur. 170; Herd. in Bull. Soc. Nat. Mosc. XLI, 3, 27 excl. 5.
macrocephala Herd.; Kom. Fl. Manchzh. III, 719; Kom. and Alis. Opred.
Rast. Dalnevost. Kr. II (1932) 1063; Kitam. in Mem. Coll. Sc. Kyoto
Univ. ser. B, XIII, 160 excl. subsp. duiensis (Fr. Schm.) Kitam.; Lipschitz
in Sp. Rast. Gerb. Flory SSSR, XIII, 124; Popov in Sp. Rast. Gerb.
Flory SSSR, XIV, 30; Popov, FI. Sr. Sib. II, 759.—S. denticulata Ldb.
Ic. pl. Fl. Ross. I (1829) 18, tab. 74; Ej. Fl. alt. IV, 27 (in nota); Ej. FI.
Ross. II, 2, 668; Herd. in Bull. Soc. Nat. Mosc. XLI, 3, 24, saltem a.
genuina (excl. B. duiensis and y. viluiensis), non S. denticulata Wall.—
S. stenophylla Freyn in Oest. Bot. Zeitschr. LII, 7 (1902) 280; LII, 8
(1902) 313.—? S. amurensis var. stenophylla Nakai in Bot. Mag. Tokyo
XLV (1931) 513 excl. syn.—S. amurensis subsp. stenophylla Kitam. in
Acta Phyt. et Geobot. IV (1935) 6.—S. odontophylla Freyn, op. cit. LI,
7 (1902) 282 (in clave) and LII, 8 (1902) 315.—S. intermedia Freynn,
op. cit. LII, 7 (1902) 280 (in clave) and LII, 8 (1902) 317 p. p.—sS.
amaurolepis Freyn, op. cit. LII, 7 (1902) 280 (in clave); LII 8 (1902)
312.—Ic.: Kom. and Alis. op. cit. (1932) Plate 318.—Exs.: GRF Nos.
3989b, 4049.

Perennial. Rhizome slender, not long, creeping; stem (30)50-
100(115) cm high, loosely flocculose-arachnoid hairy or almost glabrous,
sulcate, sometimes rather distinctly narrow-winged due to decurrent bases
of cauline leaves, simple, sometimes with floriferous branches only
above. Leaves hard, glabrous above, snow-white-tomentose or arachnoid-
woolly beneath in typical specimen, pubescence quite variable beneath,
sometimes weakly pubescent; all leaves somewhat involute, cuneate at
base, long petiolate, large, broadly lanceolate or ovate, more or less
large-toothed, usually withering before full bloom; lower cauline leaves
narrower and usually with shorter petiole, lanceolate, linear-lanceolate

450

or linear, sometimes more or less horizontal, reduced and narrower
upward; middle cauline leaves petiolate, with indistinct teeth, upper
sessile, linear-lanceolate or linear, entire or with inconspicuous solitary
teeth; all leaves decurrent on stem. Capitula tubular, in individual
corymbs, sometimes aggregated in more or less dense compound corymb,
at same height, sometimes few lateral axillary corymbs also present
besides compound corymb, sometimes inflorescence paniculate. Involucre
four- or five-rowed, imbricate, in typical specimen woolly-villous; outer
involucral bracts ovate, green or dark-colored, subobtuse or with short
cusp; inner bracts lanceolate or oblong-lanceolate, subobtuse, densely
villous, especially at apex, often glochidiate. Florets pink-violet when
dry, about 1.1—1.2(1.3) cm long, narrow part of tube 5—6 mm long,
broad part about 1.5—2 mm long, lobes of limb linear, with lustrous
glandular hairs, about 4 mm long; anther appendages ciliate. Receptacle
densely covered with scales; scales linear-subulate, glossy, about 5—6
mm long; pappus double, outer pappus bristles about 2-3 mm long,
fragile, less numerous, toothed scabrous, inner 0.9—1 cm long, numerous,
long-plumose. Young achenes glabrous. Flowering VI—VIII.

Lowland, montane and forest meadows; deciduous, coniferous, and
mixed forests; forest edges; borders of marshes.—Eastern Siberia:
Angara-Sayan, Dauria, Lena-Kolyma; Far East: Zeya-Bureya, Ussuri,
?Uda. General distribution: Northeastern China. S. amurensis Turcz.
described from grassy meadows along Amur River. Type in Leningrad.
S. stenophylla Freyn and S. odontophylla described from Zeisk. Their
types evidently are preserved in the Herbarium of Freyn in Brno(?). S.
amaurolepis Freyn described from the vicinity of Blagoveshchensk;
apparently the type is preserved there as well and an isotype in Leningrad.
S. denticulata Ldb. described from collections of Krause in eastern
Siberia, without precise indication of location. Type in Leningrad.

Note. An unusually polymorphic species, which was reported in a
paper by Freyn, V.L. Komorov, M.G. Popov and S.Yu. Lipschitz. The
shape and width of the leaves, pubescence on their lower surface, color
and pubescence of the involucral bracts, shape of inflorescence, etc., all
vary. This provided a basis for Freyn (op. cit.) to separate and describe
11 subspecies and forms of-S. amurensis, 7 of them with binominal
names. Most of the “species” and “forms” described by Freyn do not
deserve recognition as they represent descriptions of individual specimens,
apparently collected from a hybrid complex. Only S. dubia Freyn (cf.
note to this species), in my opinion, deserves recognition. Comparison
with the authentic specimen of S. denticulata Ldb. showed that this
species should be included under S. amurensis Turcz. Ledebour himself
(Fl. Ross. II, 2, 668), in his note to S. amurensis, writes that S. denticulata
is close to it, but distinguishes S. amurensis by the presence of much

45]

wider lower leaves in comparison with the upper ones, with all the
involucral bracts being obtuse and densely tomentose. At the same time,
the lower part of the stem with leaves is absent in the authentic specimen
of S. denticulata, but involucral bracts are also subobtuse, and most of
them are densely tomentose.

60. S. duiensis Fr. Schm. in Mém. Acad. Sc. Pétersb. VII sér. XII,
2 (1868) 152; Sugaw. Ill. Fl. Saghal. [IV (1940) 1875.—S. denticulata
B. duiensis Herd. in Bull. Soc. Nat. Mosc. XLI, 3 (1868) 25.—S.
amurensis subsp. duiensis (Fr. Schm.) Kitam. in Acta Phyt. et Geobot. -
IV (1935) 6; Kitam. in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII, 161;
Honda, Nom. Pl. Jap. 366.—S. susuyensis Miyabe and Miyake ex Sugaw.
Ill. Fl. Saghal. TV (1940) 1875 nomen.—Ic.: Sugaw., op. cit. (1940)
tab. 861.

Perennial. Stem 30—-60(100) cm high, erect, simple, sometimes
weakly branched only above, loosely flocculose-arachnoid-hairy or almost
glabrous, sulcate, less often weakly narrow-winged due to decurrent
leaves; fistular. Leaves dark green and weakly scabrous above, smooth,
sparsely pubescent beneath, with isolated sessile lustrous glandular hairs
and more prominent midrib, lateral veins weakly prominent; sparsely
crenate-toothed, with cuspidate teeth, less often almost entire; lower
leaves long-petiolate, oblong-lanceolate or broadly lanceolate, up to 15-
25 cm long with petiole and up to 4 cm wide, petioles 4.5—7.5 cm long,
long acuminate, abruptly narrowed at base into winged petiole; middle
cauline leaves oblong-linear or lanceolate, narrower, short-petiolate, more
or less decurrent on stem; upper cauline leaves linear-lanceolate to linear,
sessile, weakly decurrent. Capitula about 1-1.2 cm long with florets
and about 4-5 mm wide, in dense, almost leafless corymbs at stem
apex and lateral floriferous branches, often forming corymbose panicle.
Involucre four-rowed; involucral bracts pigmented, sometimes light
colored or green, arachnoid-hairy, glochidiate at apex; outer bracts
smaller, ovate, usually acuminate, without recurved saber-shaped
appendages, less often obtuse; inner bracts oblong-lanceolate, somewhat
obtuse, almost as long as pappus. Florets about 1-1.1 cm long, narrow
part of tube about 4-5 mm long, broad part 1-1.5 mm long, lobes of
limb about 3-4 mm long; anther appendages hairy. Receptacle densely
covered with white, short, about 5 mm long scales; pappus double, outer
pappus bristles about 2 mm long, fragile, scabrous; inner bristles long-
plumose, about 8-8.5 mm long, somewhat exserted from involucre.
Flowering VII-VIII.

Forests and forest edges, thickets of shrubs, meadows.—Far East:
Sakhalin. Endemic. Described from mountains in the vicinity of Dui.
Type in Leningrad.

459

452

Note. The author of this species correctly pointed out in his original

description that S. duiensis is hardly distinct in its appearance. He

compared his own species with S. acuminata Turcz. and S. sachalinensis
Fr. Schm. As is known, the latter is considered by many Japanese
botanists as a subspecies of S. acuminata. S. duiensis is also very close
to S. amurensis Turcz. It differs at once from the latter by the green
(and not snow-white-tomentose) lower surface of the leaves. S. duiensis
sharply differs from S. acuminata Turcz. by the absence of saber-shaped
appendages on the involucral bracts. The relationship between S. duiensis
and S. sachalinensis Fr. Schm. is more complex. The most reliable
characters distinguishing the former from the latter are: less densely
leafy stem, less distinct dentation of leaves which are narrower in the
type specimen of S. duiensis; less leafy corymbs; short-pointed outer
involucral bracts not extended into saber-shaped appendages (which is a
characteristic feature of S. acuminata and S. sachalinensis). A few
specimens are found among the Sakhalin materials of S. duiensis and S.
sachalinensis which spoil the “purity” of both species. These specimens,
identified as S. sachalinensis, have almost entire or weakly serrate-dentate
leaves, and also less marked saber-shaped appendages than on the
involucral bracts of typical specimens of S. sachalinensis. These
specimens are close to S. duiensis in this respect. Most probably, it
would be correct to consider these specimens as hybrids between the
two species. Compare also the note to the next species.

61. S. shiretokoensis Sugaw. in Plants of Saghal. (1937) 312
(nomen); Ej. Fl. E. and N. Mit.-rang. S. Saghal. 75 (opusc. non v.);
Honda, Nom. Pl. Jap., 369; Sugaw. Ill. Fl. Saghal. IV, 1881.—Ie.:
Sugawara op. cit. (1940) tab. 864.

Perennial. Rhizome creeping; stem 20-60 cm high, glandular-
pubescent especially in upper part, or almost glabrous, weakly leafy.
Simple, cauline leaves distinct, sometimes weakly decurrent, lanceolate,
with short apical cusp, flat along margin, shallow-toothed with teeth
terminating in thin cusp, concolorous, weakly pubescent, with scattered
sessile, glossy glandular hairs above; lower and middle cauline leaves
with truncate base, narrowed into petiole, which is as long as lamina or
shorter; upper cauline leaves sessile or short-petiolate; uppermost leaves
sessile, narrower and shorter, linear or linear-lanceolate, almost entire.
Capitula narrow-cylindrical, about 1.3-1.4 cm long including florets,
and about 4-5 mm wide, on peduncles in individual three-headed
corymbs, forming terminal, dense, almost leafless corymbose
inflorescence at apex; besides main compressed corymb, lateral axillary
corymbs may be present. Involucre imbricate, four-rowed; involucral

460

453

bracts green, violet along margin, entirely sericeous-villous; outer bracts
broadly ovate, attenuate in dark, more or less short cusp at apex, but
not with saber-shaped herbaceous appendage; inner bracts longer,
subacute, glochidiate. Florets pink-purple(?), about 1.1 cm long, narrow
part of tube about 5 mm long, as long as broad part with lobes of limb;
anther appendages hairy. Receptacle densely covered with white, linear-
subulate, glossy, about 5 mm long scales; pappus double; outer pappus
bristles 2-3 mm long, serrate, fragile, inner 8—9 mm long, long-plumose.
Flowering VII—VIII. .

Forests (larch), meadows.—Far East: Sakhalin (endemic). Described
from Sakhalin Island. Location of type specimen not known.

Note. The species is known to me only from the figure in the cited
publication of Sugawara (1940) and only on the basis of one specimen
from southern Sakhalin (S.M. Kravchenko, No. 392), which is preserved
in the Herbarium of the Botanical Institute, Academy of Sciences of the
USSR. The above description of S. shiretokoensis Sugaw. is prepared
from these two sources, because, evidently, the species diagnosis was
not published in Latin by Sugawara, and the publication in Japanese is
not available to me. In all probability, further study of S. shirotokoensis
Sugaw., including the type material, will lead to reducing it to synonymy
in S duiensis Fr. Schm. The possibility of the hybridization of S. duiensis
with S. sachalinensis and the presence of several intermediate forms
between them in the Herbarium of the Botanical Institute, Academy of
Sciences of the USSR was mentioned in the note to the description of
the previous species. It appears that in the spread of the hybrid segregates
between these species, some specimens inherited a larger number of
characters from S. sachalinensis; consequently, I refer them to this
species. Other specimens show an inclination towards S. duiensis in
their characters and, therefore, are included under this species. At present,
it appears reasonable to accept S. shiretokoensis Sugaw. as a taxon
extremely close to S. duiensis Fr. Schm., possibly, arising as a result of
its hybridization with S. sachalinensis; this conclusion is confirmed by
the shape of the outer involucral bracts and the presence of a cusp on
the outside of the bracts as well as by the toothed margin of leaves.

Series 11. Umbrosae Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 375.—Ser. Acuminatae Kitam. in Acta Phyt.
et Geobot. IV (1935) 5 p. min. p.; Ej. in Mem. Coll. Sc. Kyoto Univ.
ser. B, XIII (1937) 159 p. min. p—Ser. Tohiren Kitam. op. cit. (1935)
8 p. min. p.; Ej. op. cit. (1937) 181 p. min. p., excl. syn—Stem distinctly
winged. Capitula campanulate, sometimes large, sometimes small, on .
long peduncles, in broad corymb or panicle. Outer involucral bracts

46

—

454

ovate or triangular-ovate, attenuate into more or less long, often colored
apical cusp (saber-shaped appendage), usually recurved. Leaves,
especially basal, large, more or less toothed. Forest and meadow plants.

62. S. dubia Freyn in Oest. Bot. Zeitschr. LII, 7 (1902) 282 (in
clave); LII, 9 (1902) 349 (descriptio); Lipschitz in Sp. Rast. Gerb. Flory
SSSR, XIII, 124, in. textu; Karavaev, Konsp. Fl. Yakutii, 176.—S.
intermedia Freyn. op. cit. (1902) 280 (in clave); LII, 8 (1902) 317
(descriptio) p. p. non Turcz. (1832).—S. virescens Freyn, op. cit. LII, 7
(1902) 283 (in clave); LII, 9 (1902) 347 (descriptio)—S. contracta Freyn,
op. cit. LII, 7 (1902) 283 (in clave); LH, 9 (1902) 348 (descriptio).—?
S. zeaensis Freyn, op. cit. LII, 7 (1902) (in clave); LII, 8 (1902) 314
(descriptio).—S. rosea Kom. in Bot. Mat. Bot. Inst. Akad. Nauk SSSR,
VI, 1 (1926) 17.

Perennial. Stem 25—80 cm high, erect, angular, pubescent with curly
hairs, distinctly winged due to decurrent leaves, wings green, about 1—2
mm wide. Basal and lower cauline leaves often withering before anthesis,
broad-lanceolate, ovate or oblong-lanceolate, up to 30 cm long with petiole
and up to 6 mm wide, weakly cuneate or somewhat roundish at base,
narrowed into more or less long, slightly winged petiole, midrib of lamina
prominent, lateral veins less distinct; all leaves unequally toothed, teeth
with thin cusps; sparsely pubescent above, arachnoid-hairy beneath, less
often weakly pubescent, and throughout with fine, lustrous, sessile
glandular hairs; cauline leaves reduced upward, lanceolate, linear-
lanceolate, or linear, sessile, decurrent, usually entire, somewhat
horizontally divergent or slightly falcate. Inflorescence more or less dense,
many-headed panicle (up to 15 cm long), consisting of individual corymbs,
terminal on lateral branches, upper peduncles with corymbs of capitula
usually approximate, lower peduncles usually distant 3-10 cm long,
slightly winged, arachnoid-hairy, panicle reduced in many specimens to
terminal, more or less large corymb and fewer poorly developed lateral
branches, terminating in solitary or less numerous capitula. Capitula in
individual corymbs on 1—2 mm, less often up to 5 mm long peduncles,
campanulate, rather small. Involucre about 0.8—0.9 cm long and about 0.6—
0.7 cm wide, -arachnoid-hairy; outer rows of involucral bracts attenuate
into more or less long saber-shaped, herbaceous or dark-colored, usually
recurved apical appendage; inner bracts longer, acuminate, but usually
without appendage. Receptacle densely covered with scales, scales linear-
subulate, snow-white or sulfur-yellow, lustrous, about 5 mm long; florets
pink or purple(?), about 1 cm long, narrow part of tube about 5 mm long,
broad part covered with lustrous glandular hairs, about 2 mm long, lobes
of limb 2.5—3 mm long; anther appendages ciliate; pappus one-rowed or
double, in latter case outer pappus bristles less numerous, fragile, serrate,

462

455

about 1 mm long, inner bristles long-plumose, about 8 mm long. Achenes
about 3 mm long, glabrous, weakly narrowed toward base, light brown,
with whitish thin ribs. Flowering VII-IX.

Forests, shrubs, meadows.—Eastern Siberia: Lena-Kolyma, Dauria
(eastern part); Far East: Zeya-Bureya. Ussuri, ?Uda., Okhotsk. S. dubia
Freyn is described from Zeisk; type specimen, evidently, in the
Herbarium of Freyn in Brmo(?), S. intermedia Freyn described from Zeisk.
The type specimen is probably also there. S. virescens Freyn and S.
contracta Freyn described from Zeisk; isotypes in Leningrad; S. zeaensis
Freyn from Zeisk (from a single plant), the type specimen evidently in
the Herbarium of Freyn; S. rosea Kom. described from maritime province
(Grossevichi); type in Leningrad.

Note. Freyn’s series of “species,” the typical characters of which
are a campanulate capitulum and the presence on the outer involucral
bracts of green (herbaceous) or colored, saber-shaped, usually recurved
appendages, was found to be quite widely represented in the Far East
and in Yakutia. These “species” essentially represent only one taxon,

.for which I accept the name Saussurea dubia Freyn and include the

later described Saussurea rosea Kom. among its synonyms. To me S.
dubia Freyn appears to be a hybrid species; apparently, it evolved as a
result of hybridization between S. umbrosa Kom. (see below) and S.
amurensis Turcz. The recurved, saber-shaped appendages of the
involucral bracts were inherited from S umbrosa Kom. and the
pubescence of the lower leaf surface, as well as the smaller size of the
capitulum, nature of the pubescence on the involucral bracts, etc. from
S. amurensis. S. dubia, referred by me to series Umbrosae,
morphologically connects this series with series Amurenses. The later
species S. rosea Kom. was compared with S. dubia Freyn. V.L. Komarov
distinguished his species (S. rosea) from S. dubia by the leaves which
are arachnoid-hairy beneath with glandular hairs (the first character was
noted by Freyn also for S. dubia, while he was silent about the second,
but it is present in all the material studied by me), a smaller capitulum
and snow-white receptacular scales (for S. dubia Freyn states that the
receptacular scales are sulfur-yellow; it is more appropriate to describe
them as lustrous; according to my observations, based on the examination
of the receptacular scales of capitula from different areas of the range
of S. dubia, the scales are sometimes snow-white, sometimes sulfur-
yellow). Thus, there is no doubt about the identical nature of S. rosea
Kom. and S. dubia Freyn.

63. S. umbrosa Kom. in Tr. Peterb. Bot. Sada, XVIII, 3 (1900)
423; Kom. Fl. Manchzh. III, 739; Nakai in Bot. Mag. Tokyo, XXIX,
200 (excl. var. herbicola Nakai); Ej. Fl. sylv. Korean. XIV, 120; Kom.

463

456

and Alis. Opred. Rast. Dalnevost.. Kr. II, 1064; Kitam. in Acta. Phyt. et
Geobot. IV, 10; Ej. in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII, 192
(excl. var. herbicola Nakai); Popov, Fl. Sr. Sib. I, 758.—Saussurea
karoi Freyn in Oest. bot. Zeitschr. LI, 7 (1902) 279 (in clave); LII, 8
(1902) 310 (descriptio); Kom. Fl. Manchzh. III, 743; Kom. and Alis.
Opred. Rast. Dalnevost. Kr. II (1932) 1064; Lipschitz in Bot. Mat. Gerb.
Bot. Inst. Akad. Nauk SSSR, XVI, 453.—Ie.: Kom. op. cit. (1907) Plate
XVI.

Perennial. Rhizome creeping, somewhat thick, producing fibrous
roots; stem 20—75(100) cm high, with remains of old leaves at base,
erect, angular, sulcate, leafy, more or less winged, sometimes more or
less densely covered with hyaline, curled, articulated hairs (especially
above), sometimes almost glabrous, smooth, simple, branched at apex,
branches weakly bent or upward spreading. Leaves dark green above,
pubescent with curly hairs, particularly distinctly along margin, besides
appressed hairs, lower surface light colored, with prominent midrib and
less distinct lateral veins, pubescent with brown hairs along veins, glabrous
in remaining part, thin, sometimes thick and hard, all unequally toothed,
teeth cuspidate; basal leaves usually withering before anthesis, largest
20 cm long and 6.5 cm wide, less numerous, rosulate, elliptical, oblong-
lanceolate or broadly lanceolate, with almost roundish or somewhat
cordate, sometimes weakly truncate at base, narrowed into short winged
petiole; lower leaves like basal, oblong, oblong-lanceolate or lanceolate;
middle cauline leaves oblong-lanceolate or lanceolate; 9—-12(18) cm long
and 2.5—3 cm wide, sessile, decurrent with more or less broadly toothed
or entire wing; upper cauline leaves gradually reduced, lanceolate,
acuminate, entire or toothed, sessile. Capitula campanulate, lax, terminal
on stem or branches, in groups of three in individual corymbs, sometimes
two each on 2-6 mm long peduncles on lateral 1.5—3.5 cm long branches,
pubescent with brown partly glandular hairs, sometimes one each on
rather long, 3.5—6 mm long, branches, all together forming lax panicle.
Capitula with 1.7—2 cm long and about 1—1.2(1.4) cm wide flowers;
involucre campanulate, roundish at base, 1.1—1.3 cm long and about 1-
1.2 cm wide when dry, sometimes weakly arachnoid-hairy, glandular-
pubescent, glandular hairs on short stalks, sometimes almost glabrous;
involucral bracts four-rowed; outer bracts often reaching middle of inner
ones, ovate, acuminate into saber-shaped, usually dark colored, recurved
appendage; middle bracts, ovate, straight or recurved; inner bracts linear
of lanceolate, acuminate. Receptacle hemispherical, densely squarrose,
scales linear-subulate, lustrous, about 4—5(6) mm long. Florets dark violet,
about 1.1 cm long; narrow part of tube about 5 mm long, broad part 1-
1.5 mm long, lobes of limb linear, about 4-5 mm long; anther appendages

464

457

ciliate-hairy. Pappus double, dirty brown; outer pappus bristles short,
about 3—4 mm long, scabrous, short-plumose or finely barbate; inner
bristles 9-10 mm long, long-plumose. Immature achenes about 3 mm
long, dark brown, smooth, lustrous. Flowering VII-—VIII.

Shady places in mixed forests, moss-covered forests——Eastern
Siberia: Dauria (eastern part); Lena-Kolyma (Olekma, rarely); Far East:
Zeya-Bureya, Ussuri. General distribution: Korea. S. karoi Freyn
described from the region of the Zeisk dock; isotype in Leningrad. S.
umbrosa Kom. described from northeastern China and Korea; type in
Leningrad.

Note. Freyn (op. cit. 1902) did not compare his species S. karoi
with S. umbrosa Kom., described in 1900. Groundlessly, he compared
S. karoi with S. latifolia Ldb., a species without appendages on the
involucral bracts and phylogenetically distant from the former. In
publications of 1907 and 1932, V.L. Komarov recognized both species
(S. umbrosa and S. karoi) in the Far East. He distinguished them as
follows: in S. karoi supposedly the lower cauline leaves are petiolate
with a cordate or broadly cuneate base, but in S. umbrosa they are
sessile, uniformly narrowed towards base. A comparison of the authentic
specimens and study of materials of both species show that these
characters are not stable, and both species should be combined under
the prior name S. umbrosa Kom.

Series 12. Grandifoliae Lipsch.—Ser. Tohiren Kitam. in Acta Phytot.
et Geobot. IV (1935) 8 p. p.; Ej. in Mem. Coll. Sc. Kyoto Univ. ser. B,
XIII (1937) 181 p. p—Capitula less numerous, large, often solitary, on
long peduncles, in lax corymbs, sometimes forming lax panicle if
corymbs several, occasionally in racemose inflorescence. Involucre
globose-campanulate or turbinate; outer involucral bracts ovate or
lanceolate, with or without long, saber-shaped, less often recurved
appendages. Tall meadow or forest herbs usually with large, serrate
toothed or crenate leaves, almost triangular in type specimen; leaf base
truncate, slightly cuneate, or weakly cordate; stem wingless.

64. S. grandifolia Maxim. in Mém. Acad. Sc. Pétersb. par div. sav.
IX (1859) 169; Ej. in Mél. Biol. IX, 342 p. max. p.; Herd. in Bull. Soc.
Nat. Mosc. XLI, 3, 15; Kom. Fl. Manchzh. III, 725 excl. ¢. caudata
(Herd.) Kom.; Nakai, Fl. sylv. Korean. XIV, 118; Kom. and Alis. Opred.
Rast. Dalnevost. Kr. II, 1072; Kitam. in Acta Phyt: et Geobot. ITV (1935) -
9; Ej. in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII, 186 excl. var. caudata
(Herd.) Kom. and var. microcephala Nakai.—S. grandifolia a. genuina
seu. Maximovicziana Herd. op. cit. (1868) 15 nom. nud.; Kom. FI.

458

Manchzh. III, 726; Nakai in Bot. Mag. Tokyo, XXIX, 203.—S.
grandifolia B. tenuior seu Raddeana Herd. op. cit. (1868) 16; Kom. op.
cit. 726.—S. grandifolia y. asperifolia Herd. op. cit. (1868) 16; Kom.
op. cit. 726.—S. coarctata (Herd.) Kitam. in Iconogr. pl. As. Orient. V,
1 (1950) 461, tab. CXLII.—S. grandifolia 5. coarctata seu wilfordiana
Herd. op. cit. 16; Kom. op. cit. 726.—Ic.: Kom. and Alis. op. cit. Plate
324.

Perennial. Rhizome prostrate, with numerous roots; stem ai
cm high, erect, more or less leafy, wingless, sulcate-angular, more or
less rough due to stiff hairs, especially below, or almost glabrous, simplg,
branched at apex, branches divergent or weakly bent, puberulent. Leaves
in typical specimen hard, dark green above, light colored beneath, both
sides pubescent, scabrous, variable in shape; basal leaves withering before
anthesis; lower and middle cauline leaves almost ovate or deltoid,
attenuate-long cuspidate, more or less deeply cordate or truncate at base,
narrowed into 3—9(13) cm long petiole; lamina 8-20 cm long and 4.5—
13. cm wide, toothed-serrate; teeth cuspidate, somewhat larger toward
base of lamina; upper leaves reduced upward, short-petiolate or sessile,
ovate-triangular, ovate-rhombic or lanceolate, often cuneate at base, long
acuminate; bracteal leaves 24 cm long, linear. Capitula up to 2 cm
long with florets and about 1.2—-1.4 cm wide, 3-18, solitary or in groups
of two or three on lateral, more or less long peduncles, forming broad
corymb or panicle, sometimes in racemes. Involucre campanulate,
involucral bracts numerous, five- or six-rowed, imbricate, appressed,
arachnoid-hairy dorsally and along margin, often with dark border; outer
bracts one-fifth to one-fourth as long as inner, ovate, like middle short
acuminate with blackish cusps; middle bracts oblong, inner linear,
acuminate like middle, dark brown at apex and flocculose. Receptacle
densely squamous, scales linear-subulate, white, lustrous, 7-8 mm long.
Florets dark pink, 1.3(1.5) cm long; narrow part of tube about 0.7 cm
long, almost as long as broad part with lobes of limb; anthers prominent,
about 0.5 cm long, anther appendages ciliate. Pappus whitish, turning
yellow at base, double; outer pappus bristles short, usually less numerous,
scabrous or finely plumose, fragile; inner bristles long-plumose, 10—11
mm long. Flowering VII-VIII.

Forests, forest edges, rock crevices, shrubby thickets——Far East:
Zeya-Bureya (southeast), Ussuri. General distribution: Korea, northeastern
China. Described from Amur (north slopes in the Amur gaps through
the Burei Mountains). Type in Leningrad.

Note. A highly variable species; depending on ecological conditions,
the consistency of leaves is sometimes more firm and coriaceous, as in
plants of drier and open habitats, and sometimes softer, more delicate,
as in specimens collected from shady places. The shape of the

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inflorescence is also quite variable. Forms with few capitula may be
segregated in var. oligocephala Lipsch. A plant preserved in the

_Herbarium of Botanical Institute, Academy of Sciences of the USSR

(Loonezala Mountain in southern Sikhote-Alin, No. 1054, I.K.
Schischkin), was taken to be a new species, S. petiolata Kom. (in Herb.),
by V.L. Komarov. This lone specimen is very close to S. grandifolia
Maxim., to the shade forms of which I tentatively refer it for the present,
until new, more abundant material becomes available. The specimen
identified by V.L. Komarov as S. petiolata Kom. differs by leaf blades
that are up to 14 cm long and 11 cm wide on petioles up to 15 cm
long; smaller and narrower, densely borne capitula; inner involucral bracts
more heavily pubescent at the tip, with brushes of woolly hairs; and
smaller florets.

65. S. subtriangulata Kom. in Tr. Glavn. Bot. Sada, XXXIX, 1
(1923) 119 (nomen); Bot. Mat. Gerb. Bot. Sada, VI (1926) 18
(descriptio); Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1072.—S.
eriolepis y. caudata Herd. in Bull. Soc. Nat. Mosc. XLI, 3 (1868) 32.—
S. grandifolia var. caudata (Herd.) Kom. Fl. Manchzh. III (1907) 727;
Kitam. in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII, 187.—S. grandifolia
Maxim. in Mél. Biol. IX (1874) 342 p. p.—S. subhastata Kom. nomen
in Herb.—Iec.: Kom. and Alis. op. cit. Plate 322.

Perennial. Rhizome short, oblique, with numerous lateral roots; stem
(30)40—75(100) cm high, wingless, erect, almost glabrous, smooth or
sparsely pubescent, somewhat scabrous, sulcate-angular, leafy. Leaves
thin, tender, dark green above, light-colored beneath, usually sparsely
covered with papilliform hairs on both sides, margin ciliate, less often
more or less glabrous; basal and lower cauline leaves with up to 12 cm
long petioles, deltoid, oblong-hastate, or almost triangular, attenuate-
acuminate, hastate-cordate or truncate at base, sharp toothed, teeth finely
pointed, lamina 7—13 cm long; upper cauline leaves oblong-lanceolate
or lanceolate, short-petiolate or almost sessile, narrow cuneate at base;
uppermost leaves lanceolate-linear, almost entire. Capitula campanulate,
1.5—2.5 cm long including florets and 1.5(2) cm wide, numerous, solitary
on long peduncles, forming elongate racemes with less numerous capitula.
Involucre five-rowed, about 1.5 cm long, involucral bracts imbricate,
appressed; outer bracts ovate, villous-woolly or almost glabrous, attenuate
into narrow, long (0.5—1 cm long), dark colored, recurved, saber-shaped
appendage; inner bracts elongate, oblong, somewhat obtuse, usually
without appendage, flocculose-woolly at apex. Receptacle densely
squamose, scales linear-subulate, white, lustrous, numerous, I—1.2 cm
long. Florets violet, 1.3-1.5 cm long, narrow part of tube 0.7—0.8 cm
long, broad part 2—2.5 mm long, lobes of limb about 0.4 cm long;

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460

anthers prominent, about 0.5 cm long, anther appendages ciliate. Pappus
double, outer pappus bristles few, unequal, up to 3-4 mm long, serrate,
inner bristles long-plumose, rusty near base, about 1 cm long; achenes
about 5 mm long and about 1 mm across, narrow-cylindrical, slightly
narrowed towards base, dark mottled glabrous. Flowering VII-IX.

Shady forests, rock crevices in forests—Far East: Ussuri. General
distribution: Korea. Described from a few herbarium sheets from the
Ussuri Region. Authentic specimens in Leningrad. I designate the
specimens of Maximovicz from the region of Bruse (now Slavyanka)
Inlet as the lectotype.

Note. V.L. Komarov was inclined to consider this species closer to
S. triangulata than to S. grandifolia. But S. triangulata differs sharply
in its capitula from S. subtriangulata, although the leaves of the two
species are, in fact, similar. S. subtriangulata is extremely close to S.
grandifolia, from which it differs by the shape (narrower and elongate-
hastate) and consistency of the leaves, smaller number of capitula and,
most importantly, by the outer involucral bracts, which are furnished
with long, recurved, saber-shaped appendages. The plant that is annotated
S. subhastata Kom. (1933) in the Herbarium of the Botanical Institute,
Academy of Sciences of the USSR, does not differ at all from S.
subtriangulata. Hybridization between S. subtriangulata and S.
grandifolia is a likely possibility.

66. S. sinuata Kom. Fl. Manchzh. III (1907) 735; Kom. and Allis.
Opred. Rast. Dalnevost. Kr. II, 1071; Lipschitz in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XVII, 441.—S. mongolica auct.: Hand.-Mazz.
in Acta hort. Gotoburg. XII (1938) 320, p. p. non Franch.—Ic.: Kom.
op. cit. (1907) Plate XIV.

Perennial. Rhizome short; stem 40-90 cm high, solitary, erect,
simple, branched only above, angular-sulcate, smooth, wingless. Leaves
thin, dark green above, pale green beneath, slightly lustrous, basal leaves
usually withering before anthesis, lower and middle cauline leaves with
up to 13 cm long petioles, with short ciliate hairs on both sides,
particularly dense along margin; lamina up to 12 cm long, 10 cm wide,
oval-oblong, oval or even somewhat attenuately triangular, attenuate-
cuspidate, almost cordate or truncate at base, with two to four deep
roundish notches on each side along margin, lobes acute, two- or three-
toothed, cuspidate; uppermost leaves lanceolate or linear-lanceolate,
narrower, almost entire, cuneately narrowed toward base into short petiole
or almost sessile; bracteal leaves subulate, sessile. Capitula one to seven
(nine), solitary terminal on lateral, long (up to 7 cm long), pubescent
peduncles with stiff hairs, forming racemose inflorescence. Involucre
five-rowed, turbinate, 1.2—-1.8 cm long; outer involucral bracts ovate or

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467

Plate XXXV.

1—Saussurea sukaczevii Lipsch.; 2—S. poljakovii Glehn.

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462

oblong, with green, erect or recurved, saber-shaped, 5—15 mm long
appendages; innermost bracts linear, acuminate, with or without
appendages; all bracts puberulent or weakly arachnoid-hairy. Receptacle
densely squamose, scales linear-subulate, white, lustrous, about 7 mm
long. Florets pale pink, about 1.3(1.4) cm long; narrow part of tube
about 0.6 cm long, almost as long as broad part together with lobes of
limb; anthers prominent, 0.5—0.6 cm long, anther appendages more or
less pubescent. Pappus double, outer pappus bristles less numerous,
unequal, up to 3 mm long, serrate, fragile, inner bristles long-plumose,
about 1 cm long, straw yellow in lower part, dirty white above; mature
achenes about 7 mm long, narrow-cylindrical, slightly narrowed toward
base, dark brown, glabrous, immature achenes dark mottley. Flowering
Vil.

Broad-leaved, coniferous, and cedar-broad-leaved forests —Far East:
Ussuri. General distribution: Northeastern China. Described from
northeastern China; type in Leningrad.

Note. Close to S. grandifolia Maxim. and S. subtriangulata Kom. It
can be distinguished at once from S. grandifolia by its deeply divided
leaves (sometimes almost pinnatilobate) and long, saber-shaped, usually
recurved appendages of the involucral bracts, as a result of which the
imbricate arrangement becomes indistinct. From S. subtriangulata it
differs by the presence of deep notches in the leaves, the turbinate shape
of the capitula, and the absence of flocculose-woolly pubescence
(glochidia) at the tip of inner involucral bracts.

Series 13. Triangulatae Lipsch—Ser. Imbricatae Kitam. in Acta
Phyt. et Geobot. IV (1935) 11 p. p.; Ej. in Mem. Coll. Sc. Kyoto Univ.
ser. B, XIII (1937) 198 p. p.—Capitula on long peduncles, aggregated
in independent corymbs, forming lax panicle, sometimes on short
peduncles in appressed compound corymbs. Involucral bracts dark colored
or green; outer bracts reduced, ovate, obtuse or subobtuse. Leaves
elongate-triangular, not glandular. Stem wingless.

Members of the series resemble species of series Fauriei Kitam. in
the nature of the capitula and inflorescence.

67. S. triangulata Trautv. and Mey. in Fl. Ochot. (1856) 58; Maxim.
Prim. Fl. Amur. 167; Fr. Schm. in Mém. Acad. Sc. Pétersb. sér. VII,
XIII, 2, 152; Herd. in Bull. Soc. Nat. Mosc. XLI, 3, 33 (incl. a. genuina
Herd. (p. 33) and B. elatior Herd. (p. 34)); Kom. Fl. Manchzh. III, 737;
Nakai in Bot. Mag. Tokyo, XXIX, 203; Ej. Fl. sylv. Korean. XIV, 120;
Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1072; Kitam. in Acta
Phyt. et Geobot. IV, 11; Ej. in Mem. Coll. Sc. Kyoto Univ. ser. B,
XIII, 199 (excl. B. alpina Nakai; subsp. manshurica (Kom.) Kitam.,;

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463

var. pinnatifida (Nakai) Kitam.); Sugaw. Ill. Fl. Saghal. IV, 1873;
Lipschitz in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI, 452.—
S. derbeckii Kom. in Izv. Peterb. Bot. Sada, X, 4 (1910) 119; Kom. and
Alis. op. cit. 1072.—S. schischkinii Kom. in Bot. Mat. Gerb. Glavn.
Bot. Sada SSSR, VI, 1 (1926) 17; Kom. and Alis. op. cit. (1932) 1071.—
Ic.: Fl. Ochot. (1856) Plate 29.

Perennial. Rhizome prostrate; stem erect, 30—70(100) cm high,
angular-sulcate, glabrous or sometimes pubescent with curly hairs in
upper part, often simple, less often branched above, branches straight,
appressed or somewhat spreading. Leaves distant, thin, more pale beneath
with prominent midrib and less distinct lateral veins, toothed or shallow
crenate-toothed, teeth sharp; lamina glabrous or slightly crisped-pubescent
on both sides, sparsely ciliate; basal leaves usually withering before
anthesis, less often persistent; lower and middle cauline leaves petiolate,
petioles up to 10 cm long, gradually reduced upward, sometimes winged,
broadened at base into narrow and short sheath; lamina usually oblong-
triangular, triangular-hastate, sometimes oblong or ovate, 4-12 cm long,
acuminate, less often attenuate-acuminate, cordate at base, truncate or
cuneate more frequently in middle cauline leaves; upper cauline leaves
reduced upward, sessile on very short-petiolate, oblong, ovate or
lanceolate, acuminate, truncate-cuneate at base or weakly cuneate, less
often rounded; more or less toothed or almost entire. Capitula about 1.2
cm long and wide, almost sessile in typical specimen, terminal on stem
in globule of compressed corymbs, sometimes terminal on more or less
long, angular, slender peduncles aggregated with central corymb in
paniculate or racemose lax inflorescence, capitula 3—20. Involucre tubular,
roundish at base, 1.2(1.3) cm long and about 0.7 cm wide, four- or
five-rowed, involucral bracts imbricate, usually entirely dark-colored,
less often green, colored only at apex, without appendage, almost
glabrous, sometimes with less numerous hairs or slightly arachnoid-hairy
on dorsal surface and near tip, as well as along margin; outer involucral
bracts half as long as inner, ovate, subobtuse or obtuse like middle
bracts; inner bracts linear, subobtuse. Receptacle squamous, scales linear-
subulate, about 7 mm long, lustrous; florets purple, about 1(1.2) cm
long, narrow part of tube about 0.5(0.6) mm long, as long as broad part
together with lobes of limb; lobes of limb almost as long as broad part
of tube. Pappus smoky, double, outer pappus bristles less numerous,
short, up to 3 mm long, serrate or finely plumose, inner bristles long-
plumose, about 9 mm long; achenes narrowly cylindrical, about 6 mm
long and 1 mm wide, mature achenes dark brown, glabrous. Flowering
VU-VIIl.

Deciduous and coniferous forests, groves of Chosenia, meadows,
grassy patches near sea coast.—Far East: Zeya-Bureya, Ussuri, Uda, -

47)

464

Okhotsk regions, Sakhalin. General distribution: Korea. S. triangulata
is described from the Shantar Islands; type in Leningrad. S. derbeckii
Kom. from the De-Castri Bay; type in Leningrad. S. schischkinii Kom.
from Sikhote-Alin; type in Leningrad.

Note. Herder (op. cit., 1868, p. 34) rightly reported that S. triangulata
is extremely variable: 1) in the number, position and size of capitula
(corymb sometimes compressed, sometimes more lax, and then with a
series of lateral corymbs on more or less long peduncles, forming
paniculate or racemose inflorescence; 2) in the color of the involucral
bracts (green, dark-colored); and 3) in the size of the plant and its
leaves. The shape of the leaf base and the number of rows of involucral
bracts should also be added to this. V.L. Komarov described S. derbeckii
in 1910 and S. schischkinii in 1926 but did not compare them with
S. triangulata with which they should have been compared, being
phylogenetically closest to it. S. derbeckii is compared with
S. acuminata Turcz. and S. sachalinensis Fr. Schm., S. schischkinii 1s
compared with S. elongata DC. Among the typical characters of
S. derbeckii are the cuneate base of the lower leaves, violet-black
involucral bracts, thin leaves, etc. Comparison of S. derbeckii with
S. triangulata shows them to be identical. As regards S. schischkinii, its
author considers the dense inflorescence comprising only a few
capitula to be the main character; the same picture is observed in the
authentic specimen of S. triangulata. Cf. also the note to S. manshurica
Kom.

68. S. manshurica Kom. in Tr. Peterb. Bot. Sada, XVIII, 3 (1900)
424; Kom. Fl. Manchzh. III, 730; Kom. and Alis. Opred. Rast. Dalnevost.
Kr. II, 1072; Ling in Contr. Inst. Bot. Nat. Acad. Peip. III, 172; Sugaw.
Ill. Fl. Saghal. IV, 1873.—S. triangulata subsp. manshurica (Kom.)

‘Kitam. in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII (1937) 201, excl.

var. pinnatifida (Nakai) Kitam.—S. triangulata auct. fl. Sachal. et
Orientalis Extremi p. p.—Ic.: Kom. Fl. Manchzh. III, Plate XII; Sugaw.
op. cit. (1940) tab. 860.

Perennial. Rhizome prostrate; stem (25)50—90 cm high, erect, slender,
with narrow furrows, sometimes thick, fistular below, glabrous, simple,
branched only in inflorescence. Leaves thin, green, more pale beneath,
triangular-hastate, less often ovate or oblong, acuminate, less often
attenuate-acuminate, toothed or shallow crenate-toothed, teeth sharp;
lamina glabrous on both sides, sometimes sparsely hairy, ciliate, cordate
or cuneate, less often truncate at base, with prominent midrib and less
distinct lateral veins; basal leaves usually withering before anthesis, long-
petiolate like lower cauline leaves; middle leaves short-petiolate,
uppermost sessile, reduced, lanceolate or oblong, cuspidate, entire.

472

465

Inflorescence in typical specimen many-headed; however, often
compound; capitula solitary on long bent or erect peduncles on lateral
floriferous branches, forming dense corymbs, enclosing two or three
capitula; lateral peduncles often forming lax panicle, crowded at stem
apex or in its upper one-third part. Capitula narrow, 5 mm wide in
lower part, up to 12 mm in upper part. Involucre five- to seven-rowed,
imbricate; involucral bracts glabrous, shining, weakly pubescent only
along margin and apex; outer bracts ovate, obtuse, innermost bracts
oblong, much longer than in first outer row, obtuse; all bracts usually
entirely, or along margin dark-colored or entirely green. Receptacle
squamous, scales linear-subulate, lustrous, about 7 mm long. Florets
dark violet, about 1—1.1 cm long, narrow part of tube about 5(6) mm
long, as long as broad part together with lobes of limb; lobes of limb
3-4 mm long, slightly longer than broad part of tube; anther appendages
hairy. Pappus double, smoky, outer pappus bristles less numerous, serrate,
fragile, up to 4 mm long, inner pappus bristles long-plumose, about
0.9-1 cm long; achenes 4-5 mm long, glabrous, dark brown. Flowering
VII-VIILI.

Crevices of rocks, stony slopes, outcrops of crystalline shales in
mixed or coniferous forests, Erman’s birch forests—Far East: Zeya-
Bureya, Ussuri, Uda, Sakhalin. General distribution: Northeastern China,
Korea. Described from the Burei Mountain. Type in Leningrad.

Note. V.L. Komarov distinguished his species from S. triangulata
Trautv. and Mey. by the following characters: narrow capitula on long
peduncles, a five-rowed involucre and lax inflorescence. The extreme
forms of S. manshurica, originally designated by the author in the
description as S. manshuriensis, distinctly differ from S. triangulata by
their general appearance. The inflorescence of S. manshurica is lax,
many-headed, and with numerous floriferous branches, whereas the
inflorescence of S. triangulata is compressed, and few-headed. The
character cited for distinguishing the two species, namely, differing
numbers of rows of involucral bracts, does not hold up. Some forms of
both species are difficult to separate from each other, and constitute a
transition. In accordance with the tradition of the Flora of the USSR, |
accept the binomial name for the species, although Japanese botanists
have reduced S. manshurica to the status of a subspecies of S. triangulata.

Series 14. Fauriei Kitam. in Acta Phyt. et Geobot. IV (1935) 5; Ej.
in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII (1937) 179.—Stem more or
less winged. Capitula sessile or on very short peduncles, in dense
corymbs. Involucre narrow-tubular; involucral bracts imbricate, outer
bracts ovate or deltoid, together with middle bracts obtuse or rounded
at apex. Usually tall meadow and forest herbs.

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466

69. S. parviflora (Poir.) DC. in Ann. Mus. Paris, XVI (1810) 200;
Ej. Prodr. VI, 534; Turcz. Fl. baic.-dahur. II, 1, 110; Kryl. Fl. Zap. Sib.
XI, 2916; Lipschitz in Sp. Rast.-Gerb. Flory SSSR, XIII, 126— -Serratula
parviflora Poir. Encycl. Meth. VI (1805) 554.—Saussurea serrata DC.
in Ann. Mus. Paris, XVI (1810) 199; Ldb. Ic. pl. Fl. Ross. 1, 17, 18;
Ldb. FI. alt. IV, 25; DC. Prodr. VI, 534; Ldb. Fl. Ross. II, 2, 666;
Herd. in Bull. Soc. Nat. Mosc. XLI, 3, 18, excl. C. amurensis Herd.
and B. corymbo contracto; Kryl. Fl. Alt. III, 705; Popov, Fl. Sr. Sib. II,
759.—S. elata auct.: Turcz. in herb. (non Ldb.).—S. atriplicifolia Fisch.
ex Herd. op. cit. (1868) 19 and herb—Heterotrichum atriplicifolium
MB. ex Herd. op. cit. (1868) 19.—S. purpurata Fisch. ex Herd. op. cit.
(1868) 20.—Heterotrichum glabellum MB. ex C.A.M. in DC. Prodr. VI
(1837) 534; Herd. op. cit. 20.—Serratula multiflora Pall. ex Herd. op.
cit. (1868) 20.—Ie.: Ldb. Ic. Fl. Ross. I, tab. 71 and 72—Exs.: Smirnow
Pl. alt. No. 96 (1930) (sub nom. S. serrata DC.); GRF No. 3992.

Perennial. Rhizome prostrate; stem solitary, 20-100 cm high, erect,
simple or branched above, fistular, strongly leafy in typical specimen,
sulcate, glabrous or weakly hispid, usually more or less winged due to
decurrent leaves, less often wingless, wings more or less narrow, with
entire margin. Leaves green above, glaucous beneath, glabrous on both
sides or with stiff ciliate hairs on lower surface, especially along veins
and margin; midrib prominent, numerous lateral veins poorly noticeable;
basal leaves usually withering before anthesis, like lower cauline leaves
petiolate, elliptical or oblong-elliptical acuminate, usually serrate-toothed,
teeth small and infrequent, terminating in callose cusp, less often lamina
entire, unequal in length and width, 8-30 cm long together with petiole
and 1.5—2.5(4) cm wide; middle leaves lanceolate or elliptical-lanceolate,
attenuate-acuminate, sessile, usually strongly decurrent, cuneate at base,
not cordate; upper leaves reduced, lanceolate or linear-lanceolate, entire,
sessile, usually not decurrent. Capitula terminal on main stem and
floriferous branches in corymbs; corymbs sometimes broad, numerous,
and many-headed, forming paniculate inflorescence (var. patens Ldb.
Fl. Ross. II, 2 (1845-1846) 667), sometimes solitary, terminal dense
(var. contracta Ldb. op. cit., 667); peduncles of capitula and corymbs
pubescent, scabrous or almost glabrous, smooth. Involucre narrow-tubular,
3—3.5 mm wide, three- or four-rowed, imbricate; involucral bracts usually
entirely or in upper half dark-colored, glabrous, sometimes hairy along
margin or less often entirely floccose-tomentose at apex, especially in
inner bracts, outer bracts smaller, orbicular-ovate, obtuse or subobtuse;
inner bracts oblong, obtuse or roundish. Receptacle densly squamose,
scales linear-subulate, white, lustrous, about 6-7 mm long. Florets violet
lilac (in ssp. purpurata (Fisch.) Lipsch.), about 1—1.2 cm long, narrow
part of tube about 5—6 mm long, almost as long as broad part together

467

with lobes of limb; lobes of limb almost as long as broad part of tube;
anther appendages hairy. Pappus double; outer pappus bristles unequal,
2-4. mm long, serrate or finely plumose, inner bristles 8-9 mm long,
long-plumose; mature achenes 3—3.5 mm long, whitish or yellowish with
dark spots. Flowering VII—VIII. (Plate XXXIV, Fig. 2.)

Wet meadows, marshes, river banks, thickets of shrubs, deciduous
and mixed forests, coniferous forests, alpine meadows, tundras, stony
slopes, rocks.—European Part: Dvina-Pechora (extreme western location:
the former Ust-Sysolsk District), Volga-Kama (Ural part), Trans-Volga
Region (Ural part); Western Siberia: Upper Tobol, Ob’, Irtysh (eastern
part), Altai; Eastern Siberia: Yenisei, Angara-Sayans, Dauria, Lena-
Kolyma. General distribution: Mongolia, China (northern)? Described
from Siberia; type specimen in Geneva(?).

Note. An extremely polymorphic species; varies in the following
characters: 1) size of leaves; 2) pubescence of their lower surface;
3) cutting of margin or leaves almost entire; margin of leaf sometimes
almost flat, sometimes slightly involute; 4) leaf base; 5) shape of
inflorescence; 6) color and degree of pubescence of involucral bracts;
7) pubescence of peduncles of capitula; and 8) stem height (specimens
from alpine tundra are usually much smaller than from lower regions).
M.G. Popov (op. cit., 1959) reported hybrids of this species with S.
latifolia Ldb.

Ssp. purpurata (Fisch. ex Herd.) Lipsch. in Sp. Rast. Gerb. Flory
SSSR, XIII (1955) 127; in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XXI (1961) 375.—S. purpurata Fisch. ex Herd. in Bull. Soc. Nat. Mosc.
XLI (1868) 20.—S. serrata a. purpurata Herd. op. cit. (1868) 20.—

474 Heterotrichum glabellum MB. ex C.A.M. in DC. Prodr. VI (1837) 534;
Herd. op. cit. (1868) 20.—Serratula multiflora Pall. ex Herd. op. cit.
(1868) 30.—Saussurea acuminata Turcz. y. gracilis Herd. op. cit..(1868)
23.—Entire plant, especially stem, usually purple, somewhat woody,
glabrous, root collar sparsely covered with fibrously split sheaths; upper
cauline leaves usually narrower, lanceolate or linear-lanceolate, almost
entire, glabrous beneath, slightly involute, with weakly decurrent bases,
often not at all decurrent on stem, and then wingless in typical specimens.
Corymbs compact, less numerous, comprising capitula on short peduncles.
Involucral bracts usually longer than in type specimen, usually dark-
colored, almost glabrous, less often pubescent; florets purple. Described
from “Dauria” (Vlasov). Type in Leningrad. The subspecies is distributed
in the Trans-Baikal Region (but also found in Tomsk, Angara-Sayan);
with many forms intermediate between it and S. parviflora.

A unique variety of S. parviflora, distinguished by deeply sinuate
leaves, producing, however, a whole gamut of intermediates to simply
repand leaves was collected by V.N. Sukaczev and his collectors on

475

468

the eastern bank of Baikal, between the Ust-Barguzin Station and the
Maksimikha Station; I have described it as var. incisa Lipsch. (Bot.
Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XXI (1961) 375).

Quite a few specimens were collected from the Chuno-Angara
watershed, which, in my opinion, are hybrids between S. parviflora (Poit.)
DC. and S. stubendorffii Herd. They strongly resemble S. parviflora in
having prominently winged stems and in their leaf shape, but the lower
surface of the leaves has arachnoid-hairy pubescence and the capitula
are larger; the involucral bracts are strongly pubescent, and the florets
are bright red. These specimens are closer to S. stubendorffii in the
latter characters.

Hybrids between S. parviflora (Poir). DC. and S. controversa DC.
are known from the Balagan Region. Cf. the description in the note to
S. controversa DC.

70. S. porcii Degen in Mag. Bot. Lapok 12 (1904) 311; Wilczynski
in Mag. Bot. Lapok, XXVI, 65.—S. serrata auct.: Janka in Oest. bot.
Zeitschr. VIII, 6 (1858) 200; and auct. fl. Transsylv.; Herd in Bull. Soc.
Nat. Mosc. XLI, 3, 22 in nota, non DC.—S. parviflora auct.: Janka in
Linnaea, XXX (1859) 585; non Poir. nec DC.—S. alata auct.: Porcius
and Czetz. ap. Porc. Fl. phanerog. din. fost. distr. alu Naseud (1881)
118, non DC.

Perennial. Rhizome dark brown, woody, almost horizontal, strongly
filamentous; stems 30-80 cm high, almost glabrous or rather sparsely
arachnoid-hairy when young, smooth, densely leafy. Leaves fleshy,
narrow-lanceolate, acute, with broad base, wing-like decurrent on stem,
entire; middle cauline leaves slightly broader, lanceolate, acute, entire,
involute and sparsely toothed; upper cauline leaves shorter and narrower,
uniformly lanceolate or almost linear, acute, entire; all leaves broadly
wing-like decurrent along stem, glabrous above, young leaves rather
sparsely flocculose-arachnoid hairy beneath, however, glabrescent, lower
surface of adult leaves with few crisped hairs, slightly scabrous or almost
glabrous, pubescent or glabrous along margin. Inflorescence terminal of
more or less globose corymbs; inflorescence branches somewhat flat,
winged, slightly longer than corymbs; capitula obconical. Involucral
bracts imbricate, three- to five-rowed; outer bracts ovate-lanceolate,
obtuse; middle bracts ovate, acuminate; inner slightly oblong, lanceolate-
ovate, obtuse; all bracts green at base, dark red above middle, dorsal
surface covered with less numerous but long villous hairs, or sometimes
entirely arachnoid-hairy, woolly along margin, glabrous inside. Florets
lilac colored, five-fid above middle; anthers two times as long as
filaments, with hairy appendages at base. Receptacle squamous; pappus
double, outer pappus bristles subulate, short-plumose, inner pappus

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bristles long-plumose; young achenes dark brown, slightly compressed,
glabrous, deeply sulcate.

Marshy meadows at about 1,500 m.—European Part: Upper Dniestr.
General distribution: Central Europe. Described from specimens of
Porcius. Collected in eastern Transylvania. Type evidently in Budapest.

Note. This species is not represented in the Herbarium of the
Botanical Institute, Academy of Sciences of the USSR. Its description
is taken from Degen and Wilczynski; the latter collected S. porcii in
the Carpathians (in the eastern part of the Lyudovan Mountains).

These authors distinguish S. porcii from S. parviflora by the
following suite of characters: lanceolate (and not elliptical-lanceolate),
entire, acute leaves; larger capitula; obconical involucre (not narrow-
tubular, as.in S. parviflora); much more densely leafy stems; broadly
(not narrowly) winged broader, less acute involucral bracts; long villous
dorsally (and not glabrous or arachnoid-ciliate only along margin), less
deeply divided corolla (divided two-thirds of the length in S. parviflora);
anthers twice as long as filaments; and longer pappus. S. porcii is a
species which needs to be compared with the large amount of material
accumulated for S. parviflora.

71. S. neoserrata Nakai in Bot. Mag. Tokyo, XLV (1931) 519;
Kitam. in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII, 179.—S. serrata c.
amurensis Herd. in Bull. Soc. Nat. Mosc. XLI, 3 (1868) 19; Kom. FI.
Manchzh. III, 735.—S. serrata in Kom. and Alis. Opred. Rast. Dalnevost.
Kr. II (1932) 1064 and auct. plur. Fl. Extr.-Orient—Exs.: Karo Plantae
Amuricae et Zeaensae curav. Dérfler. No. 441 (sub nom. S. serrata
DC.).

Perennial. Rhizome prostrate; stem erect, 50-100 cm high, winged-
angular, crisped hairy, more or less glabrous above, fistular, densely
leafy, branched above with ascending branches, less often simple. Leave
coriaceous, green above, glabrous or sparsely ciliate-hairy, glaucous or
light glaucous beneath, glabrous or less often with bent, scattered,
somewhat stiff hairs and then scabrous, with prominent midrib beneath
and less distinct lateral veins; basal and lower cauline leaves withering
before anthesis; cauline leaves broadly lanceolate, elliptical-lanceolate,
12-20 cm long and up to 8 cm wide, attenuate-acute, distinctly and
sharply toothed-serrate, cuneately narrowed at base into a broad, up to
1 cm wide, petiole, wing-like decurrent; upper cauline leaves gradually
reduced, narrow-lanceolate or linear-lanceolate. Capitula about 1—1.2 cm
long and about 0.4-0.5 cm wide, rather numerous, on short peduncles,
in dense corymbs, terminal on main stem and lateral branches, forming
paniculate inflorescence. Involucre tubular, with turbinate base, imbricate,
four- or five-rowed; involucral bracts usually green, often dark red along

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margin (less often entirely colored); all bracts obtuse, dorsally smooth,
sparsely arachnoid-hairy along margin; outer bracts ovate, reduced, one-
fourth as long as inner; middle bracts oblong, inner oblong-linear. Florets
purple or pale purple, about 0.9 cm long, narrow part of tube about 5
mm long, broad part with lobes of limb about 4 mm long; anther
appendages hairy. Receptacle squamose, scales narrow-linear, lustrous,
about 6-7 mm long. Pappus double, outer pappus bristles reduced,
unequal, about 1—2 mm long, fragile, serrate or finely plumose, inner
bristles about 6-7 mm long, long-plumose; achenes 5 mm long, glabrous,
sulcate, truncate at apex, gradually tapered at base throughout, with
dark spots. Flowering VII—VIII. (Plate XXXIV, Fig. 1.)

Moist meadows, hummocky marshes, among shrubs, edges of mixed
forests, broad-leaved and coniferous forests, sedge-willow-dwarf Arctic
birch forest, thickets of filbert—Eastern Siberia: Dauria; Far East: Zeya-
Bureya, Ussuri, ?Uda Region. General distribution: Korea. Described
from Korea. Type probably in Tokyo.

Note. It hybridizes with S. umbrosa Kom. | include specimens from
the northern Sikhote-Alin (Botchi River Valley, 1924) among these
hybrids, in which the involucral bracts, especially the outer ones, have
small, leafy, saber-shaped, recurved appendages.

S. neoserrata appears in Trans-Baikal, where it begins to replace
the fairly polymorphic species S. parviflora; as it moves eastward,
it gives rise to a series of transitional forms, possibly through
hybridization.

72. S. fauriei Franch. in Bull. Herb. Boiss. V, 7 (1897) 542;
Matsumura, Ind. pl. Jap. II, 661; Kitam. in Acta Phyt. et Geobot. IV, 5;
Ej. in Mem. Coll. Sc. Kyoto Univ. ser. B, XIII, 180; Ej. in Iconogr. pl.
As. Orient. V, 1 (1950) 463.—TIc.: Kitam. op. cit. (1950) tab. CXLIII.

Perennial. Rhizome oblique, woody, developing numerous fibrous
roots; stem strong, straight, 150-200 cm high, up to 2 cm thick at base,
sulcate, winged, densely leafy, puberulent with crisped hairs, branched
above. Leaves hard; basal leaves withering before anthesis; cauline leaves
broad, ovate, ovate-oblong or elliptical-lanceolate, up to 25-30 cm long
and 5—8 cm wide, cordate, truncate, or less often cuneate at base, with
short weakly winged petiole, decurrent on stem, more or less long
attenuate, irregularly serrate-toothed, teeth poorly developed, callose at
apex, cuspidate; lamina green above, grayish pubescent beneath and
under lens with usually numerous, lustrous, scattered glandular hairs;
uppermost cauline leaves reduced, linear-lanceolate, approximate with
inflorescence, sometimes entire. Inflorescence many-branched, of
numerous dense corymbs, forming more or less broad panicle; capitula
numerous, on short peduncles, narrow-cylindrical, compactly arranged

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in individual corymbs. Involucre imbricate, five-rowed, green, 10-12
mm long and 4-5 mm wide; outer involucral bracts reduced, orbicular;
middle bracts ovate, inner linear; all bracts obtuse, slightly brownish,
and arachnoid-hairy at apex, as well as along margin. Receptacle
squamose, scales linear-subulate, lustrous, unequal. Florets 10 mm long,
narrow part of tube 5 mm long, as long as broad part with lobes of
limb; lobes of limb three times as long as broad part of tube; anther
appendages ciliate. Pappus double, outer pappus bristles numerous,
reduced, unequal, up to 3 mm long, fragile, serrate-scabrous; inner pappus
bristles long-plumose, 8—8.5 mm long; achenes immature, about 3 mm
long, with small ribs. Flowering VII—VIII.

Coastal tall herb meadows.—Far East: Sakhalin (southern Kurils,
Kunashir Island, Shikotan). General distribution: Japan. Described from
Ezo Island. Type apparently in the Herbarium of Franchet in Paris.

Note. Franchet compared this species with S. parviflora (Poir.) DC.
In my view, it is closest to S. neoserrata Nakai, from which it differs
by taller growth, leaves that are grayish pubescent. underneath (and not
glaucous and glabrescent, glabrous), less pronounced marginal teeth, as
well as more rounded bases of the blade and shorter winged petioles.
The relationship of S. fauriei Franch. and S. neoserrata Nakai requires
further study in nature.

Series 15. Latifoliae Lipsch—Stem more or less winged, densely
leafy. Leaves ovate or elliptical, green on both sides, obtuse or weakly
attenuate at apex, slightly acuminate. Capitula in dense terminal corymbs,
less often also on lateral branches; upper cauline leaves sometimes
approximate with inflorescence. Involucre imbricate; involucral bracts
in typical specimen lustrous, obtuse or subobtuse. More or less tall forest
and subalpine herbs.

Note. 1 The members of series Latifoliae are morphologically close
to the members of series Faurieae, but differ from the latter by the
larger size and shape of the capitula and less attenuate-acuminate middle
cauline leaves. Latifoliae is readily distinguished from the closely related
series Foliosae Lipsch. by the large size of the leaves, blades that are
similarly colored on both sides, and the different shape and size of the
outer involucral bracts.

Note 2. S. americana Eaton, growing in North America, is closest
to series Latifoliae. I find great phytogeographical significance in the
closeness of S. latifolia Ldb. with .S. americana Eaton, indicating a
common origin of the herb cover of the Siberian and North American
taiga. Evidently, S. americana Eaton should be placed in a different
series, Americanae Lipsch., parallel to series Latifoliae Lipsch.; similarly
the Siberian series Foliosae, corresponds to a series Pseudofoliosae

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Lipsch. in America, with the only species being S. pseudofoliosa Lipsch.
sp. nova. Its description will be given in another place.

73. S. latifolia Ldb. Ic. pl. Fl. Ross. I (1829) 17; Ej. Fl. alt. IV, 24;
DC. Prodr. VI, 534; Ldb. Fl. Ross. II, 2, 666; Herd. in Bull. Soc. Nat.
Mosc. XLI, 3, 17; Kryl. Fl. Alt. III, 704; Kryl. Fl. Zap. Sib. XI, 2914;
Popov, Fl. Sr. Sib. II, 758—S. macrophylla Bge. ex Herd. op. cit. (1868)
17.—Ie.: Ldb. Ic. pl. Fl. Ross. 1 (1829) tab. 70.—Exs.: Smirnov. PI.
alt. No. 95.

Perennial. Plants 30-120 cm high, glabrous or weakly pubescent,
with prostrate rhizome, producing one or less often two or three stems;
stem erect, simple, sulcate, winged or less often wingless (f. exalata
Kryl.), strongly leafy; leaves ovate or elliptical, acuminate, but only
slightly attenuate at apex, scabrous above, more or less pubescent
beneath, occasionally quite strongly (f. pubescens Lipsch.) or less often
glabrous, serrate-toothed and puberulent along margin, teeth terminating
in short cusp; basal and lower cauline leaves petiolate, somewhat cordate
at base, 6-15 cm long and 4—9 cm wide; upper leaves smaller, sessile
or with shorter petioles, decurrent along internodes and then latter
winged; uppermost leaves narrow-lanceolate or linear-lanceolate; width
of wings variable, less often weakly toothed, sometimes wings absent
(f. exalata Kryl.). Capitula campanulate, about 1.5—-1.8 cm long and
0.5—0.6(0.8) cm wide, slightly enlarged in fruits, in terminal, usually
dense corymbs or, with lateral corymbs, forming corymbose panicle.
Involucre four- or five-rowed; involucral bracts imbricate, light-colored
or often dark violet; outer bracts ovate, obtuse or subacute; inner bracts
oblong or lanceolate, subobtuse or acuminate; all involucral bracts more
or less pubescent, especially along margin and at apex, glabrescent and,
therefore, involucral bracts in typical specimen almost glabrous and
shining. Receptacle with spare lustrous scales; florets purple-violet, about
10(11) mm long, narrow part of tube about 5(6) mm long, almost as
long as broad part including lobes of limb; anthers violet, anther
appendages hairy. Pappus double; outer pappus bristles unequal, up to
3—4 mm long, fragile, short-plumose or serrate-scabrous, inner bristles
about 9-10 mm long, long-plumose; achenes smooth, half to two-thirds
as long as pappus. Flowering VII—VIII.

Coniferous and mixed forests; forest, subalpine and aitiad meadows,
less often moss-lichen tundras in the lower part of alpine belt—Western
Siberia: Irtysh (eastern part), Altai; Eastern Siberia: Angara-Sayans,
Dauria (southwestern part). Soviet Central Asia: Dzhungaria-Tarbagatai.
General distribution: Mongolia (Mongolian Altai). Reported by P.N.
Krylov (Flora of Western Siberia, XI, 2915) for Tarbagatai; I did not
see the specimens. Described from Altai; type in Leningrad.

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473

Note. As was mentioned above, the American species S. americana
Eaton is closest to S. latifolia. The former differs slightly from the
Siberian species by somewhat narrower leaves with more pronounced
cuneate bases, less decurrent on the stem, and thus the stem is almost
wingless, and by stronger pubescence on the involucral bracts.

Series 16. Foliosae Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 376.—Stem taller, sometimes (in alpine forms)
shorter, or plants almost without stem, more or less densely leafy up to
tip, sometimes, especially in S. foliosa Ldb., hidden under leaves; leaf
bases nondecurrent or weakly decurrent on stem. Leaves white-tomentose
beneath, less often glabrous on both sides, green, toothed in type
specimen. Capitula sometimes in terminal capitate corymbs (then corymbs
usually not observed on lateral branches), subtended by upper cauline
leaves, sometimes in more lax corymbs, then capitula on distinctly
compressed, glandular-pubescent or almost glabrous peduncles. Capitula
campanulate or tubular. Involucre indistinctly or distinctly imbricate;
outermost bracts lanceolate, usually as long as inner bracts in type
specimen or sometimes shorter; inner bracts more or less densely covered
with long, somewhat stiff or soft appressed hairs; outer and middle series
of bracts long acuminate. Transitions present from distinctly to
indistinctly imbricate involucre. Receptacle squamous.

74. S. foliosa Ldb. Ic. pl. Fl. Ross. I (1829) 17; Ej. Fl. alt. IV, 23;
DC. Prodr. VI, 535 (excl. var. B.); Ldb. Fl. Ross. II, 2, 669; Turez. FI.
baic-dahur. II, 1, 116 (excl. var. B. dentosa Turcz. and y. pygmaea
Turcz.); Kryl. Fl. Alt. II, 705; Kryl. Fl. Zap. Sib. XI, 2915.—S. alpina
foliosa Herd. in Bull. Soc. Nat. Mosc. XLI, 3 (1868) 38.—Ic.: Ldb. op.
cit. (1829), tab. 69.

Perennial. Rhizome prostrate, developing several or solitary stems;
stems erect, unbranched, 10-35 cm high, slightly pubescent or usually
glabrous, sulcate, densely leafy up to inflorescence, usually hidden under
leaves. Leaves hard, oblong-ovate or long-elliptical, subacute, involute,
toothed with cuspidate teeth; white-tomentose beneath, green and glabrous
above (var. altaica Turcz. op. cit. 1856, 116 = var. tomentosa Kryl.
Flora of Altai, 11, 1904, 705) or green on both sides, almost glabrous
or sparsely covered with short, somewhat stiff hairs (var. glabrata Kryl.
FI. Alt. (Flora of Altai), III, 1904, 705), lamina 3-9 cm long and 1.5—
3 cm wide; basal leaves usually withering before anthesis, narrowed
into short petiole; upper cauline leaves sessile, with weakly decurrent
base. Capitula narrow-campanulate, about 15 mm long and 5-6 mm
wide with florets, crowded at stem apex in dense corymb, subtended by
apical leaves; stem usually somewhat flattened below inflorescence, less

474

often besides terminal corymb, solitary capitula originate from axils of
upper leaves. Involucre usually blackish-violet, pubescent; outermost
involucral bracts in type specimens herbaceous, green, lanceolate,
acuminate, almost as long as inner bracts or slightly shorter, less often
almost half as long, more or less pubescent along margin and surface;
subsequent bracts usually dark-colored, ovate, acuminate, shorter; inner
bracts oblong, subobtuse, colored, somewhat densely hirsute, longer than
middle bracts. Receptacle squamose, scales linear-subulate, lustrous, about
6-7 mm long mostly in middle; florets pink-lilac, about 10-11 mm
long, narrow part of tube about 5—6 mm long, broad part about 2 mm
long, lobes of limb about 3 mm long; anthers violet, anther appendages
hairy. Pappus double; outer pappus bristles unequal, 1 to 4-5 mm long,
fragile, scabrous or finely plumose; inner bristles long-plumose, 8—10
mm long. Flowering VII—VIII.

Rocks, stony screes, tundras, gravelly river banks in alpine and upper
subalpine belt—Western Siberia: Irtysh (eastern part), Altai; Eastern
Siberia: Angara-Sayans (southeastern part). General distribution:
Mongolia. Described on the basis of specimens from Riddersk. Type in
Leningrad.

Note. The type material is characterized by leaves that are white-
tomentose on the lower surface; transitions to leaves glabrous or almost
glabrous on both sides are found. Var. tomentosa Kryl. was described
earlier (1856) as var. altaica Turcz.

S. foliosa Ldb. is phylogenetically very close to the North American
species S. pseudofoliosa Lipsch., which I split off from S. americana
Eaton. See also the note to S. sukaczevii Lipsch.

75. S. pricei Simps. in Journ. Linn. Soc. Bot. XLI (1913) 426; Kryl.
Fl. Zap. Sib. XI, 2919; Popov, Fl. Sr. Sib. II, 762.—S. alpina B. subacaulis
Ldb. Ic. pl. Fl. Ross. I (1829) 17; Ej. Fl. alt. 1'V, 27; DC. Prodr. VI, 535;
Ldb. FI. Ross. II, 2, 669; Kryl. Fl. Alt. Ill, 707.—S. subacaulis (Ldb.)
Serg. in Sistemat. Zam. Gerb. Tomsk. Univ. 2 (1941) 4.—S. pumilio
Fisch. ex DC. Prodr. VI (1837) 535 nomen.—S. foliosa y. pygmaea
Turcz. Fl. baic.-dahur. II, 1 (1856) 116 and Herb.—S. alpina 5. hypoleuca
a. minima Herd. in Bull. Soc.-Nat. Mosc. XLI, 3 (1868) 40.—Ic.: Simps.
op. cit. (1913) t. 21, fig. 1-7; Ldb. op. cit. (1829) tab. 73.

Perennial. Typical small alpine plants with rhizome, often many-
headed, developing floriferous shoots and rosettes of basal leaves; stems
usually densely covered at base with blackish-brown remains of previous
year’s basal leaves, (1)3—-10(12) cm high in type specimen, arachnoid-
hairy or glabrous, sometimes reddish-violet together with leaves, usually
densely leafy up to inflorescence. Leaves hard, coriaceous, distinctly
bicolored, green and glabrous above, densely white-tomentose beneath;

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475

lower and middle leaves lanceolate, oblong-ovate or oblong, indistinctly
cuneate at base, lamina 3-8 cm long and 0.5—1.2 cm wide and petiole
one-fourth to half as long as lamina, less often almost as long; lower
leaves persistent even after anthesis, often as long as inflorescence or
longer in small, particularly stemless specimens; upper leaves sessile,
not decurrent, oblong or linear-lanceolate, in most specimens approximate
with inflorescence; all leaves distinctly toothed, teeth callose with
horizontal or decurved cusp. Capitula, including florets, 1-1.3 cm long,
aggregated in groups of 5—15(20) in dense. corymbs. Involucre 5-6 mm
wide; involucral bracts light-colored, sometimes reddish-violet; outer
bracts ovate or lanceolate, usually attenuate in subulate, sometimes
recurved cusp, almost as long as innermost bracts or slightly shorter;
outer and middle bracts covered with long white hairs along margin and
outside (pubescence on several bracts sometimes glabrescent); inner bracts
densely villous above. Receptacle sparsely squamose, scales linear-
subulate, unequal, up to 3 mm long, lustrous; florets pink-violet, 8—9
mm long, narrow part of tube about 5 mm long, almost as long as
broad part with lobes of limb; anthers dark, anther appendages ciliate.
Pappus double, outer pappus bristles about 2-3 mm long, serrate-
scabrous or short-plumose, inner bristles 7-8 mm long, long-plumose;
achenes 5—6.5 mm long, ribbed, mottled, often dark-colored above.
Flowering VII—VIII.

Gravelly and moss-lichen tundras, rocks, glacial moraines and cirques
in alpine belt.—Western Siberia: Altai; Eastern Siberia: Angara-Sayans
(Sayans), Dauria (southeastern part), Lena-Kolyma (Vitim Plateau).
General distribution: Northern Mongolia. Described from the Tannu-
Ola Range in the Tuva Republic. Type in Kew(?).

Note. It is closest to S. congesta Turcz., from which it is
distinguished by the white-tomentose back side of the leaves, and to S.
sukaczevii Lipsch. It differs from the latter species by the shorter height,
the outer involucral bracts in type form subulate-acuminate to almost
equal to or slightly shorter than the innermost bracts (in typical
S. sukaczevii, involucral bracts are clearly imbricate), and more dense
corymbs. However, in the Herbarium of the Botanical Institute, Academy
of Sciences of the USSR, there is a series of specimens (hybrids?)
intermediate between S. sukaczevii and S. pricei. The limited Altai
material of S. pricei is not fully analogous to the Sayan material
(the middle involucral bracts in several specimens are distinctly shorter
than the inner ones; the outer involucral bracts are equal, shorter,
or longer than the inner; the leaves are sometimes almost entire).
Detailed comparative observations in nature are needed for both S. pricei
and S. congesta. There probably are hybrids between them. Compare
also the note to S. congesta.

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476

76. S. congesta Turcz. in DC. Prodr. VI (1837) 535; Ldb. FI. Ross.
II, 2, 662; Turcz. Fl. baic.-dahur. I], 1, 115; Herd. in Bull. Soc. Nat.
Mosc. XLI, 3 (1868) 44 (incl: a. humilior s. kiriloviana Herd. and B.
elatior s. kusnetzoviana Herd.); Popov, Fl. Sr. Sib. II, 763.

Perennial. Plants vary from almost stemless forms (2-4 cm high) to
forms with distinctly developed stem (7—12 cm high); stem often weakly
flattened, sulcate, simple, almost glabrous or pubescent especially below
inflorescence. Leaves thicker, hard (compare type material), sometimes
thinner, soft, oblong-lanceolate or lanceolate; all leaves acuminate, more
or less toothed, teeth callose at apex with horizontal or recurved short
cusps, lamina almost glabrous on both sides, with punctate glandular
hairs beneath; lower leaves usually broader (up to 1.5 cm wide), short
petiolate (in cauline leaves, petiole sometimes almost as long as lamina
and weakly woolly), narrow-cuneate at base; middle and upper leaves
almost sessile or short-petiolate, not decurrent, 0.5—1 cm wide; uppermost
leaves usually linear or lanceolate-linear in type specimen, narrow,
approximate with inflorescence. Capitula usually numerous, on short
peduncles, forming dense corymb. Involucre dark or light colored, 5—7
mm wide; involucral bracts ovate or triangular-acuminate, dorsally
pubescent as also at apex, especially inner bracts, with long, stiff, white
hairs; outer bracts almost as long as or slightly shorter than inner.
Receptacle squamose, scales linear-subulate, unequal, lustrous. Florets
(according to Truczaninow) scarlet, about 9(10) mm long, narrow part
of tube about 5 mm, almost as long as broad part including lobes
of limb; anther appendages ciliate. Pappus dirty white, double, outer
pappus bristles numerous, unequal, up to 2 mm long, serrate-scabrous,
fragile, inner bristles long-plumose, about 8 mm long. Flowering VII—
VI.

Meadow, gravelly, shrubby and moss-lichen tundras, rocks in alpine
zone.—Eastern Siberia: Angara-Sayans (Sayans), Tuva Republic, Lena-
Kolyma (? Vitim, Yablonovyi Range). Endemic. Described from alpine
zone of Tsagan-Gol (eastern Sayans). Type in Leningrad.

Note. The species is very close to S. poljakovii Glehn; compare the
discussion in the note to the latter. Specimens collected from the Vitim
Plateau are very close to S. congesta Turcz. in the nature of their
involucral bracts, shape and consistency of-leaves, etc., but the leaves
in some of them are white-arachnoid-woolly on lower side (with the
gamut of transitions to weak pubescence). With such pubescence they
approach S. pricei Simps. To all appearance, these are natural hybrids
of S. congesta Turcz. x S. pricei Simps. (= S. korotkyi Lipsch. in sched).

77. S. poljakovii (= poljakowi) Glehn. in Tr. Peterb. Bot. Sada, IV,
1 (1876) 60; Popov, Fl. Sr. Sib. II, 759.

477.

Perennial. Stem 10—20 cm high, sulcate, weakly flattened, simple,
sparsely pubescent with short crisped hairs or almost glabrous. Leaves
thin, glabrous, with more or less numerous, sessile, punctate glandular
hairs beneath, sometimes with occasional white hairs along midrib;
middle leaves ovate or ovate-triangular, acute, petiolate, petioles not
_ decurrent or slightly decurrent as weakly distinctly toothed wings; lower
leaves persistent up to anthesis, ovate, 3.5—-6 cm wide, with indistinctly
cuneate base and longer petiole; upper leaves narrower, long-acuminate,
sessile, often approximate with corymb; all leaves distinctly and
frequently toothed, teeth unequal, subulately acuminate at apex; leaf
margin more or less glandular-hairy between teeth. Capitula on glandular-
hairy peduncles, in dense corymb. Involucre distinctly three- or four-
rowed; involucral bracts almost equal; outer bracts lanceolate, terminating
from broader base into long subulate cusp. Recetacle squamose, scales
linear-subulate, unequal, 2-4 mm long, lustrous; florets about 8-9 mm
long, narrow part of tube about 4 mm long, almost as long as broad
part with lobes of limb; anther appendages ciliate. Pappus double; outer
pappus bristles minute, fragile, unequal, serrate-scabrous, inner bristles
about 8 mm long, long-plumose; immature achenes about 4 mm long,
glabrous, sulcate. Flowering VII—VIII. (Plate XXXV, Fig. 2.)

Alpine meadow patches, outcrops of stones.—Eastern Siberia: Lena-
Kolyma (along the Vitim and Barguzin rivers, and Udokan Range).
Endemic. Described from the Uksumikit Valley (Vitim River). Type in
Leningrad. ;

Note. Quite close to S. congesta Turcz. The specimens, on the basis
of which Turczaninow described S. congesta, are almost stemless, small-
leaved, and typically alpine. The specimens, on the basis of which Glehn
separated S. poljakovii, are distinctly stemmed and, evidently, originate
from lower altitudes. Glehn distinguished S. poljakovii from S. congesta
by the following characters: broader, thin ovate leaves, larger, more
strongly acuminate unequal teeth along the leaf margin, more strongly
developed stem, dense equal receptacular scales (outer ones not shorter).
S. poljakovii needs further observations in nature because, besides the
authentic specimens in the Herbarium of the Botanical Institute, Academy
of Sciences of the USSR, it is represented only by the single sheet of L.
Tyulina from the Barguzin Region (No. 768) and the two sheets of
Sviridenko from the Kalar Region. It is very probable that in the future
it will have to be combined with S. congesta Turcz. into a single species
for the following reasons. Even Herder (Bull. Soc. Nat. Mosc., XLI, 3
(1868) 45) already described two varieties in S. congesta: humilior seu
kiriloviana Herd., with broader and unequally dentate leaves, and elatior
seu kusnetzoviana Herd., with narrower and uniformly toothed
leaves. The limited material of S. congesta and S. poljakovii shows:

484

486

478

1) the shape, width, consistency and dentation of the leaves vary greatly;
2) stem height also varies (stemless, intermediate, and distinctly stemmed
forms are known in S. congesta). As regards the receptacular scales in
the authentic specimen of S. poljakovii, there is no difference from the
dwarf forms and the type material of S. congesta—on closer examination,
the receptacular scales in S. poljakovii were found to be unequal in
length. However, I have decided not to combine S. poljakovii with S.
congesta Turcz. until new material is obtained on the former species.

78. S. tilesii Ldb. Ic. pl. Fl. Ross. I (1829) 15; Ej. Fl. alt. IV, 15,
in nota; DC. Prodr. VI, 536; Ldb. FI. Ross. II, 2, 662; Lipschitz in Bot.
Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI, 450.—S. ftilesii f.
subacaulis Kom. Fl. Kamch. III (1930) 180.—Serratula tilesii Ldb. in
Mém. Acad. Sc. Pétersb. V (1815) 562.—Saussurea nuda y. tilesii Rg.
in Nouv. Mém. Soc. Nat. Mosc. XI (1859) 106.—S. ledebouri 6. tilesii
Rgl. in Herd. Bull. Soc. Nat. Mosc. XLI, 3 (1868) 41.—S. ledebouri
auct. (non Herd.): Tolmatchev in Tr. Polyarn. Komiss. Akad. Nauk SSSR,
13 (1932) 169.—S. alpina var. subacaulis auct. fl. Arct. (non Ldb.) p.
max. p.—Heterotrichum arcticum MB. ex Herd. op. cit. (1868) 43.—
Ic.: Ldb. op. cit. (1829) Plate LX.

Perennial. Rhizome oblique, woody; plants usually prostrate, (3)5—
10 cm high, rarely, up to 15 cm, densely leafy; root collar densely
covered with brownish remains of leaves; stems often weakly swollen
or flattened, sulcate, usually arachnoid hairy-tomentose, especially below
inflorescence, mixed with lustrous glandular hairs, less often almost
glabrous. Leaves oblong or oblong-lanceolate, crowded in basal part,
fairly unequal in width, from 3—5 mm to 15 mm, usually more or less
strongly attenuate-acuminate, green above, white-tomentose or arachnoid
hairy-pubescent beneath with more or less numerous lustrous glandular
hairs on both sides (sometimes absent); sometimes weakly involute,
crenate-toothed in type specimen, teeth with callose-cartilaginous cusps;
basal leaves larger, almost as long as stem in small plants, petiolate,
petioles winged, cuneate at base; cauline leaves sessile or short-petiolate;
upper leaves linear or oblong-lanceolate, usually approximate with
inflorescence, of these one to three often larger than it or as long.
Capitula in type specimen numerous, pedunculate, compactly aggregated
at stem apex in globose or semiglobose dense corymb, corymbs absent
on lateral branches. Involucre densely sericeous-villous, involucral bracts
acute, but without long cusps; outer bracts in type specimen shorter
than linear-lanceolate inner, less often as long or almost as long, and
both with numerous, long, white, squarrose, hairs, glochidiate. Receptacle
densely squamose; scales linear-subulate, white, lustrous, unequal, longest
scales about 6 mm long; florets dark pink, about 1.2 cm long, narrow

479

485 Plate XXXVI.

1—Saussurea tilesii Ldb.; 2—S. soczavae Lipsch; 3—S. pseudo-tilesii Lipsch.

480

part of the tube about 6 mm long, as long as broad part with lobes of
limb; anther appendages ciliate. Pappus double, outer pappus bristles up
to 2 mm long, persistent, short-plumose, inner bristles bicolored,
ferruginous below, white above, all bristles equal, long-plumose, about
13 mm long; achenes about 5 mm long, sulcate, glabrous, dark brown
or blackish. Flowering VII—VIII. (Plate XXXVI, Fig. 1.)

Moss, lichen and shrubby tundras, alpine meadows, stony screes,
rocks.—Arctic: Arctic Siberia; Eastern Siberia: Yenisei (northern part),
Lena-Kolyma (northern part); Far East: Chukotka, Anadyr, ?Kamchatka
(only in alpine belt). Endemic. Purportedly described from Kamchatka
on the basis of collections of Tilesius; type in Leningrad. Apparently,
the authentic specimen was collected not from Kamchatka, but from
another more northern, most probably, Arctic location.

Note. “S. tilesii” of the authors of the Kamchatka flora is sharply
distinguished from S. tilesii Ldb. s. str. and was described as a separate
species, S. pseudo-tilesii Lipsch. For details about the relationship of
this pair of species see the article of Lipschitz (op. cit. 1954). The
extreme southern outposts of the distribution of S. tilesii Ldb. on the
continent are taken to be the upper reaches of Bureya (Dusse-Alin Range)
and Sikhote-Alin. However, the study of new specimens from Dusse-
Alin showed that they represent a unique race: S. soczavae Lipsch. (cf.
below). The recently described species by Hulten from Alaska, S. viscida
Hult. is close to S. tilesii Ldb. The author characterized it by the presence
of numerous scattered, sticky hairs on the upper and lower leaf surfaces
and along the margin as well as on the stem. The specimens of S. tilesii
Ldb. collected from Wrangel Island (B.N. Gorodkov, 1938) and preserved
in the Herbarium of the Botanical Institute, Academy of Sciences of the
USSR are close to S. viscida Hult. in the nature of the pubescence, but
otherwise show no differences from typical S. tilesii Ldb. —

79. S. soczavae Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XXI (1961) 376.

Perennial. Rhizomes prostrate; stem erect, 10-18 cm high, simple,
sulcate, wingless, almost glabrous or weakly arachnoid-hairy, sparsely
leafy, internodes long. Leaves lanceolate or oblong-lanceolate, up to 6
cm long and 1.5—2 cm wide, almost cuneate at base, bicolored, dark green,
almost glabrous or with punctate glandular hairs above, white-tomentose
beneath, toothed, teeth unequal, cuspidate; basal, lower, and middle cauline .
leaves on 2—2.5 cm long petioles, weakly winged, not decurrent on stem;
middle and upper cauline leaves gradually reduced, narrow-lanceolate,
sessile, uppermost leaves approximate with inflorescence, linear or linear
lanceolate, almost entire. Capitula campanulate, 10-12 mm wide, usually

487 numerous, less often two, pedunculate, in terminal corymb, which is

481

not compact or capitate. Involucre campanulate, imbricate, dark-colored,
weakly arachnoid-hairy, glabrescent, 10-11 mm long and 10 mm wide;
involucral bracts three- or four-rowed; outer bracts ovate or lanceolate,
terminating in cusp, shorter than inner, less often almost equal; inner
bracts lanceolate, acute or subacute. Receptacle squamose, scales linear-
subulate, lustrous, unequal, up to 6-7 mm long; florets dark violet(?),
about 10 mm long, narrow part of tube about 5 mm long, as long as
broad part together with lobes of limb; anther appendages ciliate. Pappus
double, slightly exserted from involucre, outer pappus bristles unequal,
up to 5 mm long, fragile, scabrous or finely plumose, inner bristles
plumose, about 9-10 mm long; achenes glabrous. Flowering VIII. (Plate
XXXVI, Fig. 2.) .

Stony slopes of mountains,(?) alpine tundras.—Far East: Uda.
(Dusse-Alin Range). Endemic. Described from the Suluka River. Type
in Leningrad.

80. S. pseudo-tilesii Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVI (1954) 451.—S. tilesii auct. fl. Kamtsch. non Ldb::
Kom. Fl. Kamch. III (1930) 178 (incl. f. glabrata (180) and alpicola
Kom. excl. f. subacaulis Kom.); Hult. Fl. of Kamtch. IV (1930) 211.—
S. alpina 5. maxima Ldb. Fl. Ross. II, 2 (1845-1846) 669.—S. tilesii
var. exaltata Maxim. Prim. Fl. Amur. (1859) 170.—S. ledebouri «.
exaltata C.A.M. and Maxim. ex Herd. in Bull. Soc. Nat. Mosc. XLI, 3
(1868) 43.—S. nuda var. exaltata (Maxim.) Trautv. ex Fedtsch. FI. Ill.
Comm. (1906) 77.—Ic.: Hult. op. cit. (1930) 213 (florets).

Perennial. Rhizome short, oblique, woody; stem 40—60(up to 100)
cm high, erect, simple, sometimes branched only in inflorescence,
glabrous or weakly arachnoid-hairy, sulcate, wingless. Leaves ovate,
oblong-linear or broadly linear, acuminate, all leaves except uppermost
on long and weakly winged petiole (petioles of basal leaves as long as
or almost as long as lamina); lamina 5—20 cm long and up to 5-6 cm
wide, cuneate or indistinctly roundish at base, serrate-toothed, teeth with
straight or weakly bent cusp, green and almost glabrous above, finely
white-tomentose beneath. Capitula campanulate, 1-1.2 cm long with
florets and 0.8—1 cm wide, on more or less long, densely arachnoid-
hairy peduncles, aggregated in dense terminal corymb or corymbose
panicles. Involucre usually white-violet, involucral bracts arachnoid hairy-
pubescent, or almost glabrous, sometimes with scattered white hairs only
in upper part and along margin; all bracts, especially outer, attenuate at

apex into long, thin, usually recurved cusp; outermost bracts linear, linear-
lanceolate or ovate, almost as long as even longer than inner ones,
however not deforming imbricate aestivation of involucre. Florets pink-
violet, about 8—9 mm long; narrow part of tube slightly broadened at

482

base, hollow, about 4-5 mm long, almost as long as broad part with
lobes of limb; anthers blackish-blue, anther appendages ciliate. Receptacle
densely squamose, scales linear-subulate, lustrous, unequal, up to 6 mm
long; pappus double, outer pappus bristles unequal, up to 2 mm long,
serrate, scabrous, fragile, inner bristles long-plumose about 8 mm long,
brownish in lower part, with pure white tips; achenes glabrous. Flowering
VII-VIII. (Plate XXXVI, Fig. 3.)

Valley and subalpine meadows, grass plots, small patches of tundra,
forests of Erman’s birch and larch, alder thickets, thickety outcrops of
lava and slag —Far East: Kamchatka, Commander Islands. Endemic
(replaced by S. oxyodonta Hult. in the Anadyr Region and on the Kuml
Islands). Described from Kamchatka. Type in Leningrad.

Note. | am designating the specimens of Mertens from Kamchatka,
preserved in the Herbarium of the Botanical Institute, Academy of
Sciences of the USSR, as the type of the species which was not indicated
by me in my publication of 1954. S. pseudo-tilesii m. is a tall, lowland
Saussurea, replacing the short, arctic species S. tilesii Ldb. s. str., in
Kamchatka, under which it was wrongly placed by the authors of
Kamchatka flora. The type of S. tilesii Ldb. s. str. evidently does not
originate from Kamchatka. The alpine form of S. pseudo-tilesii m. from
Kamchatka is distinguished by shorter height, crowded reduced leaves,
and a compact inflorescence. However, its characteristic.feature is the
presence of a long cusp on the involucral bracts, always present in this
form. This form was described under the name S. ftilesii f. alpicola
Kom.; since I have included the latter under S. pseudo-tilesii and consider
it as a variety, a new combination had to be made: S. pseudo-tilesii var.
alpicola (Kom.) Lipsch. [Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XXI (1961) 377].

81. S. oxyodonta Hult. Fl. of Kamtch. IV (1930) 207; Kitam. in
Mem. Coll. Sc. Kyoto Univ. ser. B. XIII, 178.—S. ledebouri y. kruhseana
Herd. in Bull. Soc. Nat. Mosc. XLI, 3 (1868) 43.—S. tilesii var. kurilensis
Kudo Fl. Param. (1922) 172; Tatewaki, Vasc. Pl. North. Kuril. 305.—S.
subsinuata auct.: Kom. Fl. Kamch. III (1930) 181; Kom. Izbr. Soch.
VII, 486 p. p. (non Ldb.).—Ie.: Hult. op. cit. (1930) tab. 4a.

Perennial. Plants almost glabrous. Rhizome oblique, covered with
less numerous fibrous roots; root collar covered with remains of old
leaves; stem 10—30(50) cm high, wingless, uniformly and usually sparsely
leafy, simple, sulcate, almost glabrous or with few crisped hairs. Leaves
green above, pale beneath, weakly crisped-pubescent only along veins
and margin on both sides; basal leaves usually persistent till anthesis,
sometimes withering; lower cauline leaves long-petiolate, petioles narrow-
winged, up to 6-10 cm long in vegetative branches; leaves in type
specimen lanceolate or less often ovate and oblong-lanceolate, 6-12 cm

489

483

long and 1—2.5 cm wide, more or less attenuate-acuminate, basally
cuneate or truncate, unequally dentate, teeth acuminate; middle leaves
short-petiolate, up to 10 cm long and up to 2 cm wide; upper leaves
sessile, narrow-lanceolate or linear, with less numerous teeth or entire.
Capitula campanulate, 10-12 cm wide, numerous, in more or less
compact corymb in type specimen, on distinctly glandular hairy
peduncles, 0.5—1 cm, less often 1.5(3.5) cm long. Involucre imbricate,
usually dark-colored, arachnoid hairy-pubescent, three- or four-rowed;
outer involucral bracts lanceolate or ovate, long-attenuate-acuminate,
almost as long as inner bracts; middle bracts narrowed from broad ovate
base; inner bracts lanceolate or linear, short-acuminate, more or less
pubescent on dorsal surface, glochidiate with white hairs at apex.
Receptacle squamose, scales numerous, unequal, up to 6 mm long,
lustrous; florets pink-violet, about 10 mm long, narrow part of tube
about 5 mm long, as long as broad part with lobes of limb; anther
appendages ciliate. Pappus double, outer pappus bristles rather numerous,
unequal, up to 2—3 mm long, serrate-scabrous or finely plumose,
persistent on achenes, inner bristles long-plumose, about 8—9 mm long,
ferruginous below, white above; achenes glabrous, mature achenes dark
brown with dark spots. Flowering VII—VIII.

Moist meadows, moss-covered marshes, small patches of tundra,
birch forests, thickets of shrubs, pebbled floodplains.—Far East: Anadyr,
Kamchatka, Sakhalin (Kuril Islands). Endemic. Described from
Kamchatka. Type in Stockholm.

Note. Where it is in contact, it hybridizes with S. nuda Ldb.,
imitating S. oxyodonta Hult. in growth habit but always having a smooth
and not a chaffy receptacle. Alpine specimens of S. oxyodonta Hult. are
12-15 cm high, with smaller and usually narrower leaves, about 5 cm
long and up to 0.7 cm wide, and a lax corymb. A few vigorous specimens
from Kamchatka, with a tall stem, broader leaves with truncate-cuneate
base and a broad, winged petiole, appear to be hybrids between S.
oxyodonta Hult. and S. riederi Herd.

Series 17. Alpinae Lipsch.—Stem wingless, less often (in S.
stubendorffii) with distinct wings, weakly leafy. Leaves ovate, lanceolate
and linear, glabrous and green above, grayish beneath due to arachnoid-
hairy pubescence or white-tomentose, sometimes glabrous, green, entire
or toothed, flat or weakly revolute. Capitula ovoid or cylindrical, in
terminal, more or less compact corymb, less often in panicle, of
individual corymbs. Involucre distinctly imbricate; outer involucral bracts
ovate, often hardening subsequently, and like other bracts subobtuse,
sometimes acuminate, usually pubescent inner bracts more strongly.
Capitula base usually with one to three lanceolate appendages. Receptacle
squamous.

490

484

82. S. alpina (L.) DC. in Ann. Mus. Paris, XVI (1810) 198; Ej. ;
Prodr. VI (1837) 535 p. p. (excl. B. subacaulis Ldb.); Ldb. Fl. alt. IV,
27 p. p. (excl. B. subacaulis Ldb. and synon. Serratula angustifolia
Willd.); Ej. Fl. Ross. II, 2, 669 p. p. (excl. y. subacaulis Ldb., 5. maxima
Ldb., ¢. hypoleuca Ldb.); Herd. in Bull. Soc. Nat. Mosc. XLI, 3, 36
(excl. y. subacaulis Ldb. and Turcz. 5. angustifolia Rgl., €. hypoleuca
Ldb. cum a. maxima, b. major, c. minor, d. minima, §. foliosa); Turcz.
Fl. baic.-dahur. II, 1 (1856) 114 p. p. (excl. a. communis, B. subacaulis
Ldb.); Hegi, Ill. Fl. VI, 2 (1929) 839; Sergievskaja in Sist. Zam. Gerb.
Tomsk. Un. 2 (1941) 1, excl. var. schischkinii Serg. and var. lancifolia
Serg. p. p.; Kryl. Fl. Zap. Sib. XI, 2918; Juzepczuk in Sp. Rast. Gerb.
Flory SSSR, XIII, 98—S. decurrens Winkl. (non Hemsl.) Catal. sem.
Horti. Bot. Petropol. (1882) 16 (nom. nud.); Ej. op. cit. (1883) 16 (nom.
nud.).—S. alpina var. decurrens Rgl. ex Winkl. in Tr. Peterb. Bot. Sada,
IX, 2 (1886) 424.—Serratula alpina L. Sp. pl. (1753) 816; Willd. Spec.
pl. III (1800) 1641 excl. var. 8—Saussurea alpina sibirica C.A.M. ex
DC. op. cit. (1837) 535 and Turcz. op. cit. (1856) 115.—S. alpina B.
vulgaris Ldb. Fl. Ross. II, 2 (1845-1846) 669.—S. pohlei Gandoger in
Bull. Soc. bot. France, LVI (1909) 533.—Ic.: Hegi, op. cit. (1929) tab.
270, fig. 2 and fig. 536, 537.—Exs.: GRF nos. 1331, 2949.

Perennial. Rhizome prostrate, stem (5)8—10 cm to 20-50 cm high,
erect, solitary, less often two or more, simple or sometimes branched in
inflorescence, usually green, less often dark colored, more or less
pubescent or glabrous. Leaves green and glabrous above in type
specimen, grayish arachnoid-hairy beneath due to long, intertwined, fine
hairs (var. vulgaris Ldb.) or glabrous (var. viridis Turcz.), sometimes
with sparse sessile glandular hairs, with indistinct cartilaginous teeth or
entire, leaf margin not revolute in type specimen; basal leaves persistent
till anthesis; lower cauline leaves petiolate, ovate-lanceolate, lanceolate,
oblong or elliptical lamina, (4)8—20 cm long and 1.5—4 cm wide, usually
attenuate-acuminate, sometimes short-acuminate, less often obtuse, petiole
less often almost as long as lamina, usually one-fifth to two-thirds as
long, midrib prominent, leaf margin usually with minute horizontal or
slightly divergent carlaginous teeth; middle cauline leaves often remote,
elliptical or lanceolate, acuminate, sessile, less often weakly decurrent
(var. decurrens (Winkl.) Rgl.); uppermost.leaves lanceolate or linear,
acuminate, sessile, usually not decurrent, 2-5 cm long and 1—8 cm wide,
entire. Capitula terminal on main stem or lateral branches, in dense or
lax corymbs. Involucre distinctly imbricate, 5—7(10) mm wide, often
becoming hard; involucral bracts dark-colored, sometimes greenish; outer
bracts broadly ovate or ovate-lanceolate, subobtuse or subacute, middle
bracts ovate, subobtuse, both weakly crisped-hairy or arachnoid-hairy
along margin and on dorsal surface, sometimes almost glabrous (var.

491

485

viridis Turcz.); inner bracts oblong, subobtuse, densely pubescent with
long, white, soft, semidivergent hairs, with one to three lanceolately
acuminate appendages at base of capitulum, as long as or slightly longer
than outer involucral bracts. Receptacle densely squamose; florets violet-
pink, about 1.1 cm long; narrow part of tube about 5 mm long, almost
as long as broad part with lobes of limb; anther appendages strongly
hairy. Pappus about 9 mm long, inner pappus bristles long-plumose;
achenes 3-4 mm long, brownish. Flowering VII-IX.

Stony slopes, screes, moss and lichen covered tundras in alpine
belt; open deciduous forests; cedar-larch and spruce-larch-cedar forests,
willows of forest belt; alder thickets; grassy, hummocky tundras, rocks,
pebbles of the arctic region— Arctic: Arctic Europe, Novaya Zemlya;
European Part: Upper Dniester, Ladoga-IImen, Karelia-Lapland, Dvina-
Pechora, Volga-Kama (Ural); Western Siberia: Ob’, Upper Tobol, Irtysh
(western part), Altai; Eastern Siberia: Yenisei, Angara-Sayans, Lena-

Kolyma; Soviet Central Asia: Pamiro-Alai, Tien Shan. General

distribution: Arctic, Scandinavia, Central Europe, Altantic Europe (Great
Britain), Mongolia, Dzhungaria-Kashgaria (Kuldzha). S. alpina (L.) DC.
is described from Lapland; type, evidently, in London. S. pohlei Gandoger
is described from collections of Pohle from the Kolguev Island; type in
the Herbarium of Gandoger; isotypes(?) in Leningrad (they do not fully
conform to the diagnosis). S. decurrens Winkl. reported on the basis of
specimens raised in the Petersburg Botanical Garden from seeds by A.
Regel from Chinese Turkestan; preserved in Leningrad.

Note. An extremely polymorphic, widely distributed boreal-alpine
species requiring a monographic study based on extensive material
collected from throughout its range. Undoubtedly, it includes numerous
ecological forms correlated with the various ecological conditions, as
well as races. Among the most distinct races are: S. depressa Gren. and
Godr. (western Alps), S. angustifolia (Willd.) DC. (North America,
islands of the Bering Strait), S. densa Rydb. (North America), S. pricei
Simps. (Altai, Sayans), S. sukaczevii Lipsch. (eastern Siberia), S. ambigua
Kryl. (Altai, Tuva, Mongolia), S. stubendorffii Herd. (eastern Siberia).
Less distinct is Estonian S. alpina, described as S. esthonica Baer. ex
Rupr., which is connected by a series of transitional forms to typical S-:
alpina and, therefore, is reduced by many authors to the level of a
subspecies or variety of S. alpina (see note to this species). Some of
these races growing within the limits of the USSR, are treated here
under binomial names in accordance with the tradition of the Flora of
the USSR. The following characters of S. alpina show significant
variation: stem height; shape, size, pubescence and nature of the margin
of the leaves and the degree of leaf decurrence along stem; type of
inflorescence; shape of involucral bracts.

492

486

Like many other species of the genus, S. alpina also hybridizes.
For instance, hybrids with S. controversa DC. (= S. uralensis Lipsch.)
(Urals) and with S. tilesii Ldb. (Yakutia) are known.

An infraspecific classification of S. alpina has been worked out by
De Candolle, Ledebour, Turczaninow and Herder, and recently by
Sergievskaja. Let us mention the most clearly defined varieties: 1) var.
viridis Turez. (in DC. Prodr. V1, 535; Fl. baic-dahur. II, 114 = var.
glabra Ldb. Fl. Ross. II, 2, 669)—Leaves almost glabrous on both sides,
green, oblong-lanceolate or ovate; outer involucral bracts almost glabrous,
inner pubescent; 2) var. vulgaris Ldb. (op. cit. 669) does not in any
way differ from typical S. alpina; 3) var. krylovii Serg. (Sistemat, Zam.
Gerb. Tomsk. Univ. No. (1941) 2)—Stem 24-35 cm high; leaves oblong-
ovate or oblong, 1-3 cm broad, decurrent on stem, arachnoid-hairy
beneath; 4) var. lancifolia Serg. (op. cit. (1941) 2)}--Stem 35-60 cm
high; leaves lanceolate, more or less long-acuminate, 6—15 cm long and
4-12 mm wide, decurrent, arachnoid-hairy beneath, identical with S.
stubendorffii Herd. p. p.; and 5) var. schischkinii Serg. (op. cit. (1941)
2) undoubtedly is identical with S. stubendorffii Herd., under which it is
placed as a synonym.

83. S. sukaczevii Lipsch. in Byull. Mosk. Obshch. Isp. Prir. Otd.
Biol. LIX, 6 (1954) 71; Popov, Fl. Sr. Sib. II, 760.—S. alpina eg.
hypoleuca Ldb. Fl. Ross. II, 2 (1845-1846) 669; Herd. in Bull. Soc.
Nat. Mosc. XLI, 3 (1868) 38 and 39.—S. alpina 6. hypoleuca b. major
seu genuina Herd. Ibid. (1868) 39.—S. alpina 5. hypoleuca c. minor
Herd. Ibid. (1868) 40.—S. denticulata Turcz. Fl. baic.-dahur. II, 1 (1856)
115 (mon Wall. nec. Ldb.).—S. alpina var. leucophylla Turcz. ex
Herd. op. cit. (1868) 38.—S. foliosa B. dentosa Turcz. in DC. Prodr. VI
(1837) 536 p. p.; Turcz. op. cit. (1856) 116 p. p.; Herd. op. cit. 38 in
syn. p. p.—S. serrata var. pygmaea Turcz. in herb. and ex Herd. op.
cit. (1868) 40 in syn.—S. alpina a. communis Turcz. op. cit. (1856)
114 and herb.

Perennial. Rhizome oblique, with fibrous roots; root collar covered
with brown-black, partially split remains of old leaves; stem erect,
solitary, simple, not winged, green or often red-violet, weakly arachnoid
hairy-pubescent or almost glabrous, sometimes, mainly below corymb,
flattened, 12—30(35) cm high; high mountain specimens up to 7 cm
high (var. pumila Lipsch.). Leaves green and glabrous above, densely
white tomentose beneath with prominent midrib; all leaves shallow-
toothed with teeth more or less sparse, callosely-pointed, horizontal or
weakly deflexed, often slightly revolute; basal leaves rersisting till
anthesis, sometimes withering; lower cauline leaves in type specimen
lanceolate, less often oblong-ovate, acuminate, up to 12 cm long with

493

487

petiole and (0.5)1—1.5(2) cm wide, petioles half to two-thirds as long as
lamina, less often almost as long; middle and upper cauline leaves
resembling lower leaves in shape, usually narrower, sessile or subsessile,
not decurrent; uppermost leaves linear or linear-lanceolate, acuminate,
sessile, 2-4 cm long and 1-5 mm wide, entire. Capitula cylindrical or
narrow-campanulate, on 1-3 cm long peduncles in type specimen
(sometimes peduncles poorly developed), usually adjoining. somewhat
lax terminal corymb. Involucre imbricate, 5—7 mm wide, involucral bracts
dark-colored, sometimes green; outer bracts ovate, often becoming hard,
attenuate toward apex, acuminate, middle bracts ovate, obtuse, both more
or less arachnoid-hairy along margins and on dorsal surface; inner bracts
oblong, subobtuse, usually glochidiate with white hairs in upper part
with (one) two or three lanceolate bracteal leaves present below capitula
which are as long as outer involucral bracts or longer. Receptacle densely
squamose, scales linear-subulate, lustrous, unequal, up to 6 mm long;
florets pink-violet, about 1.2 cm long, narrow part of tube about 5.5—6
mm long, almost as long as broad part together with lobes of limb;
anther appendages ciliate. Pappus double, outer pappus bristles unequal
(up to 3 mm long) serrate-scabrous; inner bristles about 1 cm long,
long-plumose; achenes mottled. Flowering VII—VIII. (Plate XXXV,
Fig. 1.)

Sphagnum moss-marshes; grass areas, gravelly and moss-lichen
tundras, rocks, stony screes in alpine zone and in upper part of forest
zone.—Eastern Siberia: Angara-Sayans, Yenisei (southern part), Dauria.
Endemic. Described from Kultuk-on-Baikal. Type in Leningrad.

Note. The species, particularly its alpine variety (var. pumila
Lipsch.), is close to S. pricei Simps., from which it differs by greater
height, imbricate involucral bracts, with the outer ones distinctly shorter
than the inner ones, and also by longer, attenuate-lanceolate leaves and
receptacular scales up to 6 mm long. Hybrids between S. sukaczevii and
S. pricei are possible in their areas of contact. Cf. the note to the latter
species.

84. S. esthonica Baer ex Rupr. in Beitr. Pflanzenk. Russ. Reichs,
IV (1845) 21; C.A.M. in Mém. Acad. Sc. Pétersb., VI sér., sc. natur.
VII, Bot. 5 (diagn. emend.); Juz. in Sp. Rast. Gerb. Flory SSSR, XIII,
99, No. 3950.—S. alpina subsp. esthonica (Baer ex Rupr.) Kupff. in
Korrespondenzbl. Naturforsch.-Ver. zu Riga, XLV (1902) 94.—S. alpina
auct. fl. Pribalt. Ross.: Wiedemann and Weber, Beschr. der. phanerogam.
Gawiachse Est-, Liv-u. Kurlands (1852) 483; Klinge, Fl. v. Est-, Liv-u.
Curl. (1882) 317; Lehmann, Fl. v. Poln.-Livl. (1895) 268.—Ic.: Kupff.
op. cit. fig. B.—Exs.: Eesti taimed Tartu Ulik. Botaanikam. Nos. 27,
27a.

494

488

Perennial. Stem 30—40(60) cm high, less often shorter, solitary, green
or violet, slender, glabrous or, with weak and easily rubbed off
pubescence mainly in upper part. Leaves in type specimen narrow-
lanceolate, young leaves flocculose-pubescent, glabrescent or subglabrous,
narrowed at base and weakly cuneate, not decurrent, long acuminate,
less often scarcely acuminate; lower cauline leaves long-petiolate, oblong-
lanceolate, up to 20 cm long and 1-2 cm (less often up to 3.5 cm)
wide, toothed with callosely pointed teeth; middle cauline leaves like
lower but narrower and on shorter petioles, gradually reduced toward
stem apex, usually less distinctly toothed, usually more than 10 on a
stem; uppermost leaves sessile, reduced, narrow-linear, entire, slightly
involute. Capitula (3)10-15 (less often up to 25) in paniculate
inflorescence, comprising individual lax corymbs; peduncles of capitula
usually longer than latter. Involucre imbricate, light green or dark-colored;
involucral bracts villous due to numerous soft hairs; outer bracts reduced,
ovate or rotund, subacute; middle and inner bracts ovate-oblong, scarcely

acuminate. Receptacle densely squamose, scales linear-subulate, lustrous,

about 6—7 mm long; florets (according to observations of A. Vag in
nature) lilac-colored with strong blue tinge, about 1.3 cm long, narrow
part of tube about 6 mm long, almost as long as broad part together
with lobes of limb; anther appendages ciliate. Achenes cylindrical,
glabrous; pappus dirty white, double, outer pappus bristles numerous,
deciduous, serrate, up to 3-4 mm long, inner bristles plumose, about
11-12 mm long. Flowering VII—VIII.

Marshy meadows, marshes, scrubs——European Part: Baltic Region
(Estonian SSR); ?Ladoga-IImen. Described from Estonia. Type in
Leningrad.

Note. Of the species aggregate S. alpina; it mainly differs from the
typical (Lapland) specimens of S. alpina by a taller stem, narrower basal
leaves, numerous (often exceeding 10), weakly pubescent (often glabrous)
cauline leaves, a lax paniculate inflorescence with peduncles longer than
the capitula. Kupffer and Juzepczuk considered S. esthonica an endemic
race of the Estonian SSR. The first author assumed that the range of S.
esthonica is sharply isolated from the northern localities of S. alpina
(known to him from the former Olonetsk and Arkhangelsk provinces).
Juzepczuk realized the mistake of Kupffer, as bridging localities of S.
alpina were subsequently found in the Leningrad and Novgorod
provinces. However, Juzepczuk did not refer the latter specimens to
true S. alpina. At the same time, an examination of the accumulated,
rather extensive herbarium collections from the northwest European Part
of the USSR shows that-a series of plants is found there, particularly
growing on limestone and peat marshes, which are intermediate between
S. esthonica and S. alpina s. str. Some of them are so close to the

489

former that they could quite rightly be referred to it. These questions
can be finally resolved only after special studies in nature. It appears
that S. esthonica is not a local Estonian race. On the other hand, S.
esthonica is morphologically quite close to one of the east Siberian
races of S. alpina—S. stubendorffii Herd., which is also found in Yakutia.
Morphologically, S. esthonica differs from S. stubendorffii chiefly as
follows: leaves beneath in type glabrous or very weakly pubescent; lower
leaves narrowly lanceolate (1-2 cm wide); cauline leaves usually more
than 10, not decurrent on stem; inflorescence consists of fewer capitula
because the floriferous branches are fewer; florets lilac-colored with
bluish tinge, and not purple. All these differences between the species
are relative in nature. The chief reason preventing us from combining
the two species is their distinct ranges. Following the practice of the
Flora of the USSR, they are treated here under binominal names.
Formally speaking, it would be more correct to call our plants S. alpina
subsp. esthonica (Baer ex Rupr.) Kupff. and S. alpina subsp. stubendorffii
(Herd.) Lipsch.

85. S. stubendorffii Herd. in Bull. Soc. Nat. Mosc. XLI, 3 (1868)
26; Freyn in Oest. bot. Zeitschr. LII, 7, 281; Popov, Fl. Sr. Sib. II,
760.—S. denticulata Ldb. y. viluiensis Herd. op. cit. (1868) 26.—S.
alpina var. schischkinii Serg. in Sist. Zam. Gerb. Tomsk. Univ. 2 (1941)
2; Lipschitz in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XXI, 377.—
S. alpina var. lancifolia Serg. op. cit. (1941) 2.—S. amurensis auct. fl.
Sib. (non Turcz.) p. p.—S. serrata auct. fl. Sib. p.p. non DC.—S. alpina
auct. fl. Sib. p. p.

Perennial. Stem erect, 30—70(80) cm high, slightly floccose-arachnoid
hairy-pubescent or glabrous, sulcate, less often simple, often branched
above or even from middle; sometimes narrow-winged due to decurrent
leaves, usually wingless. Leaves soft, lanceolate, oblong-lanceolate or
even almost ovate, green and weakly scabrous above, more or less
densely grayish or whitish flocculose-arachnoid-hairy (pubescence
glabrescent); basal and lower cauline leaves 2.5—-3 cm wide, persistent
at anthesis, petiolate, distinctly toothed, teeth callose, remote; upper
leaves narrow, linear or linear-lanceolate, almost entire, sessile or
subsessile together with middle leaves, more or less decurrent on stem
and sometimes forming narrow wing. Capitula pedunculate, in more or
less numerous corymbs terminating on pubescent peduncles; individual
corymbs usually unequal in height, forming more or less lax paniculate
inflorescence. Capitula in type specimen up to 1.5(2) cm long. Involucre
ovoid or cylindrical, four- or five-rowed; involucral bracts imbricate,
light-colored or dark, covered with long hairs; outer bracts ovate,
subacute, inner linear-lanceolate, acuminate. Receptacle squamose, scales

496

490

linear-subulate, lustrous, up to 6 mm long; florets red, about 1.2 cm
long, narrow. part of tube about 6 mm long, as long or almost as long
as broad part together with lobes of limb; anther appendages ciliate.
Pappus double, outer pappus bristles less numerous, serrate, fragile, about
2-3 mm long, inner bristles long plumose, about 1 cm long; achenes
cylindrical. Flowering VII-—VIII.

Coniferous and deciduous forests, willow stands, meadows, peat
bogs.—Western Siberia: Ob’ (eastern part), Altai; Eastern Siberia: Angara-
Sayans, Yenisei, Lena-Kolyma. Endemic. Described from the eastern
part of Siberia, from the Khorma and Biryusa rivers. Type in Leningrad.

Note. Herder compared this species with S. serrata DC. (© S.
parviflora), S. denticulata Ldb. and S. amurensis Turcz. distinguishing
it from them by larger capitula and red florets. Among the type material
is a sheet with the label (written in the hand of C.A. Meyer) that says:
“Saussurea Stubendorfiana m. Semina misit Dr. Stubendorf. Cult. in
h[orto] b.[ot.] Petropol. 1850.” The plant on this sheet differs from the
authentic wild plants of S. stubendorffii Herd. by more coriaceous leaves,
with more dense, snow-white, almost tomentose indumentum beneath,
relatively small capitula, weakly pubescent involucral bracts and one-
headed peduncles of the individual corymbs, and it is close to S.
sukaczevii Lipsch. It appears that S. stubendorffii Herd. is an east Siberian
and Yakutian race of S. alpina, also entering Altai, in habit often
convergent with S. serrata and S. amurensis; many of the floristic
taxonomists of Siberia wrongly identified it sometimes as S. serrata,
sometimes as S. amurensis. It differs sharply from S. parviflora (= S.
serrata) by the pubescence on the lower leaf surface, larger capitula,
and the absence of a distinct wing on the stem. It is more difficult to
distinguish S. stubendorffii from S. amurensis, especially its Trans-Baikal
forms, which have somewhat lost the “purity” of the species. S.
stubendorffii, in comparison with S. amurensis, has softer leaves that
are arachnoid hairy-flocculose on the lower side (and not white-
tomentose), a stem with less distinct wings from decurrent leaves, larger
capitula, and involucral bracts (especially inner) less villous-pubescent.
Among the materials referred by me to S. stubendorffii, these differ
from the typical specimens: 1) specimens from the Irkut River Valley
(Torskaya and Tunkinskaya ravines, V.I. Smirnov, 1931), characterized
by cupuliform capitula, in a more compact terminal corymb, densely
villous from ferruginous involucre hairs, outer involucral bracts narrower
and more strongly acuminate, and also a deeper violet. Evidently, it is
a separate race (S. turczaninovii Lipsch. in herbario), but the material at
my disposal is inadequate for a final conclusion about the specific status
of this entity; and 2) specimens from the Upper Angara Basin (vicinity
of village Verkhne-Angarsk, V.N. Sukaczev and G.I. Poplavskaya, 1912,

497

491

No. 1250, 4 sheets; N.V. Shipczinsky, 1912, No. 439) are distinguished
in type by stiffer leaves, entire and not toothed leaves (basal and middle
cauline leaves sessile or subsessile), and reddish stems (S. angarensis
Lipsch. in herbario). Specimens intermediate between the two species,
including a set of characters of both parents, have been collected in the
Yenisei region, in placés of contact between S. stubendorffii Herd.
and typical S. parviflora (Poir.) DC.; these are undoubtedly natural
hybrids.

86. S. ambigua Kryl. ex Serg. in Sistemat. Zam. Gerb. Tomsk.
Univ. 2 (1941) 2; Kryl. Fl. Zap. Sib. XI, 2920.—S. frigida Fisch. in
herb. and ex Herd. in Bull. Soc. Nat. Mosc. XLI, 3, 37, pl. tschuiens.—
S. alpina auct. fl. Sib. p. min. p.

Perennial. Rhizomes obliquely ascending or vertical; stem erect,
solitary, sometimes 2.5—3 cm high (var. pumila Lipsch.), or much taller
(4-20(25) cm high), almost glabrous or flocculose-pubescent in short-
stemmed plants. Leaves pale green, almost glabrous on both sides (var.
subglabra Serg. op. cit. (1941) 2), sometimes grayish arachnoid-hairy
beneath due to more or less thin tomentum of intertwined hairs, usually
with scattered (especially beneath) lustrous, sessile, yellowish glandular
hairs; basal and lower cauline leaves oblong-ovate, short-petiolate,
petioles one-eighth to half as long as lamina and together with it 5-13
cm long and 1—2.5(3) cm wide, with more or less frequent callose teeth
and short cilia in some, especially in short-stemmed forms; middle leaves
ovate-oblong, oblong or lanceolate, acuminate or less often subobtusely

- acuminate, sessile, nondecurrent or slightly decurrent on stem in

acaulescent specimens. Capitula 4-12, aggregated in compact-terminal
corymbs; less often with one or two (three) lateral corymbs besides
terminal corymb, arising from axils of upper leaves (var. ramosa Lipsch.);
sometimes several (or one) linear-lanceolate bracteal leaves borne at
base of corymb. narrower than outer involucral bracts. Involucre
imbricate, 5-8 mm wide, pale in type specimen, less often violet; outer
involucral bracts ovate or oblong-ovate, acuminate, pubescent; inner
bracts oblong, subobtuse, usually strongly villous-pubescent with white
deflexed hairs; peduncles, and sometimes also involucral bracts, bearing
solitary, lustrous, sessile, glandular hairs. Receptacle densely squamose;
scales linear-subulate, lustrous, about 4 mm long; florets dark pink,
turning red when dry, about 1.1-1.2 cm long, narrow part of corolla
tube about 5-6 mm long, as long as broad part with lobes of limb;
anther appendages hairy. Achenes 3-4 mm long, brownish or light-
colored, with dark brown spots and solitary glandular hairs; pappus dense,
white, double, outer pappus bristles less numerous, fragile, small, serrate,
inner hairs up to 10-11 mm long, plumose. Flowering VIJ-IX.

498

492

Stony and saline steppes, saline ‘meadows.—Western Siberia: Altai
(southeastern part), Angara-Sayans (Tuva Republic).—General
distribution: Mongolia. Described from the upper reaches of the Chuya
River. Type in Leningrad.

Note. It differs from the Siberian forms of S. alpina by a shorter
stem, oblong-ovate, outer involucral bracts one-third to half as long as
the inner, covered with more or less numerous, sessile, yellowish
glandular hairs below, and by more dense pappus.

87. S. saxosa Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVI (1954) 457.

Perennial. Plants woody; rhizome woody, blackish-brown, deeply
penetrating stones, longitudinally fibrously divided, many-headed at apex,
imbricately covered with remains of previous year’s leaves; sometimes
lateral caudices terminating in rosettes of leaves; stems 4—15(20) cm
high, slightly ascending at base, sulcate, arachnoid-hairy, densely leafy
up to apex. Leaves hard, oblong or ovate-oblong, grayish-green on both
sides, densely arachnoid-hairy (more strongly beneath) and with sessile
glandular hairs, with sparse cartilaginous small teeth, often weakly
involute, sometimes almost entire, with prominent midrib beneath,
continued into petiole in lower leaves, and with less prominent lateral
veins; leaf base weakly cuneate; all leaves acuminate, less often obtusely
acuminate; basal leaves distinctly petiolate, petioles weakly winged, 5—
7(8) cm long (including petioles, about 2-4 cm long) and (1)1.5—2 cm
wide; middle leaves sessile or subsessile, usually weakly decurrent on
stem like basal leaves; uppermost leaves reduced, lanceolate or linear,
usually approximate with inflorescence, about 2—2.5 cm long and 1.5
mm wide. Capitula on about 5 mm long peduncles, terminal on stem in
dense corymb. Involucre imbricate, terminal on stem in dense corymb.
Involucre imbricate, about 1—1.1 cm long, 0.7—1 cm wide; involucral
bracts pale, sometimes dark-colored, pubescent; one or two lanceolate,
acuminate, bracteal leaves below capitulum, as long as outer involucral
bracts, sometimes almost as long as inner involucral bracts; outer bracts
ovate, acuminate; inner bracts longer, linear-lanceolate or lanceolate,
acuminate, villous in upper half due to long, white, semideflexed hairs,
sometimes forming tuft at apex of bracts. Receptacle not densely
squamose, scales linearly subulate, lustrous, unequal, small in center,
about 1-2 mm long, larger along margin, 3-4 mm long; florets about
1.1-1.2 cm long, narrow part of tube about 5—6 mm long, as long or
almost as long as broad part with lobes of limb; anther appendages

hairy. Pappus double, outer pappus bristles small, about 2-3 mm long,

serrate-scabrous, inner bristles long-plumose, about 8-10 mm long;
achenes mature, about 5 mm long, dark brown, with white ribs, shining,
covered with glandular hairs. Flowering VII—beginning of IX.

493

RNs
SS
FSS

°
SF

, U3.

499 Plate XXXVII.

1—Saussurea salsa (Pall.) Spreng.; 2—S. dahurica Adams.

500

494

Stony slopes, rocks (in the crevices), screes.—Central Asia: Pamiro-
Alai (Zeravshan and Turkestan ranges). Endemic. Described from the
Zeravshan Valley. Type in Leningrad.

Note. A species distinctly demarcated from the polymorphic species
S. alpina, closest to the Mongolian species S. klementzii Lipsch.

88. S. angustifolia (Willd.) DC. in Ann. Mus. Paris, XVI (1810)
200; DC. Prodr. VI, 535; Ldb. Fl. Ross. II, 2, 668; Hult. Fl. of Alaska
and Yukon, X, 1625; Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVI, 451; Polunin, Circumpolar Arctic Flora (1959) 453 excl.
S. viscida Hult.—Serratula angustifolia Willd. Sp. pl. Ill, 3 (1800)
1642.—Saussurea borealis Fisch. ex Herd. in Bull. Soc. Nat. Mosc.
XLI, 3 (1868) 38 (nomen) and in Herb.—Ie.: Polunin, op. cit. (1959)
453.

Perennial. Small woody shrubs; stems 3.5—10(12) cm high, simple,
sulcate, somewhat pubescent or glabrous. Leaves entire, usually more
or less revolute, glabrous on both sides, sometimes sparsely woolly
beneath; basal and cauline leaves linear or oblong-linear, uppermost
leaves linear, approximate with inflorescence; all leaves narrow, in type
specimen 2-3 mm wide, subobtuse. Capitula small, about 1 cm long,
on distinct peduncles in terminal, relatively compact corymbs. Involucre
three-rowed, pubescent; outer involucral bracts ovate, subobtuse or obtuse;
inner bracts lanceolate or oblong-lanceolate, longer than outer bracts,
acuminate, both pubescent with long, often squarrose, with hairs on
dorsal surface, along margins and at apex. Receptacle densely squamose,
scales linearly-subulate, lustrous, up to 5 mm long; florets about 1—1.1
cm long, narrow part of tube about 5 mm long, as long as broad part
with lobes of limb; anther appendages ciliate. Pappus double, outer
pappus bristles small, fragile, serrate-scabrous, inner bristles plumose,
about 9 mm long; achenes glabrous. Flowering VII—VIII.

Mossy tundras, grassy swales.—Arctic: Chukotka (islands of Bering
Strait, for example, Arakamchechen); Gulf of St. Lawrence; near the
Chukotsk hot springs). General distribution: North America (Alaska,
Canada).

Note. The Arctic race of S. alpina is quite close to S. viscida Hult.
(see below), included under S. angustifolia by some authors (N. Polunin).
S. angustifolia DC. differs from S. viscida Hult. by having glabrous or
flocculose and not glandular-hairy stem and leaves; basal leaves linear
with revolute margins (elliptical, lanceolate or broadly lanceolate in S.
viscida). The earlier reports of the occurrence of this North American
species in eastern Siberia and in the vicinity of Ayan are mistaken. The
plants from Ayan and eastern Siberia identified-as S. alpina var.
angustifolia Rgl., are not identical with the true S. angustifolia, distributed

501

495

only in the extreme Northeast of the USSR (cf. above), and I have
described them as a separate race—S. pseudoangustifolia Lipsch.

89. S. viscida Hult. Fl. of Alaska and Yukon, X (1950) 1627;
Polunin, Circumpolar Arctic Flora, 453 p. p. sub nom: S. angustifolia
agg.—sS. alpina var. ledebouri Gray, Synopt. Fl. N. Amer. I, 2 (1886)
397 quoad pl. ex Alask—S. alpina f. angustifolia Kjellman in Vega
Exp. Vet. Iaktt. 2 (1883) 14, excl. synon.—Ic.: Hult. op. cit. (1950)
figsiGre:

Perennial. Dwarfish plants, 5-15 cm high; stem simple, weakly
arachnoid-hairy and covered with viscid hairs. Lower leaves elliptical-
lanceolate, obtuse or acute, entire or indistinctly toothed, sessile or with
extremely short petiole, covered with scattered, viscid, many-celled hairs
on both sides, margins ciliate and with dense viscid hairs; upper leaves
like lower but narrower. Capitula on short peduncles, aggregated in
compact terminal corymbs. Involucre two- or three-rowed; outer
involucral bracts ovate, acuminate or subobtuse, more or less densely
pubescent; inner bracts longer, lanceolate, acuminate, densely pubescent
with white semi-erect hairs. Receptacle squamose; scales linear-subulate,
lustrous, unequal, up to 6 mm long; florets dark pink(?); narrow part of
tube about 5 mm long, as long or almost as long as broad part with
lobes of limb; anther appendages ciliate. Pappus double, outer pappus
bristles scarcely noticeable, small, fragile, inner bristles about 9 mm
long, plumose; achenes glabrous, ribbed. Flowering VII—VIII.

Tundra.—Arctic: Chukotka (village of Uelen, Naukan). General
distribution: Alaska. Described from St. Lawrence Island. Type in
Stockholm.

Note. Besides S. angustifolia, it is also close to S. tilesii; see the
note to the latter about specimens from the Wrangel Island. Viscid hairs
are not always present on the stem and leaves.

90. S. pseudoangustifolia Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVI (1954) 452.—S. alpina [var.] angustifolia Rgl. in Mém.
Soc. Nat. Mosc. XI (1859) 107.

Perennial. Root collar covered with remains of previous year’s leaves.
Stem erect, solitary, 10—-15(20) cm high, sulcate, arachnoid-hairy,
sometimes glabrescent, usually with floriferous branches in upper part.
Leaves lanceolate or lanceolate-linear, weakly but more or less distinctly
toothed, often weakly involute, arachnoid-hairy beneath and with
scattered, small, lustrous glandular hairs; basal and lower cauline leaves
1-1.3 cm wide, long-petiolate, sometimes somewhat cuneate at base;
upper leaves 3-5 mm wide, linear in type specimen, less often, linear-
lanceolate, sessile or subsessile, not decurrent on stem. Capitula solitary

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496

or less numerous (2-5), in latter case lateral capitula on more or less
long peduncles of various length; capitula usually up to 2—2.3 cm long
and 1—1.5 cm wide, central capitulum of corymb usually larger than
laterals. Involucre four- or five-rowed, imbricate, with one to three linear
bracteal leaves at its base; outer involucral bracts ovate or ovate-
triangular, acuminate, pubescent, inner bracts oblong-lanceolate,
acuminate, more or less villous. Receptacle squamose, scales linear-
subulate, equal, 1.1 cm long, narrow part of tube about 5 mm long, as
long or almost as long as broad part together with lobes of limb. Pappus
double, outer pappus bristles few, fragile, serrate-scabrous, unequal, up
to 4-5 mm long, inner bristles plumose, about | cm long; achenes

’ glabrous. Flowering VII-—VIII.

Open forests, thickets of Japanese stone pine meadows, marshes on
alpine summits.—Far East: Okhotsk; Eastern Siberia: Lena-Kolyma
(eastern part). Endemic. Described from the vicinity of Ayan. Type in
Leningrad.

Note. A separate eastern Siberian race of S. angustifolia DC., for
which it was incorrectly taken to be by a number of authors. In habit it
also approaches S. tschuktschorum Lipsch., from which it differs by
having a squamose receptacle. The old reports (Willdenow, Pallas, etc.)
that the range of S. angustifolia is in eastern Siberia are mistaken.

Series 18. Kuschakevicziae Lipsch.—Plants almost stemless, small,
sometimes with distinctly developed but short stem. Basal leaves rotund,
ovate or elliptical, usually in rosette, surrounding inflorescence in
stemless forms and as long as inflorescence. Capitula aggregated in large
or small number in dense terminal corymb. Receptacle squamose, with
small; sometimes corymb. Receptacle squamose, with small, sometimes
fragile scales. Alpine plants. .

Note. S. ovata Benth. and S. caprifolia Iljin and Zapr.
phylogenetically gravitate to this series. However, the latter species differs
so much by its smooth receptacle and larger solitary or few capitula
that it is considered later under section Depressae Clarke. The differences
between the species included in the series are difficult to delineate, but
are well expressed in overall appearance.

_ 91. S. kuschakeviczii Winkl. in Tr. Peterb. Bot. Sada, XI, 1 (1889)
170.—S. alpina var. kuschakeviczi Winkl. ex O. Fedtsch. Ibid., XXI, 3

(1903) 358.—S. kuschakeviczii Winkl. var. caulescens O. Fedtsch. op.

cit., XXVIII, (1908) 170.

Perennial. Dwarfish, small, stemless plants or with short, 2—5(8) cm
high, sometimes flattened, flocculose, less often almost glabrous stem |
(var. caulescens O. Fedtsch.). Leaves, especially beneath, arachnoid-

503

497

hairy pubescent, sometimes glabrescent, toothed, teeth cartilaginous at
apex; basal leaves more or less rosulate, surrounding inflorescence in
stemless forms, as long as inflorescence or larger, variable in shape and
size from ovate to elliptical and even oblong-lanceolate; all leaves short-
petiolate, petioles usually much shorter than. lamina, together 1.5—5(7)
cm long, 1—-1.5(2) cm wide; cauline leaves distinctly developed only in
forms with stem (var. caulescens O. Fedtsch.), like basal leaves sessile;
upper cauline leaves reduced, 3-5 mm wide, lanceolate, approximate
with inflorescence. Capitula 3-20, numerous in type specimen, aggregated
in dense terminal corymb. Involucre narrow-campanulate, indistinctly
imbricate, tomentose or glabrous, subsequently becoming hard (always
?); outer and middle involucral bracts ovate, acuminate; inner bracts
lanceolate or oblong-lanceolate, acuminate, both usually light colored,
less often pigmented. Receptacle squamose, scales few, linearly subulate,
lustrous, unequal, fragile; florets 1.1-1.2 cm long, narrow part of tube
about 5—(6) mm long, as long or almost as long as broad part with
lobes of limb; anther appendages hairy. Pappus double, outer pappus
bristles up to 3 mm long, unequal, serrate, rather numerous, easily
shedding, inner bristles long-plumose, 11-12 mm long; achenes 3-4 mm
long, dark brown, glabrous. Flowering VIIIJ-IX.

Moraines, alpine grassy areas, pebbled areas, meadows above the
upper boundary of forests, rocks.—Soviet Central Asia: Pamiro-Alai
(Pamir, Alai and Trans-Alai ranges); Tien Shan; Dzhungaria-Tarbagatai
(Dzhungarian Alatau). Endemic. Described from Kyzyl-Art. Type in
Leningrad.

Note. A high-mountain, Soviet Central Asian race of S. alpina,
differing from it by growing close to the ground, having basal leaves
that are rosette-like, surrounding the inflorescence, and having a less
distinctly developed, imbricate involucre. C. Winkler reports a smooth
receptacle in the original description of S. kuschakeviczii. Study of the
authentic material revealed the presence of a few scales on the receptacle.
The plants collected subsequently from Kyzyl-Art also have squamose
receptacles. S. kuschakeviczii Winkl. is closest to S. caprifolia Ijin and
Zapr., from which it differs by having a squamose receptacle, larger
number of smaller capitula, and less rotund toothed leaves. Earlier known
only from Pamir and the Kyzyl-Art Pass, it subsequently was found in
other regions of Alai, in the Central Tien Shan and Dzhungarian Alatau.

92. S. ovata Benth. in Henderson and Hume, Lahore to Yarkand
(1873)325.—Ie.: op. cit. (1873); 326, tab. colorata.

Perennial. Rhizome developing stems and sometimes rosettes of
leaves; stems 2.5-10 cm long, sometimes 15—25 cm in individual
specimens, glabrous in type specimens or less often more or less

504

498

pubescent, sulcate, usually with fewer leaves. Leaves fleshy, glabrous
on both sides (arachnoid-hairy in dwarf specimens) and with scattered,
small, lustrous glandular hairs (visible under lens), ovate, toothed or
even shallow sinuate-toothed, teeth cartilaginous at apex; basal and lower
cauline leaves distinctly petiolate, 4-10 cm long (including petiole) and
2-4.5 cm wide, truncate at base or indistinctly cuneate, usually with
cusp, petiole narrow-winged; upper leaves significantly reduced, sessile,
approximate with inflorescence. Capitula narrow-cupuliform, on short
peduncles, three to many, aggregated in dense capitate corymbs at stem
apex. Involucre imbricate, 8-10 mm long and 5-8 mm wide, pubescent
with stiff hairs; outer involucral bracts broader, ovate, with short apical
cusp or roundish; inner bracts narrower, subobtuse and often glochidiate
at apex. Receptacle covered with smooth short scales, scales about 1—2
mm long; florets dark pink, narrow part of tube shorter than its broad
part together with lobes of limb; anther appendages ciliate. Pappus
double; outer pappus-bristles less numerous, serrate-scabrous, fragile,
small, 2-3 mm long, inner bristles long-plumose, about 10 mm long;
achenes 3 mm long, dark brown, glabrous. Flowering VII-—VIII.

Stony and clayey slopes; alpine meadows.—Soviet Central Asia:
Pamiro-Alai (near Irkeshtam). General distribution: Kashgaria. Described
from the Yarkenda Region. Type in Kew(?).

Note. Because of the limited material, I am referring the specimens
from the Talass Alatau (“stony screes in the upper reaches of the
Dzhebogly-Su River, 3500 m above msl, 4, VIII, 1931, No. 934”. N.V.
Pavlov) to S. ovata Benth. They are characterized by stems 3-5 cm
high, arachnoid-hairy stem and leaves, smaller capitula (involucre 5—7
mm long) with slightly exserted florets (possibly, this is explained by
the fact that the florets were damaged by frost). Evidently, this is a
separate species (S. talassica Lipsch. ined. in Herb. Inst. Bot. Acad. Sc.
URSS), but larger collections are required to resolve the question finally.

Bentham reported incorrectly in the original diagnosis of S. ovata
that the receptacle of this species is smooth; all unquestioned specimens
of S. ovata examined by me (from Kashgaria) show a squamose
receptacle, but the scales are 1—2 mm long and not dense.

Series 19. Salsae Lipsch.—Plants of saline and salty habitats,
somewhat bitter to taste. Stem covered with hardened remains of previous
year’s leaves at base. Leaves fleshy, sometimes lower leaves lyrate-
pinnatisect, with large terminal lobe almost sagittate at base and sinuate-
large toothed along margin, upper leaves often entire, sometimes deeply
pinnatisect; usually (but not always) decurrent. Capitula in corymbs,
sometimes numerous, aggregated in branched, lax, paniculate-corymbose
inflorescence, sometimes less numerous, terminal, dense. Outer involucral

505

499

bracts not herbaceous and not distinguished from inner bracts by
consistency. Anther appendages hairy. Pappus snow-white, soft.

Note. Besides S. salsa, I also include under this series S. turgaiensis
B. Fedtsch., S. daurica Adams, S. kaschagarica Rupr., and tentatively
S. pseudosalsa Lipsch. from Kashgaria, as well as S. faminziniana
Krassn., endemic to the USSR. The affinity of the latter species is not
clear to me.

93. S. salsa (Pall.) Spreng. Syst. veg. III (1826) 381; Ldb. FI. alt.
IV, 22 excl. syn. S. elongata DC. and Poir.; Boiss. Fl. or. III, 566; O.
and B. Fedtsch. Perech. Rast. Turk. IV, 232 excl. syn. plur.; Grossh. FI.
Kavk. IV, 168; Fl. Yugo-Vost. VI, 386; Kryl. Fl. Zap. Sib. XI, 2912
excl. var. laciniosa (Kryl.) Serg.; Popov, Fl. Sr. Sib. II, 757.—S. salsa

var. fastigiata Winkl. in O. and B. Fedtsch. op. cit. (1911) 232, nomen.—

Serratula salsa Pall. Reise, I, Anhang (1771) 502 sub nomine “Planta
salsa ambigui generis”; nomen specificum conf. Pall. Reise, III (1776)
607, 635; MB. FI. taur.-cauc. II, 266.—S. salina Pall. Reise, III (1776)
314.—-Heterotrichum salsum MB. FI. taur.-cauc. III (1819) 551.—
Saussurea crassifolia DC. in Ann. Mus. Paris, X (1810) 201; Ej. Prodr.
VI, 533; Spreng. op. cit. 380; Ldb. Fl. Ross. II, 2, 665, excl. 8. papposa
Turcz.; Bge. in Mém. Acad. Pétersb. sav. étrang. VII, 353 excl. var. B.
muricata Bge.; Kryl. Fl. Alt. III, 703 excl. f. laciniosa Kryl.
Perennial. Root collar more or less densely covered with remains
of previous year’s leaves. Stems 15-50 cm high, solitary or several,
sulcate, pubescent or almost glabrous, branched above or from middle,
distinctly winged (wings of variable width, entire or toothed) due to
decurrent leaves,- sometimes weakly winged, often wingless. Leaves
somewhat thick, fleshy, more or less scabrous due to short stiff hairs or
almost smooth, glabrous, covered with numerous punctate glandular hairs
beneath, more weakly noticeable above; all leaves fairly polymorphic;
especially in respect of serration; basal and lower cauline leaves larger,
petiolate, broadened at base in sheaths, together with petioles 5—35(!)
cm long, 2-12 cm wide, lyrate-pinnatisect, with large, usually sagittate
at base, and sometimes boldly sinuate-toothed, sometimes entire terminal
lobe; lateral lobes numerous, triangular, usually entire, retrorse, gradually
reduced toward base; cauline leaves oblong, oblong-linear or lanceolate,
entire or with less numerous teeth along margin, sessile, usually
decurrent; uppermost leaves narrow and weakly decurrent. Capitula with
florets 1—-1.2 cm long, numerous, forming lax corymbose-paniculate
inflorescence. Involucre imbricate, 4-5 mm wide, glabrous or arachnoid-
hairy, involucral bracts obtuse; outer bracts ovate, inner oblong, pink or
red colored. Receptacle densely squamous, scales uneven, sometimes
44.5 mm long, linearly subulate, lustrous; florets pink, 1-1.1 cm long

506

500

with narrow part of tube, almost as long as or somewhat shorter than
broad part with lobes of limb; anther appendages hairy. Pappus double,
outer pappus bristles unequal, serrate, up to 3 mm long, fragile, inner
bristles plumose, about 9 mm long; achenes smooth, compressed.
Flowering VI-IX. (Plate XXXVII, Fig. 1.)

Saline lands, alkaline steppes and meadows, reed thickets.—
European Part: Black Sea Region (Novomoskovsk and Melitopol
regions), Crimea (Perekop), ?7Lower Don, Lower Volga, Trans-Volga
Region, Volga-Kama (southeastern); Caucasus: Cis-Caucasia, southern
Transcaucasia? Dagestan (according to Grossheim); Western Siberia:
Upper Tobol, Ob’ (southern part), Irtysh, Altai; Eastern Siberia: Angara-
Sayans; Soviet Central Asia: Aralo-Caspian Region, Balkhash Region,
Kara-Kum, Kyzyl-Kum, Amu-Darya, Syr-Darya, Tien Shan, Pamiro-Alai,
Dzhungaria-Tarbagatai. General distribution: Iran, Dzhungaria-Kashgaria,
Mongolia. The type specimen originated from saline lands near Samara
and around the salt mines near Lake Iletskoe (Pallas). Type in Leningrad.

Note. An extremely polymorphic species which was first reported
by Ledebour (F7. alt. IV, 23). It varies in the form of leaves (their size,
serration), presence or absence of wings on the stem formed by decurrent
leaves, scabrous or smooth stem and leaves, pubescence of the involucral
bracts, form of the inflorescence (few terminal corymbs or numerous
lax corymbs, forming corymbose panicle), etc. I consider the following
taxa, phylogenetically close to S. salsa, as independent species: S. daurica
Adams, S. turgaiensis B. Fedtsch. and S. kaschgarica Rupr. Populations
with white florets (albinos) are found in S. salsa; this is f. albiflora
Lipsch. (vicinity of the Aul Health Resort in the Altai, V.I. Vereshchagin,
1921). The taxonomic status of S. crassifolia B. muricata Bge. |.c. 1s
not clear to me. It was described from Zeravshan (its type specimen is
not found in the Herbarium of the Botanical Institute). This variety is
close, if not identical, to var. aspera Serg. (Kryl. Fl. Zap. Sib., XI,
2912). Specimens resembling S. salsa, but differing from it in several
characters, have been collected from the Kurgan-Tyubensk Region
(“Clayey Plain between Kurgantyube and Lyagman,” 2.VIII.1913, No.
659, B.A. Fedchenko; same place, 15.VIII.1913, No. 1040, I.
Bornmiiller). Unfortunately, these specimens were collected at an
immature stage, which does not allow us to come to a definitive opinion
about their taxonomic rank. They are characterized by a less leafy,
glabrous, wingless stem up to 70 cm high; elongated, strongly pinnatisect
lower leaves, some lower lobes of which are bipinnatisect, curved, and

' upward-pointed, and the upper lobe, as in true S. salsa, is larger and

sagittate at the base. Apparently, it is a separate race (or ecological
variety?). Pamir specimens differ slightly from the typical ones and are
split off by me into the separate var. pamirica Lipsch. var. nova: stems

507

501 |

15—20(25) cm high, few, simple, less often branched, usually distinctly
winged from decurrent leaves; leaves elongate, runcinate or sinuate-
toothed, with indistinctly developed upper lobe; corymbs of capitula
compact; inner involucral bracts acuminate (type: toward north of
Murgab, 1954, No. 590, P. Polyakov). In the Zaisan Region, very unusual
forms of S. salsa are found, which are intermediate between it and S.
laciniata Ldb., a species with appendages on the involucral bracts and,
therefore, included under subgenus Theodorea (Cass.) Lipsch.

94. x S. paradoxa Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 378.—S. salsa (Pall.) Spreng. x-S. lanciniata
Ldb.

Perennial. Root collar covered with remains of sheaths of previous
year’s leaves; stems solitary or few, 25-35 cm high, sulcate, usually
weakly winged due to decurrent leaves, slightly pubescent with short
hairs, pubescence easily rubbed off, and then stems almost glabrous.
Basal and lower cauline leaves very similar to leaves of S. salsa, long-
petiolate, with petiole up to 18 cm long, up to 5 cm wide, broadened at
base into short sheath; lamina somewhat thick, more or less scabrous
due to short, hard, appressed hairs, ovate, elliptical or oblong, crenate-
toothed, especially in lower part, teeth terminating in callose cusp; middle
and upper leaves reduced, entire. Capitula in corymbs, terminal on stem
and less numerous lateral branches, like capitula of S. laciniata. Involucre
up to 10 cm long, middle and inner involucral bracts broadened into
membranous pinkish apical appendage. Receptacle densely squamous,
scales linearly subulate, white, lustrous, unequal; florets pink, about 1.2
cm long including narrow part of tube, almost as long as broad part
together with lobes of limb. Pappus white, double, outer pappus bristles
short, fragile, inner bristles plumose, about 8-9 mm long, slightly exserted
from involucre. Flowering VII-IX.

Saline lands, alkaline meadows.—Western Siberia: Irtysh (Zaisan
District). General distribution: Mongolia. Described from the Takyrka
River. Type in Leningrad.

Note. As can be seen, the plant has inherited its vegetative
morphological characters (stem, leaves) from S. salsa and its capitulum
features (presence of membranous appendages on the involucral bracts)
from S. Jaciniata and is, therefore, undoubtedly, a natural hybrid between
them. This hybrid is also known from Mongolia (Ladygin, No. 410,
Island on Lake Kobdo).

95. S. daurica Adams in Nouv. Mém. Soc. Nat. Mosc. III (1834)
251; Lipschitz in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX],
378.—Serratula davurica Adams errore ex Ldb. FI. Ross. II, 2 (1845-

502

1846) 666.—Saussurea papposa ‘Turcz. in DC. Prodr. VI (1837) 534;
Turcz. Fl. baic.-dahur. II, 1, 108; Herd. in Bull. Soc. Nat. Mosc. XLI,
3, 10; Kom. in Tr. Peterb. Bot. Sada, XXV, 2, 742; Serg. in Sistemat.
Zam. Gerb. Tomsk. Univ. 2 (1941) 4-5 (descriptio emend.); Krylov, Fl.
Zap. Sib. XI, 2913; Popov, Fl. Sr. Sib. I], 757.—S. crassifolia B. papposa
Ldb. Fl. Ross. II, 2 (1845-1846) 666.—Exs.: GRF No. 1330.

Perennial. Root slender, one- or less often two- or three-headed.
Stems solitary, sometimes two or three, slender in type specimen, (5)10—-
15(20) cm high, erect, usually simple, less often branched, almost
glabrous or sparsely pubescent, not winged. Leaves somewhat thick,
almost smooth or weakly scabrous due to short stiff hairs, lanceolate or
oblong; basal and lower cauline leaves petiolate, with petiole 3-9 cm
long, 1-3 cm wide, terminal lobe often indistinct, if fully developed,
then oblong, entire or shallow crenate-toothed, lateral lobes two or three
pairs, sometimes absent; cauline leaves two to five, sessile or short-
petiolate, semiamplexicaul, nondecurrent on stem, oblon, runcinate,
sometimes entire, 4-7 cm long and 0.6—1.5 cm wide; uppermost leaves
smaller, entire. Capitula in semicapitate or capitate corymbs. Involucre
10-12 mm long, 5-6 mm wide; outer involucral bracts ovate, inner
oblong, acuminate, almost glabrous or puberulent along margin, often
reddish. Receptacle squamous, scales linear-subulate, lustrous; florets
pink, up to 1.4 cm long, narrow part of tube about 8 mm, broad part
with lobes of limbs about 6 mm long. Pappus snow-white, outer pappus
bristles short, serrate, fragile, inner bristles plumose, about 11-12 mm
long, strongly exserted from involucre at fruiting; achenes with narrow
top edge (corona). Flowering VII-IX. (Plate XXXVII, Fig. 2.)

Saline lands, desert steppes, saline meadows.—Western Siberia: Altai
(southeastern part, Chuya steppe), Eastern Siberia: Angara-Sayans
(Khakassia, Tuva); Dauria. General distribution: Mongolia, China.
Described from Dauria; Adams’ type evidently is lost; the type of S.
papposa Turcz. originates from the Selenginsk saline lands and is
preserved in Leningrad.

Note. Adams’ name has priority. He described this species in the
genus Saussurea and not in the genus Serratula. Ledebour erroneously
cites the name “Serratula davurica Adams.”

Popov (op. cit. 1959) reported that forms of S. salsa transitional to
S. papposa (= S. daurica) are found in the Selenginsk Dauria; I did not
see them. S. salsa is the well defined Daurian-Mongolian race. It can
be distinguished easily from the latter by the lower stature, aggregated
corymbs of capitula, with a dense, snow-white pappus, strongly exserted
from the involucre at fruiting, giving a globose or hemispherical
appearance to the inflorescence, smaller, nondecurrent leaves, acuminate
inner involucral bracts, and achenes with a narrow border-corona.

503

96. S. turgaiensis B. Fedtsch. in Fedde, Repert. VIII (1910) 497;
Fedtsch. in Tr. Pochv.-Bot. Eksp. Pereselench. Upravl. Ch. II, Bot. Issl.
1908, 7, 242; O. and B. Fedtsch. in Perech. Rast. Turk. IV, 232; FI.
Yugo-Vost. VI, 386.—S. salsa var. pinnatifida Winkl. in O. and B.
Fedtsch. op. cit. 232 nomen and in herb.—S. salsa var. bipinnatifida
Rgl. and Schm. in O. and B. Fedtsch. op. cit. 232 nomen and in herb.—
S. crassifolia f. laciniosa Kryl. Fl. Alt. HI (1904) 703.—S. salsa var.
laciniosa (Kryl.) Serg. in Kryl. Fl. Zap. Sib. XI (1949) 2912.—Ie.:
Fedtsch. Rast. Turk. (1915) 751.

Perennial. Roots strongly thickened, woody, with many caudices
above, covered with remains of hardened sheaths of previous year’s
leaves. Stems more or less numerous, (15)20—30(40) cm high, branched
from middle or above, sulcate, sometimes angular-sulcate, pubescent,
wingless, sparsely leafy. Leaves more or less covered with stiff hairs,
numerous, mainly crowded in lower part of plant, pinnatisect, short-
petiolate, broadened at base in sheath; lamina of basal and lower cauline
leaves oblong, pinnatisect or even bipinnatisect, lower lobes ovate,
divided in linear lobules, upper lobes linear, almost undivided; cauline
leaves reduced, pinnatisect, on very short petioles or sessile; uppermost
leaves small, sometimes with small number of lateral teeth, less often
almost entire. Capitula numerous, in corymbs, aggregated in lax
paniculate inflorescene. Involucre imbricate, about 10 mm long,
involucral bracts usually reddish; outer bracts smaller, ovate, inner
oblong, both puberulent especially along margin (often glabrescent),
subobtuse. Receptacle squamose, scales linear-subulate, lustrous; florets
pink, about 1.2 cm long, narrow part of tube about 7 mm long, broad
part with lobes of limb about 5 mm long. Pappus double,
outer pappus bristles short, serrate, fragile, inner bristles plumose,
somewhat exserted from involucre; achenes glabrous. Flowering VII-—
VIII. (Plate XXXVII, Fig. 1.)

Saline lands, saline meadows, thickets of chee grass (Lasiagrostis
splendens), limestone outcrops.—European Part: Trans-Volga Region
(southeastern part); Western Siberia: Upper Tobol, Irtysh, Altai; Soviet
Central Asia: Aralo-Caspian, Balkhash Region, Kyzyl-Kum, Tien Shan
(?), Pamiro-Alai (Alai Valley, form transitional to S. salsa). Described
from former Turgai District (border of Naurzum-Karagai forest and the
former Akmolinsk District). Type preserved in Leningrad.

Note. Albino populations (f. albiflora Lipsch.) are also found. It
differs from S. salsa (Pall.) Spreng. by pinnatisect and more strongly
scabrous leaves owing to short stiff hairs. A few specimens transitional
to S. salsa are found in which the leaves are indistinctly pinnatisect.

97. S. kaschgarica Rupr. in Mém. Acad. Sc. Pétersb. VII sér. XIV,
No. 4 (1869) 54; O. and B. Fedtsch. Perech. Rast. Turk. IV, 233.

510

504

Perennial. Root collar covered: with more or less shining or smooth,
hardened, dark brown sheaths of basal leaves. Stems several, 20—25 cm
high, more or less ascending at base, sulcate, weakly pubescent, sparsely
branched, wingless. Leaves pinnatifid or pinnatisect, scabrous due to
numerous short and stiff hairs; basal leaves petiolate, up to 15 cm long
with petioles, petioles broadened in sheaths, lobes of lamina acute, long
or short-toothed; cauline leaves reduced, short-petiolate or almost sessile,
not decurrent. Capitula in dense corymbs, terminal on stem and branches,
if branched. Involucre deep purple, about 12 mm long and 8-10 mm
wide, imbricate, four-rowed; outer involucral bracts ovate; subobtuse;
inner bracts oblong or oblong-lanceolate, with somewhat subacute apex,
sparsely pubescent with stiff hairs on dorsal surface, along margin, and
apex, later glabrescent. Receptacle squamose, scales linearly-subulate,
lustrous, unequal; florets pink, about 1.4 cm long, narrow part of tube
almost as long as or slightly shorter than broad part with lobes of limb;
anther appendages hairy. Pappus double; outer pappus bristles unequal,
3—5 mm long, stiff, serrate, persistant, inner bristles about 10-11 mm
long, plumose; achenes about 5 mm long, glabrous. Flowering VIII.

Saline lands, outcrops of varicolored rocks.—Soviet Central Asia:
Tien Shan. Endemic. Described from the Suukta Valley. Type in
Leningrad.

Note. A species of clear phylogenetic affinity to S. salsa, from which
it was distinguished by Ruprecht on the basis of scabrous, weakly
decurrent leaves, larger and wider capitula, compactly aggregated on
the tips of unbranched or weakly branched stems, and a receptacle that
is not squamose, but narrow-setose. It was incorrectly stated in the
diagnosis of Ruprecht that the receptacular bristles are “sometimes
plumose”; the receptacular bristles (scales) in Saussurea are never
plumose. Evidently, Ruprecht considered the deciduous outer pappus
bristles remaining on the receptacle as plumose receptacular bristles.
The collections from central Tien Shan made since Osten-Sacken’s
collections, on the basis of which S. kaschgarica was described, are not
identical with the authentic specimens; they differ by having smaller
capitula, pale (and not purple) involucral bracts, and more scabrous
peduncles and branches of the inflorescence. But since these specimens
(Issyk-Kul Basin, No. 281, 1953, P.P. Polyakov; Naryn-Tau mountains,
1926, No. 1191) are quite close to S. kaschgarica Rupr., I include them
under the latter. The presence of a well-developed outer row of pappus
hairs that persist on the achene is a very characteristic feature of both
the authentic material of S. kaschgarica Rupr. and these similar
specimens.

98. S. faminziniana Krassn. in Bot. Zap. IJ, 1 (1887) 71; in Zap.
Russk. Geogr. Obshch. po Obshch. Geogr. XIX, 338; Lipschitz in Bot.

51

—

505

Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX, 387.—S. colorata Wink].
in Tr. Peterb. Bot. Sada, XI, 1 (1889) 167.

Perennial. Rhizome many-headed, forming turf; caudices covered
with remains of previous year’s leaves, partially fibrously divided; stem
(2)5—12 cm high, prostrate or ascending, pubescent or glabrescent, simple,
sometimes branched above. Leaves lanceolate, 2 to 17 cm long (including
petiole) and 0.3—1.2 wide, scabrous, arachnoid-hairy or less often almost
glabrous on both sides, large toothed, or pinnatidentate, less often almost
entire, teeth with small cartilaginous cusp; basal leaves gradually narrowed
in petiole, petiole almost as long as lamina or nearly half as long and
broadened at base in hard sheath; lower cauline leaves short-petiolate,
upper sessile, weakly amplexicaul. Capitula on short peduncles, in compact
corymbs, campanulate, about 1.2—1.5 cm long and up to 1.2 cm wide.
Involucre imbricate, light colored or dark, arachnoid-hairy, later sometimes
glabrescent; outer involucral bracts ovate, inner lanceolate, obtuse.
Receptacle glabrous, not squaamose sometimes with less numerous scales;
florets pale purple, about 1.3—-1.4 cm long, narrow part of tube about 6—
7 mm long, broad part with lobes of limb almost as long as narrow part;
anther appendages ciliate. Achenes about 4 mm long, glabrous, smooth,
with short apical corona; pappus double, snow-white; outer pappus bristles
short-plumose, persistent on achenes, up to 4-5 mm long, inner bristles
long-plumose, about 11 mm long. Flowering VIII.

Sands (saline), stony slopes, moraines, alkaline patches in alpine
steppe.—Soviet Central Asia: Pamiro-Alai (eastern Pamir), Tien Shan.
General distribution: Dzhungaria-Kashgaria (eastern Tien Shan). S.
faminziniana Krassn. was described from Sary-Dzhas (central Tien Shan).
Type in Leningrad. S. colorata Winkl. was described from Eastern Pamir
(Lake Kara-Kul). Type in Leningrad.

Note. As shown by my studies, S. faminziniana Krassn., for which
the author of the species reports the presence of receptacular scales, and
S. colorata Winkl., whose receptacle was described as smooth (without
scales), are identical, representing a single species. The receptacle of
this species is sometimes smooth (without scales) and sometimes
squamose. In this case, I see reason for using the artificial section
Depressae Clarke. The affinity of S. faminziniana Krassn. to series Salsae
is doubtful. ;

Series 20. Karaartschae Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 379.—Stem covered with hardened leaf sheaths
at base, wingless, somewhat leafy. Leaves succulent, extremely variable
from almost undivided to lyrate-pinnatisect. Capitula in type specimen
solitary or few, and then in lax corymbs. Outer involucral bracts
herbaceous, strongly acuminate with more or less bent cusp. The series
includes a single species.—S. karaartscha Saposhn.

512

506

99. S. karaartscha Saposhn. in Bot. Mat. Gerb. Glavn. Bot. Sada,
VI, 12, (1926) 13 1k

Perennial. Entire plant glabrous. Roots woody, many-headed, with
more or less numerous stems, densely covered at base with sheaths of
previous year’s leaves. Stems 12—25(35) cm high, ascending, slightly
flexuous, glabrous, sulcate, usually branched, less often simple. Leaves
fleshy, hard when dry, fairly variable on same plant; basal leaves long-
petiolate, petiole narrow, somewhat winged, broadened at base in sheath,
lamina 13-14 cm long (including petiole) and 2-4 cm wide, deeply
lyrate, with ovate and triangular lateral lobes terminating in small callose
cusp, and larger and broader, triangular or ovate terminal lobe, sometimes
entire, sometimes toothed, cuneate at base, sometimes only one terminal
lobe developed, and lateral lobes reduced to less numerous teeth; lower
cauline leaves also petiolate, lyrate; upper leaves sessile or subsessile.
Capitula solitary or two to seven on stem, terminal on main stem and
lateral branches arising from axils of upper leaves. Involucre 1.3—1.5
cm long and 0.6—1 cm wide, five-rowed, glabrous; outer involucral bracts
in type specimen like bracteal leaves, herbaceous, green; subsequent
bracts green, ovate, long-acuminate; inner bracts lanceolate or oblong,
reddish in upper part, long-acuminate, exserted from outer bracts;
acuminate tips of involucral bracts sometimes slightly bent. Receptacle
densely squamose, scales linear-subulate, lustrous, half or one-third as
long as involucral bracts; florets purple, 1.3—1.4 cm long, narrow part
of tube about 0.6 cm long, broad part with lobes of limb about 0.8 cm
long; anther appendages ciliate. Pappus double; outer pappus bristles
serrate-scabrous, unequal, small, inner bristles long-plumose, much longer
than outer. Immature achenes about 0.4 cm long. Flowering VII—VIII.
(Plate XXXVIII, Fig. 2.)

Stony slopes, coastal pebbles——Soviet Central Asia: Tien Shan.
Endemic. Described from the region of Kara-Archa (Tien Shan). Type
in Leningrad.

Note. Undoubtedly, it is phylogenetically close to S. salsa (Pall.)
Spreng., from which it differs by solitary or few capitula and the presence
of herbaceous, strongly acuminate outer involucral bracts, as well as by
the form of the leaves. Perhaps, it is also confused with S. faminziniana
Krassn. but it differs from the latter mainly by lyrate-pinnatisect leaves,
herbaceous, strongly acuminate outer involucral bracts and by form. From
S. blanda Schrenk., to which also it may be close, it differs by the
absence of pubescence on the leaves (leaves of S. blanda are strongly
scabrous from numerous stiff hairs along the margin), herbaceous,
strongly acuminate outer involucral bracts, and form of inflorescence.

Section 4. Jurineiformes (Lipsch.) Lipsch. in Bot. Zhurn. XLIII, 3
(1958) 452 pro serie—Semishrubs with numerous strongly and repeatedly

513

507

branched, virgate, erect, more or less glandular pubescent stems. Leaves
less numerous, small, lanceolate. Capitula numerous, solitary terminal
on branches and branchlets, forming lax paniculate inflorescence.
Involucre imbricate; involucral bracts with hard, callose cusps covered
with hairs. Receptacle densely squamose, scales subulately-linear,
unequal, lustrous. Achenes with small fringe (corona). Pappus white,
one-rowed, easily shedding, pappus-bristles equal or almost equal,
plumose, connate at base in ring.

100. S. chondrilloides Winkl. in Tr. Peterb. Bot. Sada, XI, 1 (1889)
169; Lipschtiz in Bot. Zhurn. XLIII, 3 (1958) 452.—Jurinea
chondrilloides (Winkl.) O. Fedtsch. in Perech. Rast. Turk. IV (1911)
296.—Saussurea aphylla Rech. f. Symb. Afghan. II (1955) 167.—Ie.:
Rech. f. Ibid. (1955) fig. 114.

Perennial. Semishrub, up to 75 cm high, with numerous, strongly
and repeatedly branched, virgate, erect, more or less glandular-hairy
stems; branches cylindrical, finely sulcate, pale green, terminating in
solitary capitula. Leaves few, small, coriaceous, lanceolate, about 2 cm
long and scarcely 2-3 mm wide, acuminate, sometimes weakly involute,
entire or with two or three teeth; upper cauline leaves significantly
reduced; uppermost leaves filiform-linear, weakly divergent; lamina with
small sessile glandular hairs on both sides. Capitula medium-sized,
turbinate, numerous, solitary terminal on branches and branchlets, forming
lax paniculate inflorescence. Involucre about 10—11 mm long; involucral
bracts imbricate, five- or six-rowed, appressed and short-pubescent,
sometimes sparsely glandular-hairy, with indurescent yellow or brownish
cusp covered with spinulose hairs; outer bracts reduced, inner elongate,
both lanceolate. Receptacle densely squamose, scales subulate-linear,
lustrous, almost as long as middle involucral bracts; florets pale pink
(?), numerous with small glandular hairs on outer surface; narrow part
of tube about 5 mm long, broad part including five-fid limb about 8—9
mm long; anther appendages woolly-hairy. Achenes 5 mm long, with
narrow corona at apex, comprising prominent, small, sometimes unequal
spinules; pappus easily shedding, simple, about 10 mm long, pappus
bristles white, soft, plumose, connate at base into ring. Flowering VIII-—
IX.

Stony deposits, pebbles, outcrops of conglomerates, outcrops of
varicolored rocks.—Soviet Central Asia: Pamiro-Alai. General
distribution: Iran (Afghanistan). S. chondrilloides Wink. is described
from Darvaz; type in Leningrad. S. aphylla Rech. f. is described from
Afghanistan. Type in Vienna.

Note. Comparison of the type specimens of S. aphylla Rech. f. and
S. chondrilloides Winkl. showed their complete identity. The transfer of

514

508

S. chondrilloides Wink\. to the genus Jurinea by O. Fedtschenko
(incidentally, without justification). was mistaken. S. chondrilloides has
a pappus of equal or subequal plumose, basally connate, one-rowed
bristles, as well as a receptacle covered with linear-subulate scales, and
undoubtedly should be placed in the genus in which C. Winkler described
it. But based on the presence on the achenes of a narrow crown of
small spinules, the presence at their tips of the involucral bracts of
indurescent cusps, the growth habit of the plant (semishrubs with
numerous virgate branches), and other characters, it should be placed in
a separate section. S. popovii Lipsch., distributed in Dzhungaria, is close
to S. chondrilloides, being distinguished by the form of its involucral
bracts, nature of the achenes (black with 4 white, slightly winged, ribs)
and other characters.

Section 5. Jacea Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVI (1954) 441.—Stem densely leafy. Capitula solitary terminal
on stem and lateral branches, in latter case numerous, usually surrounded
by uppermost leaves. Involucral bracts coriaceous, straw-yellow, with
chestnut or black border. Receptacle squamous, scales 1.5--2 cm long,
as long as or slightly longer than inner involucral bracts. Outer pappus
bristles short (when present), plumose like inner ones. Achenes linear.

101. S. masarica Lipsky in Tr. Peterb. Bot. Sada, XVIII, 1 (1900)
81; O. and B. Fedtsch. Perech. Rast. Turk., IV 234.

Perennial. Stems numerous, strongly virgately branched from base,
erect, sulcate, canescent or almost glabrous, densely leafy, wingless,
30-50 cm high. Leaves lanceolate or oblong-ovate, bicolored, green
above, snow-white beneath, appressed-hairy, up to 4—5 cm long and 1.5
cm wide, with short cusp, short-petiolate, petioles weakly winged; upper
cauline leaves sessile or subsessile; all leaves weakly toothed, more
often undivided; base of cauline leaves roundish or indistinctly cuneate.
Capitula with florets up to 1.5(2) cm long and 1—1.2 cm wide, solitary,
less often in twos, terminal on main stem and numerous lateral branches
that may be simple or branched in turn, forming lax and leafy panicle.
Involucre four-rowed, involucral bracts coriaceous, hard, imbricate,
sparingly and flocculosely arachnoid-hairy, especially along margin, or
almost glabrous, straw-yellow, with clear brown border, less often entirely
brownish; outer bracts smaller, ovate, inner much longer, linear-
lanceolate, both acuminate, with very short cusp. Florets reddish when
dry, about 14 mm long, narrow part of tube about 8 mm long, broad
part with short lobes of limb about 6 mm long; anther appendages
setaceous, about 1 mm long. Receptacle squamose, scales linear-subulate,
up to 1.5 cm long, as long as inner involucral bracts; pappus about 1.2

5]

Nn

509

cm long, white, pappus bristles connate at base into ring; outer bristles
shorter, but all of same shape, plumose; achenes immature, about
4-5 mm long, linear, glabrous. Flowering VII—VIII. (Plate XXX, Fig.
2.)

Borders of glaciers, stony slopes.—Soviet Central Asia: Pamiro-Alai
(Darvaz, Karategin). Endemic. Described from Darvaz and Karategin.
Type in Leningrad.

Note. A typical species of the Himalayan alliance, together with S.
jacea (Klotzsch) Clarke and S. tadshikorum \\jin and Gontsch. forming
a natural group of Saussurea.

102. S. tadshikorum Iljin and Gontsch. in Izv. Tadzh. Bazy Akad.
Nauk SSSR, Bot., I, 1 (1933) 67.

Perennial. Stem 40—50 cm high, wingless, sulcate, flocculose, almost
glabrous above, strongly leafy up to tip, branched from base; branches
numerous, slender, in turn branched. Leaves numerous, somewhat hard,
ovate-oblong or ovate, 2—2.5 cm long and 1—1.4 cm wide, narrowed
into very short, broadly cuneate petiole, almost entire or remotely and
indistinctly toothed, weakly involute, short acute; lamina concolorous,
green on both sides, more or less covered with small glandular hairs
beneath; lower leaves somewhat arachnoid-hairy, uppermost reduced.
Capitula 15-20 mm long and 8-12 mm wide, solitary terminal on
numerous branches, forming lax, almost paniculate inflorescence.
Involucre almost cylindrical, four-rowed, imbricate; involucral bracts
coriaceous, hard, straw-yellow, with dark brown border, weakly ciliate-
arachnoid hairy or almost glabrous; outer and middle involucral bracts
ovate or oblong-ovate, with indistinct midrib, narrowed into a short cusp;
inner bracts linear, acuminate, much longer than outer. Florets reddish
(?) (according to M.M. Iljin, whitish, which is not distinct in the
herbarium material), about 14 mm long, narrow part of tube about 8
mm long, broad part with lobes of limb about 6 mm long, lobes of limb
1.5—2 mm long; anther appendages setaceous, about 1 mm long.
Receptacle squamose, scales linear-subulate, lustrous, up to 18-20 mm
long; pappus about 12 mm long, pappus bristles long-plumose, connate
at base into ring, outer hairs shorter, but also plumose; achenes linear,
5—6 mm long. Flowering VII.

Pebbles, stony slopes.—Soviet Central Asia: Pamiro-Alai. Endemic.
Described from the Vakhsh Range. Type in Leningrad.

Note. Besides the three sheets of the type material, there are no
other collections in the Herbarium of the Botanical Institute, Academy
of Sciences of the USSR. The species is very close to S. masarica
Lipsky and S. jacea (Klotzsch) Clarke. It differs from S. masarica by
its leaves, green on both sides with minute punctate glandular hairs

516

510

beneath, and from S. jacea by numerous, smaller capitula, and also by
smaller leaves, which are smooth or weakly scabrous on both sides.

Section 6. Depressae Clarke. Comp. Ind. (1876) 221; Kitam. in
Mem. Coll. Sc. Kyoto Univ. ser. B, XIII (1937) 148.—Sect. Gymnoclyne
Franch. in Journ. de bot. II (1883) 311; Kitam. in Acta Phyt. et Geobot.
IV (1935) 4.—Capitula solitary or aggregated at apices of stem and its
branches into compact corymbs. Involucral bracts without appendages.
Receptacle glabrous, without scales. Pappus two-rowed, outer pappus
bristles reduced, fragile, inner bristles oblong, plumose; achenes without
apical fringe (corona).

Note. The distinctive feature of the species grouped into this section
is the absence of scales on their receptacles. In other respects, the species
included in this section do not differ sharply from the species included
in other sections of the genus. As a result, the section is an artificial
one and, likely, will be reduced in future. Of the saussureas growing
within the USSR, it includes S. nuda Ldb., S. caprifolia Ijin and
Zaprjagaev, S. tschuktschorum Lipsch.

103. S. nuda Ldb. Ic. pl. Fl. Ross. I (1829) 15; Fl. alt. IV, 16 in
nota; DC. Prodr. VI, 536; Ldb. Fl. Ross. II, 2, 662; Rgl. in Mém. Soc.
Nat. Mosc. XI, 106 (saltem a. genuina and B. subsinuata, excl. y. tilesii);
Kom. Fl. Kamch., III, 182; Hult. Fl. of Alaska and Yukon, X, 1626.—
S. ledebouri a. nuda Rgl. ex Herd. in Bull. Soc. Nat. Mosc. XLI, 3
(1868) 42.—S. subsinuata Ldb. Ic. pl. Fl. Ross. I (1829) 15; Ej. FI. alt.
IV, 16; DC. Prodr. VI, 536; Ldb. Fl. Ross. II, 2, 662; Kom. Fl. Kamch.
Il], 181; Hult. Fl. Kamtch. IV, 210; Kitam. in Mem. Coll. Sc. Kyoto
Univ. Ser. B, XIII, 148.—S. ledebouri 8. subsinuata Rgl. ex Herd. in
Bull. Soc. Nat. Mosc. XLI, 3 (1868) 42.—S. nuda var. subsinuata B.
Fedtsch. Fl. II. Comm. (1906) 77.—S. alpina auct.: Lessing in Linnaea,
VI (1831) 87.—?S. morozeviczii B. Fedtsch. Fl. II. Comm. (1906) 77.—
Ic.: Ldb. op. cit. (1829) tab. 61, 62.

Perennial. Stem 10—20 (up to 40) cm high, erect or ascending in
lower part, wingless or with narrow wings, glandular pubescent or almost
glabrous, sulcate, covered at base with remains of old: leaves. Leaves
thicker, fleshy, sometimes thinner, green on both sides, often light-colored
and usually crisped-pubescent beneath, especially densely along leaf
margin, less often almost glabrous, sinuate-toothed to entirely toothed —
or almost undivided, teeth with callose tip, or subacute, less often
roundish-obtuse; basal and lower cauline leaves petiolate, petioles winged,
lamina oval, oblong or lanceolate, together with petiole 3.5—12 cm long,
(0.5)1.5—2 cm wide (individual basal leaves reaching 25 cm in length
and 3 cm in width, petioles 12.5 cm long), cuneate at base or less often

511

517 Plate XXXVIII.

1—Saussurea turgaiensis B. Fedtsch.; 2—S. karaartscha Saposhn.

518

512

roundish; middle cauline leaves short-petiolate or subsessile, weakly
amplexicaul or sometimes slightly decurrent on stem; upper leaves
gradually reduced, linear or lanceolate, sessile. Capitula 10-16 on
peduncles, aggregated into a terminal more or less dense corymb, less
often capitula solitary or in small number; peduncles short, sometimes
up to 4-5 cm long, ferruginous or white arachnoid-woolly, mixed with
lustrous glandular hairs; bracteal leaves linear, cuspidate. Involucre
campanulate, 11-13 mm long and wide; involucral bracts four-rowed,
light or dark-colored; outer bracts as long or almost as long as inner
ones, all lanceolate, cuspidate, ferruginous or whitish villous on dorsal
surface, especially along margins, usually (but not always) ferruginous-
glochidiate at apex. Receptacle smooth, without lustrous scales; florets
pinkish-violet, about 10-12 mm long, narrow part of tube as long or
almost as long as its broad part with lobes of limb; anther appendages
ciliate. Pappus weakly ferruginous, double; outer pappus bristles serrate,
small, fragile, inner bristles long-plumose, about 9-10 mm long;
achenes linear, about 6 mm long, 1 mm wide, glabrous. Flowering VII—
Wile

Meadows and grassy patches in the forest and alpine zone, mossy
marshes, stony outcrops, rocks, sea coast—Arctic: Chukotka, Anadyr;
Far East: Kamchatka and Commander Islands (northern part), Okhotsk,
Uda, Sakhalin (western part opposite the ‘liman’ of the Amur River)
(Kurils). General distribution: Alaska. S. nuda Ldb. was described from
Kamchatka and Cape Espenberg, S. subsinuata from Kamchatka; types
preserved in Leningrad. S. morozeviczii B. Fedtsch. was described from
the Commander Islands; E. Hultén saw the type specimens of the
latter and identified this species with S. subsinuata: at present, the type
of S. morozeviczii is missing from the Herbarium of the Botanical
Institute.

Note. A polymorphic species. Varies in shape and incision of leaves
and their pubescence, as well as decurrence along stem; stem height;
number of capitula (alpine forms are sometimes unicapitulate—f. uniflora
Lipsch., or with few capitula; low altitude forms usually with larger
number of capitula aggregated in terminal, and rarely also lateral,
corymbs); color of involucral bracts (green or anthocyanin-colored).
Evidently, this variation is related to the amplitude of fluctuations in
ecological conditions where the species grows. A comparison of the
type specimens of S. nuda and S. subsinuata leaves no doubt about
their identity. S. nuda also figures incorrectly alongside S. subsinuata
Ldb. in the manuals on the flora of Kamchatka of Komarov and Hultén. -
Later (in 1950), in the Flora of Alaska and Yukon, Hultén, independently
of me, came to the conclusion that the two species are the same and
need to be combined into a single species under the name S. nuda Ldb.

513

The specific epithet nuda, as rightly pointed out by Hultén, “symbolizes”
its most typical and stable character, the smooth, nonsquamose receptacle.
Evidently, S. nuda forms hybrids with S. oxyodonta Hult., where they
come in contact; these specimens have small scales on the receptacle
and more acute and frequent teeth along the leaf margin, but the nature
of the involucre is the same as in S. nuda. Such, for example, are the
few specimens from the Anadyr Territory, where both species are found.
It appears to me that S. nuda is the closest phylogenetically to S.
oxyodonta, but always has a smooth receptacle and almost equal
involucral bracts. A few specimens with dense, snow-white, tomentose
indumentum on the lower leaf surface and a smooth receptacle were
found among the collections from the Bering islands. These are
undoubtedly hybrids between S. nuda and S. pseudo-tilesii, as the range
of the latter species extends to the Commander Islands, where these
species grow together and may form natural hybrids.

104. S. tschuktschorum Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVI (1954) 452.—S. angustifolia auct., p. p.

Perennial. Stem 6—10(12) cm high, simple, weakly flattened, finely
sulcate, very sparsely pubescent or almost glabrous, sparsely covered at
base with remains of previous year’s leaves. Leaves linear or lanceolate-
linear, usually narrow, about 2 mm, less often up to 5 mm wide, entire,
more or less convolute, arachnoid-hairy beneath. Capitula pedunculate,
aggregated in terminal, relatively dense corymbs; bracteal leaves one to
three, approximate, linear longer than capitate corymbs. Involucre up to
1 cm long, imbricate, with soft squarrose hairs; involucral bracts
lanceolate, outer short, inner bracts longer than outer, both subobtusely
acuminate, often terminating with a tuft of white hairs. Receptacle
smooth, without scales; florets about 10-11 mm long, narrow part of
tube as long as broad part with lobes of limb. Pappus brownish, double;
outer pappus bristles 1.5—2 mm long, fragile, serrate-scabrous; inner hairs

519 plumose, about 10 mm long; achenes dark brown, glabrous, immature,
soft. Flowering VII.

Arctic: Chukotka, Anadyr. Endemic (?). Described from the Anadyr
River. Type in Leningrad.

Note. In appearance it resembles S. angustifolia (Willd.) DC., but
differs by a smooth receptacle, 1-3 linear subtending leaves exceeding
the inflorescence, and narrower involucral bracts. Phylogenetically it
undoubtedly is close to this species, but is included under section
Depressae on the basis of the smooth receptacle.

105. S. caprifolia [ljin and Zapriagaev in Izv. Tadzh. Bazy Akad.
Nauk SSSR, Bot. I, 1 (1933) 69-70.—Ie.: Ibid. 68.

520

514

Perennial. Miniature plant, stemless or with 2.5 cm high stem, in
latter case stem arachnoid-hairy, covered by rosulate basal leaves, basal
leaves roundish or almost so, shortly spinescent, entire or less often
with solitary, indistinct, spinescent small teeth, roundish or broadly
cuneate at base, distinctly petiolate, petioles broadened at base into short
sheaths covering root collar; all leaves weakly pubescent above, finely
tomentose beneath, with distinct midrib, with more or less numerous
lustrous, small glandular hairs on both sides; cauline leaves reduced,
sessile, one to three. Capitula less often solitary, often aggregated in
groups of two or three (five), terminal. Involucre cupuliform, somewhat
imbricate, about 1.2 cm long and about 1 cm wide; outer involucral
bracts dark-colored, ovate, acuminate, cusp short, sometimes weakly bent,
arachnoid-hairy, inner bracts slightly longer than outer and middle, oblong
or lanceolate, acuminate, pubescent, with tuft of soft hairs at apex.
Receptacle smooth, without scales; florets pale pink, about 1.2 cm long,
narrow part of tube almost as long as broad part with lobes of limb
covered with lustrous glandular hairs; anther appendages ciliate. Pappus
double; outer pappus bristles less numerous or isolated, serrate, small,
fragile, inner bristles long-plumose, about 1—1.1 cm long; achenes 4—
4.5 mm long, oblong-linear, glabrous. Flowering VIIJ-IX.

Stony screes in alpine zone.—Soviet Central Asia: Pamiro-Alai
(Darvaz Range). Endemic. Described from the Kurgovady Pass, 3800—
3900 m. Type in Leningrad.

Note. A striking, acaulescent species, particularly close to
S. kuschakeviczii Winkl., differing from it by having more rounded,
entire leaves, fewer capitula, which in the type are much larger,
and, most importantly, an entirely smooth receptacle. Because of
its smooth receptacle, it is formally put in section Depressae Clarke.
Phylogenetically it undoubtedly is close to series Kuschakevicziae
Lipsch.

Subgenus 6. Theodorea (Cass.) Lipsch. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XXI (1961) 379.—Gen. Theodorea Cass. Dict.
sc. nat. LIII (1828) 463; Benth. and Hook Gen. Pl. IJ, 472.—Sect.
Theodorea (Cass.) DC. Prodr.. VI (1837) 536; Endlich. Gen. pl. 468;
Turcez. Fl. baic.-dahur. II, 1, 118; Franch. in Bull. Herb. Boiss. V, 534;
Nakai in Bot. Mag. Tokyo, XXIX, 194 (pro subgen. sub falso nom.
Theodorea (Cass.) DC.), 207 (pro subgen. sub falso nom. Theodorea
Cass.); Kitam. in Acta Phyt. et Geobot. IV, 3; Kitam. in Mem. Coll. Sc.
Kyoto Univ. ser. B, XIII, 142; Popov, Fl. Sr. Sib. Il, 755.—Sect. Elatae
Hook. f. Fl. Brit. Ind. III (1882) 373 p. p.; Pflanzenfam. IV, 5, 320 p.
p.—Rhizome biennial, fusiform, or plants perennial. Stem tall, branched,
more or less densely leafy. Capitula numerous, in corymbs. Involucre

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imbricate; involucral bracts, at least inner, with membranous, usually
colored, sometimes more or less toothed apical appendages. Receptacle
densely squamose; anther appendages as small number of hairs. Pappus
double; outer pappus bristles reduced, fragile, inner ones plumose,
achenes without corona.

Series 1. Amarae Lipsch.—Subsect. Theodorea-vera Kitam. in Acta
Phyt. et Geobot. IV (1935) 3 p. p.; Ej. in Mem. Coll. Sc. Kyoto Univ.
ser. B, XIII (1937) 143 p. p—Stem wingless. Leaves simple, entire or
sinuate-toothed, but not pinnatisect. Capitula cylindrical or narrow
campanulate. Outer rows of involucral bracts with dark green teeth or
three-fid apical appendages, others (middle and outer) terminating into
colored membranous appendages.

106. S. amara (L.) DC. in Ann. Mus. Paris, XVI (1810) 200; Ldb.
Ic. pl. Fl. Ross. I, 16; Ldb. Fl. alt. 1V, 21; DC. Prodr. VI, 536; Ldb. FI.
Ross. II, 2, 664; Herd. in Bull. Soc. Nat. Mosc. XLI, 3, 47 p. max. p.;
Kryl Fl. Alt. III, 702; Kom. Fl. Manchzh. III, 2, 719; Fl. Yugo-Vost.
VI, 387; Kryl. Fl. Zap. Sib. XI, 2910; Popov, Fl. Sr. Sib. II, 755.—
Serratula amara L. Sp. pl. I (1753) 819; willd. Sp. pl. HI, 3, 1644.—
Saussurea marginata Borszcz. in Zap. Akad. Nauk, VII, Appendix 1
(1865) 31 nomen.; Trautv. in Tr. Peterb. Bot. Sada, VIII, 2, 482.—_S.
glomerata Poir. Encycl. Meth.; Suppl. V (1827) 71; Ldb. op. cit. (1829)
17; Ldb. op. cit. (1833) 21; DC. op. cit. (1837) 536; Ldb. op. cit. (1845—
1846) 664; Turcz. Fl. baic.-dahur. II, 1, 118; Kom. op. cit. 724.—
S. amara var. glomerata (Poir.) Trautv. in Bull. Soc. Nat. Mosc. XXXIX,
1 (1866) 369; Kryl. Fl. Zap. Sib. XI, 2910—S. amara B. glomerata
Trautv. in Herd. op. cit. 48.—S. glomerata Poir. var. angustifolia Freyn
and var. intermedia Freyn in Oest. bot. Zeitschr. XLV, 11 (1895)
432.—? S. centauroides Tausch in Flora, XI, B, 2, No. 31 (1828) 484
(secund cl. Ledebour = S. glomerata Poir.}—? S. macrocephala Less. in
Linnaea, VI (1831) 87 (apud cl. Ledebour FI. Ross. 665 est synon. S.
glomeratae; ego specimina non visa; secundum cl. Kitamura (1937) 146
= §. japanica DC.).—? S. scabra Less. in Linnaea, IX (1834) 180 (apud
cl. Ledebour op. cit. est synon. S. glomeratae, secundum cl. Kitamura
= §. japonica DC.).—S. gmelini Hort. Dorp. ex Herd. op. cit. 48.—S.
amara var. integrata DC. in Ann. Mus. Paris, XVI (1810) 200.—Ie.:
Ldb. Ic. pl. Fl. Ross. I (1829) tab. 67, 68—Exs.: GRF Nos. 1228a,
3988.

Perennial. Entire plant glaucescent, usually scabrous, less often
smooth. Stem erect, wingless, distinctly sulcate, branched above,
sometimes simple, weakly pubescent, or often almost glabrous, 15-60
em high. Leaves fairly variable in shape and incision; basal and lower

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516

cauline leaves long-petiolate, elliptical or oblong-elliptical, acuminate,
entire or sinuate-toothed, teeth unequal, sometimes attenuate, lamina 5—
20(28) cm long and 1.5—8(12) cm wide; cauline leaves short-petiolate
or sessile, sometimes auriculate or scarcely decurrent, lamina smaller,
oblong or lanceolate, entire; all leaves green on both sides, or somewhat
pale beneath with numerous glandular hairs, usually scabrous due to
somewhat pointed tubercles. Capitula campanulate,. 1-1.5 cm wide or
smaller, or cylindrical (in var. glomerata), less often solitary terminal
on stem, usually forming more or less dense corymbose-paniculate
inflorescence. Involucre of unequal, imbricate bracts, sometimes weakly
obtuse, sometimes (often) pubescent; outer involucral bracts smaller,
more or less lanceolate, with dark green, somewhat thick, oval or oblong
teeth or three-fid appendages; middle bracts oblong or oblong-linear,
broadened at apex into roundish, membranous, toothed, pink appendages
two or three times or somewhat (var. glomerata) broader than bract;
innermost involucral bracts linear, with or without narrow appendage.
Florets pink or less often white (f. albiflora Lipsch.), about 15 mm
long, narrow part of tube almost one and one-half to two times as long
as its broad part with lobes of limb; anther appendages weakly pubescent.
Receptacle densely squamose scales linear-subulate, lustrous, unequal,
up to 7 mm long. Pappus double; outer pappus bristles serrate, small,
numerous, unequal, although fragile, but persistent on achenes, inner
bristles plumose, about 10 mm long; achenes about 3 mm long, smooth,
without corona. Flowering VII-IX.

Saline and alkaline soils, saline meadows.—European Part: Middle
Dnieper. Lower Volga, Trans-Volga Region, Volga-Kama; Western
Siberia: Upper Tobol, Irtysh, Altai; Eastern Siberia: Angara-Sayans,
Dauria, Lena-Kolyma; Soviet Central Asia: Aralo-Caspian Region,
Balkhash Region, ?Kara-Kum, ?Kyzyl-Kum, Tien Shan, Dzhungaria-
Tarbagatai. General distribution: Mongolia, China (north).

Note. From time to time, populations with white corollas (albinos)
are found, (f. albiflora Lipsch.), or one-headed plants occur (f.
monocephala Lipsch.). The characters cited by Ledebour (Fi. alt. IV,
22) for distinguishing S. amara from S. glomerata prove unstable when
extensive material from different parts of its natural range are examined...
A unique race is found in Dauria, which is characterized by much smaller
and narrower capitula with fewer florets (these capitula are 2/3 as wide
as in typical S. amara), in lax and irregular corymbs, simulating a
panicle, by the indistinct smaller appendages of the middle involucral
bracts, the latter being narrower in the type specimen than in true S.
amara, and also by narrow lanceolate leaves. However, this race produces
a great many forms transitional to typical S. amara, and, therefore, I do
not recognize it. Apparently, if S. amara and S. glomerata are recognized

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SG |

as separate species, then the name S. glomerata sensu Poiret should be
applied to this race and not to the numerous forms that cluster around
S. amara, which have been treated by a series of Russian authors,
including Trautvetter, sometimes as S. glomerata and sometimes as S.
amara var. glomerata. Because of the lack of material from Japan, I am
not clear about the relationship of S. glomerata with S. japonica (Thunb.)
DC. (© Serratula japonica Thunb.) non auct. plur. It appears: that S.
japonica (Thunb.) DC. is identical with S. glomerata Poir. or is quite
close to it. Since much confusion has been created in relation to S.
japonica and many authors, including in this country, have incorrectly
used this name for another species, i.e., S. pulchella Fisch. (cf. below),
as was correctly pointed out by Handel-Mazzetti, it is better to
discontinue the use of the name S. japonica. Kitamura (Mem. Coll. Sc.
Kyoto Univ. ser. B, XIII (1937) 146), however, continued to use this
name for the island form (or species?) of “S. amara,” without comparing
the Japanese material with S. glomerata and without including the latter
among the synonyms of his own S. japonica. But he has included a
number of species among the synonyms of S. japonica (Thunb.) DC.
(for example, S. macrocephala Less., S. scabra Less.) that Ledebour,
based on his acquaintance with it in the Berlin Herbarium, referred
without question to S. glomerata Poir.

Series 2. Pulchellae Lipsch.—Subsect. Theodorea-vara Kitam. in
Acta Phyt. et Geobot. IV (1935) 3 p. p.; Ej. in Mem. Coll. Sc. Kyoto
Univ. ser. B, XIII (1937) 143 p. p—Stem more or less winged or
wingless. Leaves pinnatisect. Capitula almost globose or broadly-
campanulate with large, colored, membranous appendages surrounding
involucral bracts, usually beginning from outermost bracts.

107. S. pulchella Fisch. in DC. Prodr. VI (1837) 537; Lindl. in
Bot. Regist. XXVIII, t. 18; Ldb. Fl. Ross. II, 2, 664; Turcz. Fl. baic.-
dahur. II, 1, 119; Maxim. Prim. fl. amur. 171 p. p. excl. var. Jatifolia
Maxim.; Rgl. in Mém. Acad. Pétersb. sér. VII, IV, 4, 93-94 (incl. a.
subinterga Rgl. y. pinnatifida Rgl. 5. ovata Rgl.); Herd. in Bull. Soc.
Nat. Mosc. XLI, 3, 50-51 excl. syn. pl.; Nakai in Bot. Mag. Tokyo,
XLVI, 615; Hand.-Mazz. Symb. Sin. VII, 4, 1145 in nota; Kitam. in
Mem. Coll. Sc. Kyoto Univ. ser. B, XIII, 143; Popov, Fl. Sr. Sib. I],
757.—Heterotrichum pulchellum Fisch. in Mém. Soc. Nat. Mosc. III
(1812) 71.—Serratula pulchella Sims. in Bot. Mag. LII (1825) tab.
2589.—Theodorea pulchella Cass. in Dict. sc. nat. LIII (1828) 465.—
Saussurea dissecta Ldb. Ic. pl. Fl. Ross. I (1829) 16, tab. 66; Ej. FI. alt.
IV, 19.—S. pulchella var. lineariloba Nakai in Bot. Mag. Tokyo, XLVI
(1932) 616; Kitam. op. cit. 146.—S. japonica auct. plur. non DC.;

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518

Maxim. in Mél. Biol. LX, 3, 337 (p. p., conf. observatio cl. Maximovicz
ad S. pulchella); Kom. Fl. Manchzh. III, 727, p. p.; Kom. and Allis.
Opred. Rast. Dalnevost. Kr. II, 1063, 1174.—S. pulchella Fisch. var.
subintegra Rgl. Tent. Fl. Ussur. (1861) 93; Herd. in Bull. Soc. Nat.
Mosc. XLI, 3, 50; Nakai in Bot. Mag. Tokyo, XLVI, 615.—S. japonica
var. subintegra (Rgl.); Kom. Fl. Manchzh. II (1907) 729; Kom. and
Alis. Opred. Rast. Dalnevost. Kr. II, 1174.—S. pulchella var. pinnatifida
Rgl. Tent. Fl. Ussur. (1861) 94; Herd. in Bull. Soc. Nat. Mosc. XLI, 3,
50.—S. japonica var. pinnatifida (Rgl.) Kom. op. cit. 729; Nakai in
Bot. Mag. Tokyo, XXIX (1909) 209.—S. pulchella var. ovata Rgl. Tent.
Fl. Ussur. (1861) 94; Herd. op. cit. (1868) 51.—S. japonica var. ovata
(Rgl.) Kom. op. cit. 729.—? S. pulchella var. alata Rgl. Tent. Fl. Ussur.
(1861) 93.—? S. japonica var. alata (Rgl.) Kom. Fl. Manchzh. III (1907)
729.—S. japonica var. pterocaulis Franch. in Bull. Herb. Boiss. V, 7
(1897) 536.—Ie.: Bot. Mag. LII (1825) tab. 2589; Ldb. Ic. pl. Fl. Ross.
I, tab. 66; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, Plate 317.—
Exs.: GRF Nos. 3993, 3994.

Perennial. Root vertical, fusiform, bearing one or two stems. Stem
25-100 cm high, erect, strong, sulcate-angular with reddish furrows,
simple or branched above, pubescent or glabrous, often wingless, less
often more or less winged due to decurrent leaf bases, somewhat densely
leafy. Leaves numerous, appressed to stem or spreading, pinnatisect,
pinnatipartite, more or less deeply pinnatifid or almost entire, with
shallow sinuses along margin; hard or soft, scabrous above due to short
tubercles and sessile glandular hairs, flocculose beneath with numerous
punctate glandular hairs; basal and lower cauline leaves long-petiolate
in typical form; upper leaves short-petiolate or almost sessile; lamina
wide, oblong or elliptical, pinnatisect into more or less narrow linear or
lanceolate-linear, elongate, acuminate lobes, usually secondarily divided,
undivided or toothed, secondary lobules or teeth cuspidate; midrib
prominent on leaf axis, lateral veins solitary on primary lobes; cauline
leaves gradually reduced above, uppermost leaves lanceolate or linear,
with pinnatilobate or pinnatifid margin, sometimes undivided. Capitula
numerous in type specimen, on slender, 1-5 cm long pubescent
peduncles, aggregated in more or less dense corymbs, sometimes forming
dense panicle, compising up to 100 capitula, usually corymbs with 5—
20 capitula; capitula almost globose or broadly campanulate, roundish
at base. Involucre 10-15 mm wide; involucral bracts numerous, weakly
pubescent; outermost bracts one-rowed, reduced, two-fifths as long as
inner (including its appendages), linear, green, subsequent, middle and
inner bracts gradually longer, oblong in lower part or ovate, yellowish,
lustrous, narrowed above in distinctly developed, membranous, roundish,
toothed, pink-violet appendage. Receptacle densely squamose, scales

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p19

linear-subulate, lustrous, up to 6—7 mm long; florets.purple, 12-13 mm
long, narrow part of tube 7-8 mm long; broad part with lobes of limb
about 5 mm long, lobes of limb with lustrous glandular hairs; anther
appendages hairy. Pappus double; outer pappus bristles numerous,
reduced, scabrous, although fragile yet persistent on achenes, inner
bristles plumose, 7-8 mm long; achenes 3.5—4 mm long, truncate at
apex, weakly narrowed at base, glabrous, with reddish longitudinal
speckles. Flowering VII-—VIII.

Dry and meadow steppes, flooded meadows, shrub thickets,
wastelands, forest edges.—Eastern Siberia: Dauria; Far East: Zeya-
Bureya, Ussuri, Uda, Sakhalin. General distribution: Japan, Korea, China,
Mongolia (northeastern). Described from Dauria. Type in Leningrad. S.
dissecta was described from Dauria; type in Dauria.

Note. As was explained by Nakai (Bot. Mag. Tokyo, XLVI (1932)
614) and independently by Handel-Mazzetti (Symb. Sin. VI, 4 (1936)
1145) from a study of the authentic specimen of Serratula japonica
Thunb. (= Saussurea japonica (Thunb.) DC.) preserved in Uppsala, the
plant described by Thunberg is close to or identical with Saussurea
glomerata Poir., 1.e., it belongs to our series Amarae. S. japonica sensu
Thunb. and sensu DC. does not have anything in common with the
Saussurea examined, for which many authors have wrongly used this
name. The use of the latter name for the species under consideration
has caused considerable confusion; therefore, it should be discarded.
Saussurea pulchella is the valid name for this species, which was
understood and recognized by Maximovicz (1859) and Regel (1861).
Notwithstanding the opinion of Nakai (Bot. Mag. Tokyo, XXV (1911)
58; XXIX (1915) 209), S. dissecta Ldb. is a direct synonym of S.
pulchella Fisch. This is evident from the comparison of the authentic
specimens of both species originating from Dauria. Ledebour himself
later included his S. dissecta among the synonyms of S. pulchella Fisch.
In the Trans-Baikal Region, from where S. pulchella was described, it
is represented by uniform material; its lamina is bipinnate or simply
incised into lanceolate-linear or linear-elongate lobes. Saussurea pulchella
y. pinnatifida Rgl., described from the Far East, judging from the original
specimens is typical S. pulchella as understood by Fischer. As the Far
East is approached, plant populations are found together with typical S.
pulchella, that are distinguished by: 1) a winged or wingless stem and,
particularly, 2) the form of the leaves (width, length but mainly their
incision; the leaves vary greatly—they are sometimes pinnate, sometimes
more or less entire or toothed and constitute the gamut of transitional
forms), and 3) the number of capitula (one can distinguish two series of
populations: few-headed—paucicephalae Lipsch., and many-headed—
pluricephalae Lipsch.).

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520

Infraspecific classification of S. pulchella has been attempted by
C.J. Maximovicz (op. cit. 1859), E. Regel (1861), V.L. Komarov (1907,
1932) and others. The main varieties are the following:

Var. Subintegra Rgl.—Leaves weakly decurrent; basal and lower
cauline leaves pinnatifid; upper leaves entire or weakly dentate,
lanceolate, lanceolate-linear, sometimes ovate. Identical with it is var.
ovata Rgl. having, according to the author, a winged stem, pinnatifid
lower leaves, the middle and upper leaves ovate and almost entire, and
the uppermost lanceolate. In my view, var. ovata Rgl. has no taxonomic
significance, because Regel’s varieties were established on the basis of
two uncorrelated characters: the wingedness of the stem and the cutting
of the lamina. These unrelated characters used by him as the basis for
the infraspecific classification of S. pulchella, are not restricted to the
varieties described by him. Thus, var. alata Rgl., described at the same
time by Regel, which I include as well, with some doubt, among the
synonyms of subintegra Rgl., likewise has a winged stem as is seen
also in some specimens of other varieties described by other authors,
for example, var. latifolia Maxim. (see below). Among new collections
there are no specimens identical with the authentic specimens of var.
ovata Rgl. and alata Rel.

Var. dentata Kom. Fl. Manch. III (1907) 729 has pectinate to
coarsely toothed lower and middle cauline leaves and lanceolate,
acuminate upper cauline leaves. It forms transitions to var. subintegra
Rel.

The unique var. latifolia Maxim., first described by Maximovicz
from the lower reaches of the Amur and subsequently found in Sakhalin,
deserves the greatest attention. This apparently is a separate race of S.
neopulchella Lipsch.

108. S. neopulchella Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 380.—S. pulchella B. latifolia Maxim.
Prim. fl. amur. (1859) 171; Sugaw. Ill. Fl. Saghal. IV, 1869, tab. 858.—
S. pulchella 8. alata auct. (non Rgl. 1861).—S. japonica B. alata
Rgl. p. p. and y. latifolia Maxim. ex Kom. Fl. Manchzh. III (1907)
HDD:

Perennial. Stem winged due to decurrent leaves (wings sometimes
entire or weakly toothed), branched, up to 150 cm high. Leaves large,
isolated leaves up to 25 cm wide, strongly scabrous beneath due to
numerous bristles; basal leaves (if present) and lower cauline leaves
pinnatipartite; middle cauline leaves pinnatifid; uppermost leaves almost
undivided; lobes of pinnate leaves varying from oblong, sinuate-toothed
and subobtuse to lanceolate, almost entire, acute; venation prominent,
midrib strongly prominent, lateral veins continued into leaf lobes, slightly

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521

weaker. Capitula numerous; involucre densely arachnoid-pubescent.
Flowering VII—VIII.

Tall herb meadows; chains of coastal rocks.—Far East: Ussuri
(eastern part), Zeya-Bureya, Sakhalin. Described from Sakhalin. Type
in Leningrad.

Note. The specimens identified by many authors as var. alata are
not identical with Regel’s variety, Var. alata Rgl. non auct., despite
having a winged stem, seems to be closer to var. subintegra Rgl.; I
consider them identical. Most specimens taken by the authors to be var.
alata are typical representatives of S. neopulchella.

Series 3. Sclerodontae Maxim. in Mel. Biol. XI (1881) 254.—Plants
with distinctly winged (due to decurrent leaves) or wingless stem,
sometimes stemless. Leaves pinnati- or bipinnatisect. Teeth of leaves
and sometimes involucral bracts with cartilaginous cusp.

109. S. alata DC. in Ann. Mus. Paris, XVI (1810) 202; Ldb. Ic. pl.
FI. Ross. I, 15; Ej. Fl. alt. 1V, 16; DC. Prodr. VI, 537; Ldb. Fl. Ross. II,
663; Turcz. Fl. baic.-dahur. II, 1, 120; Herd. in Bull. Soc. Nat. Mosc.
XLI, 3, 45 (excl. y. pygmaea Fisch. 6. runcinata, ¢. albiflora, &.
lanciniata); Popov. Fl. Sr. Sib. I, 755.—S. laciniata var. lasiocephala
Turez. ex DC. Prodr. VI (1837) 537; Turcz. op. cit. (1856) 120.—S.
alata §. aspera Ldb. Fl. Ross. UH, 2 (1845-1846) 663; Herd. op. cit.
45.—S. alata DC. a. genuina Herd. op. cit. (1868) 45.—Heterotrichum
squalidum MB. ex DC. Prodr. VI (1837) 537 nomen.—Serratula squalida
Steph. ex Ldb. FI. alt. [V (1833) 17 nomen.—S. salsuginosa Sievers ex
Herd. op. cit. (1868) 45.—S. decurrens Ldb. Ind. sem. hort. Dorp. suppl.
(p. a. 1811) 4 (opusc. non v.); Ldb. op. cit. (1845-1846) 663 pro syn.—
Ic.: Ldb. Ic. Pl. Fl. Ross. I (1829) tab. 63.

Perennial. Entire plant strongly or weakly scabrous due to somewhat
thick articulate hairs with stiff base. Root collar sparsely covered with
remains of hard sheaths of basal leaves. Stem erect, 20—35(50) cm,
sometimes about 100 cm high, simple or branched, sulcate-angular,
distinctly winged due to decurrent leaves, wings broad or narrow, toothed.
Leaves pinnatisect; basal and lower cauline leaves petiolate with
sheathing base, lamina strongly scabrous, with numerous sessile glandular
hairs, almost lyrately and deeply pinnatifid or pinnatisect, with unequal,
triangular, lanceolate, or oblong, lateral lobes, sinuate-toothed, teeth
unequal terminating in catilaginous cusp; upper cauline leaves oblong,
pinatifid, cristate or less often almost undivided, sessile, with decurrent
base, acute, strongly scabrous along margin, bracteal leaves linear-
lanceolate. Capitula 8-10 mm wide, in dense, frequent corymbs of less
numerous heads terminating on stem and its branches forming paniculate
inflorescence with strongly branched stem. Involucre imbricate, oblong

522

or ovoid; involucral bracts strongly arachnoid-hairy in type specimen
often glochidiate with white hairs; outer bracts herbaceous, oblong, acute,
slightly deflexed; inner bracts coriaceous, linear-lanceolate, subobtuse,
with indistinct purple, membranous, toothed apical appendage or tuft of
hairs, usually larger than outer. Florets reddish, about (11)13-14 mm
long, narrow part of tube slightly longer or almost as long as broad part
with lobes of limb; lobes of limb about 3 mm long; anther appendages
comprising few hairs. Receptacle densely squamose, scales linear-
subulate, lustrous, unequal, 4-5 mm long; pappus double, outer pappus
bristles unequal, up to 5 mm long, serrate, persistent on achenes, inner
bristles long-plumose, equal, about 10 mm long; achenes glabrous.
Flowering VIII.

Saline lands, saline meadows and sands.—Eastern Siberia: Angara-
Sayans (Tuva), Dauria. General distribution: Mongolia. Type specimen
comes from Siberia (‘“Sibiria salsuginosis”), is preserved in Geneva.
Type of S. decurrens Ldb. in Leningrad.

Note. Ledebour (FI. Ross. II, 2, 661) considers S. alata DC. in the
group of saussureas characterized by almost equal involucral bracts, i.e.,
he split off this species from others which are closely related (e.g., S.
laciniata, S. runcinata, etc.), which have, according to Ledebour,
membranous appendages at the tip of the involucral bracts. S. alata,
although lacking distinct appendages, nevertheless, has been and
undoubtedly should be included under subgenus Theodorea on the basis
of the sum total of characters. The species is variable in the roughness

- and cutting of the leaves, size of the capitula, shape and length of the

528

involucral bracts in relation to each other. The strongly scabrous plants
with deeply pinnatisect leaves and aggregated capitula were separated
by Ledebour as var. B. aspera Ldb. The specimens from Tuva, on the
basis of which B.K. Schischkin described a separate variety, in my view
should be referred to S. alata DC. and not to S. laciniata Ldb. As a
result, a new combination has to be made: S. alata var. pinnatifida
(Schischk.) Lipsch. hoc. loco. S. laciniata var. pinnatifida Schischk. in
Izv. Tomsk. Univ. (1914). Stems more slender, weakly leafy, narrow-
winged or wingless; leaves pinnatifid (their lobes narrower than in the
type specimen) or sinuate-toothed; outer involucral bracts with strongly
recurved tips.

110. S. crepidifolia Turcz. in Bull. Soc. Nat. Mosc. XX, 2 (1847)
47; Ej. Fl. baic.-dahur. II, 1, 120; Popov, Fl. Sr. Sib. II, 756—S.
runcinata B. dentata Ldb. Fl. Ross. II, 2 (1845-1846) 663.—S. alata
DC. y. pygmaea Fisch. ex Herd. in Bull. Soc. Nat. Mosc. XLI, 3 (1868)
46.—S. polypodiifolia Turcz. (non DC.) nomen in herb. and ex Ldb.
op. cit. (1845-1846) 663 and Herd. op. cit. (1868) 46.

523

Perennial. Very similar to S. alata DC. Root collar covered with
remains of leaf sheaths of previous year’s basal leaves partially fibrously
divided. Stem 5—10(15) cm high, sulcate-angular, slightly narrow-winged,
weakly pubescent or almost glabrous. Leaves almost glabrous, weakly
pubescent only along margin and veins, with sessile, lustrous, glandular
hairs on both sides; basal and lower cauline leaves petiolate with
sheathing base, discontinuously pinnatisect, segments usually oblong,
toothed, teeth terminating in cartilaginous cusp, sometimes lamina of
basal leaves almost undivided, roundish or oblong, with teeth along
margin or only in upper part; cauline leaves pinnatifid, with lanceolate,
small-toothed lobes, decurrent, forming narrow wing. Capitula narrowly
oblong, almost cylindrical, 7-10 mm wide, densely aggregated at the
apex of stem and lateral branches; individual corymbs slightly distant
from the main corymb, with two or three capitula. Involucre 10-12 mm
long, almost glabrous, involucral bracts hard, gradually longer from outer
to inner; outer bracts short, gradually narrowed from broad base into
narrow, divergent green apex; inner bracts longer, lanceolate, broadened
at apex into small, membranous toothed, colored appendage. Florets
pink, about 12-13 mm long, narrow part of tube as long or almost as
long as its broad part with lobes of limb; anther appendages sparsely
hairy. Receptacle densely squamose; scales unequal, linear-subulate,
lustrous, about 4—5 mm long; pappus double; outer pappus bristles serrate,
unequal, 4—5 mm long, persistent on achenes, inner bristles long-plumose,
9-10 mm long; achenes about 4 mm long, glabrous, dark brown.
Flowering VII-—VIII. (Plate XXXIX, Fig. 1.)

Saline lands, alkaline areas of sea coast, thickets of chee grass
(Lasiagrostis splendens).—Eastern Siberia: Dauria; Lena-Kolyma (mouth
of Lena, Adams). General distribution: Mongolia. Type specimen of
the species comes from Dauria, is preserved in Leningrad.

Note. Turczaninow considered his species intermediate between S.
alata and S. laciniata; he distinguished it from the former by the shape
of the outer involucral bracts and the absence of indumentum and from
the latter by the less incised leaves. S. crepidifolia Turcz. undoubtedly
is phylogenetically close to S. runcinata DC., from which it differs by
having a narrow-winged stem, broader lobes of the pinnatisect leaves,
outer involucral bracts distinctly shorter than the inner, and other
characters. Popov (op. cit. 756) reported the occurrence of eee
forms between these two species; they are unknown to me.

111. S. runcinata DC. in Ann. Mus. Paris, XVI (1810) 202; Ej.
Prodr. VI, 537; Ldb. Fl. Ross. [I, 2, 663 excl. B. dentata Ldb.; Turcz.
Fl. baic.-dahur. II, 1, 119; Popov. Fl. Sr. Sib. II, 756.—S. alata 6.

529 runcinata Herd. in Bull. Soc. Nat. Mosc. XLI, 3 (1868) 46.—S.:

524

polypodiifolia Turez. (non DC.) pl. exs.; nomen in Ldb. op. cit. 663 p.
p.—S. polypodiifolia Turcz. (non DC.) B. angustifolia Turcz. ex Herd.
op. cit. (1868) 46 and in herb.—Serratula dissecta Steph. ex Herd. op.
cit. (1868) 46 and in herb. nomen.—Heterotrichum dissectum MB. ex
Herd. op. cit. (1868) 46 and in herb. nomen.

Perennial. Similar to S. crepidifolia Turcz. Stem (10)15—25(30) cm
high, basally covered with remains of sheaths of previous year’s leaves,
slender, erect, wingless, almost glabrous, sulcate-angular, branched.
Leaves glabrous; basal, lower and middle cauline leaves pinnatisect with
distinct linear and lanceolate, unequal, acute lobes terminating in
cartilaginous cusp, often recurved, usually entire, less often with few
small teeth, terminal lobe fairly variable in shape, from linear and acute
to broader, subobtuse or acute; basal and lower cauline leaves long-
petiolate with sheathing base; upper leaves linear, narrow, undivided or
small-toothed in lower part. Capitula on more or less long peduncles
aggregated at stem and lateral branch apices in few-headed, dense, and
frequent corymbs, if stem branched, then corymbs forming many-headed
corymbose-paniculate inflorescence. Involucre up to 1 cm wide; outer
involucral bracts herbaceous, linear or linear-lanceolate, narrowed from
ovate base, shorter or as long, sometimes even larger than inner bracts,
not recurved; inner bracts coriaceous, broadened into membranous,
toothed, purple appendage. Florets purple and white (var. albiflora Herd.
op. cit. (1868) 47), about 10 mm long, narrow part of tube almost as
long as broad part including lobes of limb; limb with isolated lustrous
glandular hairs; anther appendages sparsely hairy; florets in individual
capitula up to 14 mm long, with narrow part of tube about 8 mm long,
broad part including lobes of limb about 6 mm long. Receptacle densely
squamose, scales linear-subulate, lustrous, unequal, up to 4-5 mm long;
pappus double; outer pappus bristles serrate or finely plumose, unequal,
up to 5 mm long, persistent on achenes, inner bristles long-plumose,
about 9 mm long; achenes glabrous. Flowering VIII-IX. (Plate XX XIX,
Fig. 2.)

Alkaline meadows, saline steppes, saline lands—Eastern Siberia:
Dauria. General distribution: Mongolia. Type of the species originates
from Siberia, without precise indication of locality (evidently, the
specimens were collected in Dauria). Type in Geneva(?).

Note. A characteristic species, the main distinctive feature of which
is the presence of herbaceous outer involucral bracts. Hybridizes with
S. salsa (Pall.) Spreng.

112. S. laciniata Ldb. Ic. pl. Fl. Ross. I (1829) 16; Ej. Fl. alt. IV,
18; DC. Prodr. VI, 537; Ldb. Fl. Ross. II, 2, 663; Kryl. Fl. Zap. Sib.
XI, 2909; Grubov, Konsp. Fl. Mong. 276.—S. alata DC. &. laciniata

525

530 Herd. in Bull. Soc. Nat.. Mosc. XLI, 3 (1868) 46.—Ie.: Ldb. Ic. pl. FI.

532

Ross. I (1829) tab. 64.

Perennial. Entire plant glaucous or yellowish green. Root collar with
more or less numerous caudices, densely covered with brownish remains
of sheaths of basal leaves; caudices producing floriferous shoots or less
often rosettes of leaves. Stems (5)15—40 cm high, solitary, straight,
sometimes weakly bent, woody, sulcate-angular, winged almost from
base to tip, with toothed wings, terminating in few-headed corymb or
branched almost from base, with branches bearing frequent, dense,
corymbs forming paniculate inflorescence. Leaves with numerous sessile
glandular hairs and scattered, somewhat thick, articulate hairs with stiff
base; basal leaves petiolate with sheathing base, lamina oblong,
bipinnatipartite almost up to base of axis into oblong-ovate or oblong
lobes, further divided up to half or deeper into lanceolate lobules (less
often only toothed), terminating in long, white, cartilaginous cusp, leaf
rachis sometimes toothed; cauline leaves sessile, few, lower leaves like
basal, gradually reduced toward stem tip; uppermost leaves slightly
incised and decurrent, forming toothed wing. Capitula terminal on stem
and branches in dense corymbs. Involucre imbricate; outer involucral
bracts ovate, undivided or toothed, slightly arachnoid-hairy and with
lustrous glandular hairs, broadened at base, decurved and with one or
(when involucral bracts toothed) several cartilaginous cusps; inner bracts
erect, broadly-linear, terminating in small, almost roundish, toothed,
membranous, purple appendage. Florets pale purple, about 12 mm long,
narrow part of tube as long or almost as long as its broad part including
lobes of limb, covered with isolated lustrous glandular hairs; anther
appendages sparsely hairy. Receptacle densely-squamose, scales linear-
subulate, lustrous, unequal, up to 3.5 mm long; pappus double; outer
pappus bristles numerous, unequal, up to 4 mm long, short-plumose,
fragile, but persistent on achenes, inner bristles long-plumose, about 9—
10 mm long. Achenes glabrous. Flowering VII-—VIII.

Alkaline soils, quartzites.—Western Siberia: Upper Tobol
(sporadically in Tselinograd Region), Irtysh (southeast); Central Asia:
Balkash Region. General distribution: Mongolia. Described from
Chingistau Mountains (Semipalati Region). Type in Leningrad.

Note. It produces hybrids with S. salsa (Pall.) Spreng.; see note to
S. paradoxa Lipsch. Occurs sporadically. It varies in the branching of
the stem, which is sometimes simple and unbranched (f. simplex Lipsch.)
and sometimes strongly branched (f. ramosa Lipsch.); the cutting of the
leaves which are pinnate or bipinnate, less often pinnatifid; shape and
depth of incision of the lateral lobes of the leaves; and in the number of
capitula in the inflorescence. Two varieties are recognized.

533

526

1. Var. pygmaea Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XXI (1961) 380.—Plants with numerous caudices; stem slender,
3.5—-7 cm high; leaves pinnatisect, lobes narrow. Capitula solitary or in
threes; involucral bracts appressed, usually not recurved, outer and middle
bracts woolly at apex.—Karaganda Region, Karakalinsk District, vicinity
of the Zhosaly Mountains. “White alkali soil,” 13.VII.1953. L.
Demchenko.

2. Var. latifolia Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XXI (1961) 380.—Stem winged, thick, 10-15 cm high; leaves
pinnatifid, with large, broad, toothed lobes, teeth terminating in
cartilaginous white cusps. Capitula large, in groups of threes in corymbs,
crowded by leaves usually at base—Northern slope of Tarbagatai.
Southern slopes of the Dzhilytau Mountains. 25.VII.1906. V.
Resnichenko.—This variety is transitional to S. alata DC. The
relationship of the latter with S. laciniata Ldb. requires further study. It
is quite possible that these two species will be combined in the future,
because intermediate forms between them are observed.

113. S. ceterahifolia Lipsch. in Byull. Mosc. Obshch. Isp. Prior.,
Otd. Biol. LIX, 6 (1954) 73.—S. runcinata auct. (non DC.);
Sobolevskaya, Konsp. Fl. Tuvy, (1953) 195.

Perennial. Almost stemless, many-headed, canescent; stems dwarf,
1.5—3(4) cm high. Leaves usually longer than capitula, crowded near
root collar, densely covered with brown remains of sheaths of previous
year’s leaves, oblong, short-petiolate, arachnoid hairy-tomentose and with
more or less numerous, lustrous, sessile glandular hairs, lamina pinnatifid
with spreading, toothed lobes, teeth acute, terminating into white
cartilaginous cusp; leaf rachis toothed. Capitula one to three, terminal
on short stem. Involucre 8-10 mm wide, imbricate; outer involucral
bracts more or less villous, almost cymbiform, broadened at base, three-
toothed at apex, teeth with cartilaginous white cusp; inner bracts straight,
broadly linear, terminating in small, almost roundish, membranous, dark
colored (dark red) toothed appendage. Receptacle densely squamose,
scales linear-subulate, lustrous, unequal, up to 3 mm long; florets pink
(or pale purple?) about 10 mm long, narrow part of corolla-tube as long
as broad part including lobes of limb, decorated with individual lustrous
glandular hairs; anther appendages sparsely hairy; pappus double; outer
pappus bristles unequal, up to 4 mm long, serrate, fragile, inner bristles
about 7-8 mm long, long-plumose; achenes about 3 mm long, glabrous,
with indistinct corona at apex. Flowering VII.

Saline places.—Eastern Siberia: Angara-Sayans (Tuva). General
distribution: Mongolia. Described from Tuva. Type in Leningrad.

Plate XXXIX.

Saussurea crepidifolia Turcz.; 2

531

—S. runcinata DC.

1

534

528

Note. The phylogenetic closeness of S. ceterachifolia Lipsch. to S.
laciniata Ldb. is beyond doubt. It differs from the latter by having a
prostrate stem (plants almost stemless), less cutting of the leaves, fewer
capitula, and nonrecurved outer involucral bracts.

Series 4. Robustae Lipsch.—Stem winged throughout length due to
decurrent leaves. Leaves entire in type specimen, less often pinnatilobate
or with few teeth along margin. Teeth of leaves and involucral bracts
without cartilaginous cusp. Florets 16—20 mm long.

114. S. robusta Ldb. Ic. pl. Fl. Ross. I (1829) 16; Ej. Fl. alt. IV,
19; DC. Prodr. VI, 537; Ldb. Fl. Ross. II, 2, 664; Kryl. Fl. Zap. Sib.
XI, 2909; Lipschitz in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XVI, 446.—S. excelsa M. Pop. in Fl. Almat. Gos. Zapoved. (1940) 47
nomen (= S. altissima M. Pop. nomen in herb.).—Ic.: Ldb. (1829) tab.
65.

Biennial, perennial. Plants with vertical root; root collar thick,
covered with remains of previous year’s leaves, producing one or several
stems. Stems strong, (25)40—80 cm high, erect, nbbed-angular, distinctly
winged due to decurrent cauline leaves, wings more or less broad, entire
or toothed, scabrous together with stem due to scattered, somewhat thick,
crisped hairs and sessile glandular hairs, strongly branched from base or
in upper half; lower axillary branches, sterile, usually more or less
numerous. Leaves decurrent throughout stem length, scabrous on both
sides and along margin due to numerous, somewhat thick, crisped hairs
with stiff base, and also covered with glandular hairs; basal leaves
withering before anthesis; lower cauline leaves oblong, middle and upper
leaves lanceolate, acuminate, undivided, or with few teeth along margin,
less often pinnatilobate, 6-10 cm long and 0.7—3.5 cm wide; upper leaves
reduced. Capitula on long peduncles, large, with florets (2)2.5-3 cm
long and 1.5—2 cm wide, aggregated in strong corymbose-paniculate
inflorescence, less often capitula terminal on stem in corymbs if stem
unbranched. Involucre campanulate, imbricate, many-rowed; outer
involucral bracts greenish, linear-lanceolate, pubescent, acute, decurved,
and aristate; middle and inner bracts linear or linear-lanceolate, with
membranous, pink, more or less toothed, apical appendage, densely
white-tomentose, verrucose-villous near transition to appendage; all
involucral bracts with prominent midrib, sometimes covered with
pubescence. Florets lilac purple, about 16-17 mm long, narrow part of
tube about 10 mm long, broad part including lobes of limb about 6—7
mm long; anther appendages sparsely hairy. Receptacle densely
squamose; scales linear-subulate, white, lustrous, unequal, up to 7 mm
long; pappus dual; outer pappus bristles unequal, 3-4 mm long, serrate-

535

529,

scabrous, persistent on achenes, inner bristles long-plumose, equal, about
10-11 mm long; achenes dark brown, thin ribbed, 3.5-4.5 mm long,
glabrous. Flowering VIIJ-IX.

Saline meadows, sands, thickets of reed, chee grass with pea
scrub, bottomlands, alkaline and saline soils—Western Siberia: Irtysh;
Soviet Central Asia: Aralo-Caspian Region, Balkhash Region,
Dzhungaria-Tarbagatai, Tien Shan. Endemic. Described from the
meadows of the Chingistau Mountains (former Semipalati Region). Type
in Leningrad.

115. S. prostrata Winkl. in Tr. Peterb. Bot. Sada, IX, 2 (1886)
518; Lipschitz in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI,
446.—S. robusta Ldb. var. discolor Rgl. and Schmalh. ex O. and B.
Fedtsch. Perech. Rast. Turk. IV (1911) 235 (nomen) and in herb.

Biennial or perennial. Plants with vertical roots; root collar thick,
producing one or two to nine stems. Found as two morphologically
distinct forms, but according to personal observations, growing together;
one stemless, procumbent, often creeping (var. exscapa Winkl. ex
Fedtsch., I.c. (1911) 235, nomen and herb.; Lipsch in Bot. Mat. Gerb.
Bot. Inst. Akad. Nauk SSSR, XXIi (1961) 380 (description), and other
with stem (var. erecta Winkl. ex Fedtsch. op. cit. nomen and herb.;
Lipsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XXI (1961) 381
(descriptio)). Stem in second form ascending or erect, 10-60 cm long,
angular, sulcate, strongly winged due to decurrent cauline leaves, wings
usually narrow, entire or toothed, like stem scabrous due to somewhat
thick, oblong, crisped hairs with stiff base, and with numerous sessile
lustrous glandular hairs; stem branched in upper half, less often simple;
sterile shoots in axils of lower cauline leaves absent or few. Leaves
decurrent throughout stem length; basal leaves rosulate, withering before
anthesis; middle and upper leaves lanceolate or lanceolate-linear,
acuminate; uppermost leaves reduced; all leaves pinnate-toothed, less
often undivided or with fewer teeth, scabrous on both sides and along
margin due to numerous setaceous hairs. Capitula with florets 1.5—2 cm
long and 1—1.5 cm wide, pedunculate and aggregated in corymbose-
paniculate inflorescence. Involucre campanulate, imbricate, outer
involucral bracts greenish, lanceolate, woolly-pubescent, sometimes
glabrescent, acuminate, not recurved or slightly recurved; middle bracts
somewhat villous, with slightly broadened cuspidate dark-colored apical
appendage; inner (together with middle) bracts more elongate, linear,
with membranous, violet-pink roundish, toothed, apical appendage,
broader than bracts, verrucose-villous inside near transition to appendage;
all bracts with distinct, often dark-colored, prominent midrib. Florets
purple, 17—18(20) mm long, narrow part of tube longer than broad part

530

including lobes of limb; lobes of limb narrow-linear, about 5 mm long;
anther appendages hairy. Receptacle densely squamose, scales linear-
subulate, white, lustrous, unequal, about 7-8 mm long; pappus white,
double; outer pappus bristles serrate-scabrous, unequal, up to 5 mm long,
fragile, but persistent on achenes; inner bristles long-plumose, about 12
mm long; achenes dark brown, finely ribbed, glabrous, about 3.5 mm
long. Flowering VII—VIII.

Saline meadows, alkali and saline lands.—Central Asia: Tien Shan.
General distribution: China (Sinkiang, Kuldzha). Described from the
Kokkamyr Plain east of Lake Sairam. Type in Leningrad.

Note. Without much justification, Winkler compared it with S.
laciniata Ldb. The species is quite close to S. robusta Ldb., which it
replaces in Tien Shan.

GENUS 1583. Polytaxis Bge.':?

Bge. in Bot. Zeitung. I, 15 (14 April 1843) 256, descr. abbrev.
germ.; ejusd. Delect. sem. Hort. bot. Dorpat. (30 November 1843) 8;
ejusd. in Linnaea, XVIII (1844) 156; ejusd. Beitr. zur. Kenntn. der.
Flor. Rasslands (1851) 194; ejusd. in Mém. Acad. Pétersb. sav. étr. VII
(1854) 370. .

Capitula homogamous, cupuliform, on long peduncles, aggregated
in lax corymbose-paniculate inflorescence. Involucral bracts appressed,
herbaceous, membranous along margin, imbricate. Receptacle flat,
squamose, scales lustrous, white, flat, linear-lanceolate, acute, slightly
shorter than inner involucral bracts. Florets less numerous, pink or purple,
tubular, cylindrical, narrowed below, two times broader in upper half
and slightly bent, bisexual, deeply five-toothed, zygomorphic (with
variable depth of indentation); teeth linear, acute, somewhat falcate,
sparsely puberulent on outer side. Filaments short, glabrous. Anthers
with appendages; basal appendages plumose like brush, apical oblong,
acuminate, linear, weakly falcate. Style with two short, lanceolate,
subobtuse, deflexed branches, densely pubescent below branches on outer
side with longer, above with short, semiappressed hairs. Achenes with
areole of basal attachment, cylindrical, throughout six-ribbed, dark brown,
darker in furrows, ribs transversely rugose-cristate, pubescent with
scattered, white, soft, long, below bent and above with verrucose hairs,
each rib with cristate margin at apex, ribs continued into flexible incurved

'Treatment by V.P. Botschantzev.
*From the Greek words polys—many, and taxis—series, line; named on the basis of
the many-rowed pappus.

536

537

531

hooks at base of achene, covering areole, achene above ribs with entire,
unequal-toothed, short corona enclosing double, white, persistent pappus.
Outer pappus comprising many rows of sparse scabrous bristles,
outermost bristles very short, inner increasingly longer from periphery
to center. Inner pappus one-rowed, two times as long as outer, connate
at base into tube appressed-hairy on outer side, bristles free and long
plumose above.

Annual succulent herbs, pubescent with capitate glandular hairs on
rather long articulate stalks. Basal leaves numerous, broad- and short-
petiolate, broadly obovate-lanceolate or oblanceolate, acute, undivided
and entire or with small number of sharp and large teeth at apex; cauline
leaves less numerous, smaller, sessile, semiamplexicaul, broadly-
lanceolate, acute, entire. Stem branched, usually from base.

The genus has two species growing in Soviet Central Asia.

Type species: P. lehmannii Bge.

1. Stem short (8-36 cm). Basal leaves entire. Capitula with 3—10-
florets. Florets pink. Involucral bracts densely glandular-hairy on
dorsaltisurfaceicis kta. ok, meee 1. P. lehmannii Bge.

+ Stem taller (15-55 cm). Basal leaves usually with less numerous,
large, sharp teeth. Capitula larger, with 5—20 florets. Florets purple.
Involucral bracts glabrous .............eseeeeees 2. P. winkleri [ljin

1. P. lehmannii Bge. Delect. Sem. Hort. Dorpat. (1843) 8; ejusd.
in Linnaea, XVIII (1844) 156; ejusd. Beitr. zur Kenntn. der. Flor.
Russlands (1851) 194; ejusd. Mém. Acad. Pétersb. sav. étr. VII (1854)
370; Boiss. Fl. or. III, 584; Ind. Kew. fasc. I], 1261 (sub jurinea Cass.
sine combin.); O. and B. Fedtsch. Perech. Rast. Turk. IV, 304; Nikitin
in Fl. Turkm. VII, 214.

Annual. Stem 8-36 cm high, often branched from base, leafy,
together with leaves pubescent with capitate glandular hairs on rather
long articulate stalks. Basal leaves many, 2.5—-8.5 cm long and 1.7—2.8
cm wide, on broad, soft, petioles, broadly obvate-lanceolate or broadly
oblanceolate, acute, entire; cauline leaves less numerous, strongly
reduced, 0.4-0.5 cm long and 0.2—2 cm wide, sessile, semiamplexicaul,
broadly-lanceolate, or lanceolate, acute, entire. Capitula with 3—10-florets,
at anthesis about 23 mm long and 7 mm wide, in lax paniculate-
corymbose inflorescence. Involucral bracts herbaceous, green, pinkish
at apex, with three to seven longitudinal, distinct, dark brown veins and
wide membranous border, pubescent dorsally with numerous capitate
glandular hairs on short articulate stalks; outer bracts throughout, .inner
ones near apex ciliate-fimbriate, outer bracts broadly ovate, obtuse,
lowermost among them about 3.5 mm long and 1.7 mm wide; inner -

532

bracts broadly lanceolate, acute,-uppermost among them about 16 mm
long and 4 mm wide. Receptacular scales about 9 mm long and 0.3-0.6
mm wide. Florets pink, about 18 mm long, in lower part 0.6 mm, in
upper 1.25 mm wide. Achenes (excluding pappus) about 7.5 mm long
and 1.8 mm thick; outer pappus bristles 0.25 mm (outer) to 7.5 mm
(inner) long; inner pappus about 14 mm long, connate at base into tube
up to 3—-5.5 mm high between individual hairs. Flowering IV—VI.

On the outcrops of Tertiary, varicolored, saline crusts in lower hills
and in outliers —Soviet Central Asia: Kyzyl-Kum, Amu-Darya; Pamiro-
Alai (Kugitang Range). Endemic. Described from the vicinity of the
Bakali Well in the Kyzyl-Kum desert. Type in Leningrad.

2. P. winkleri Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
VII, 3 (1937) 52, Fig. 1.—P. lehmannii Bge. var. longipedunculata
Winkl. in O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 304, nom.
nud.—Jurinea polytaxis Winkl. var. longipedunculata Winkl. in O. and
B. Fedtsch. Ibid, in syn.

Annual. Stem 15-55 cm high, usually branched from base, leafy,
together with leaves pubescent with capitate glandular hairs on rather
long articulate stalks. Basal leaves many, 2.5-11 cm long and 1-4.5 cm
wide, on broad and short petioles, broadly obovate-lanceolate or broadly
oblanceolate, acute, with few sharp and large teeth in upper part, less
often entire; cauline leaves less numerous, strongly reduced, 0.3-8 cm
long and 0.1—2.5 cm wide, sessile, semiamplexicaul, broadly lanceolate
or lanceolate, acute, entire. Capitula with 5—20 florets, at anthesis about
27 mm long and 10 mm wide, in very lax paniculate-corymbose
inflorescence. Involucral bracts herbaceous, green on outer side, purple
inside, with three to seven longitudinal indistinct veins and wide
membranous border, dorsally glabrous, outer bracts throughout, inner
ones near apex ciliate-fimbriate; outer bracts broadly ovate, obtuse,
lowermost of them about 3 mm long and 1.75 mm wide; inner bracts
lanceolate, acute, uppermost among them about 18 mm long and 3 mm
wide. Receptacular scales about 12 mm long and 0.3—-0.6 mm wide;
florets purple, about 22 mm long, in lower part 0.75 mm, in upper part
2 mm wide. Achenes (excluding pappus) about 7.8 mm long and 1.8
mm thick; outer pappus bristles 0.25 mm (outer) to 9 mm (inner) long;
inner pappus about 18 mm long, connate at base into tube over 3—5.5
mm high between individual hairs. Flowering IV—V.

On outcrops of Tertiary varicolored saline crusts in lower hills —
Soviet Central Asia: Pamiro-Alai (southern Kazakhstan). Endemic.
Described from the Kyzyl-Tumshuk Mountains in southern Tadzhikistan
(in original description, the mountains mistakenly named “Kzyltysh-
Shup”). Type in Leningrad.

538

333)3)
GENUS 1584. Jurinea Cass.':?

Cass. in Bull. Soc. Phill. Paris (1821) 140.

Capitula with many florets, occasionally with fewer florets. Involucre
usually cupuliform, ovoid or obhemispherical, sometimes almost globose,
less often oblong, imbricate, less often with almost equal, long appressed
bracts, or outer, and often middle bracts, distinctly recurved, inner bracts
always erect, herbaceous or almost coriaceous, more or less arachnoid-
hairy or glabrous, with sessile glandular hairs (less often without them)
and usually with hardly visible appressed bristles above (especially on
inner bracts), along margin, involute (inner) and ciliate. Receptacle flat,
usually with dense glabrous, smooth and acuminate scales. Florets
homogamous, bisexual, all fertile; corolla regular or slightly obliquely
five-fid, always narrowed toward base, pink, purple or dark purple, in
exceptional cases cream-colored, almost always with sessile glandular
hairs, glabrous; anthers glabrous, hastate, syngenesious; apical appendages
of anthers narrow hastate, basal appendages setaceous, membranous,
caudate, mostly divided or split along length, often flexuous, filaments
free, glabrous and smooth, occasionally somewhat tuberculate, but not
hairy, inserted at point of narrowing in species with abruptly narrowed
tube, and in lower part in species with gradually narrowed tube; style
glabrous, stigma bifid almost up to base and with tuft of short bristles
at base, branches of stigma arcuate, covered with short papillae, mostly
slightly grooved above. Pappus many-rowed, bristles unequal, two to
five inner larger than others, bristles serrate, barbate, or plumose,
innermost two to five bristles supporting pappogenus, broader and shorter-
barbate, connate into ring (in species with deciduous pappus), enclosing
pappogenus developing on membranous bottom, and with nectary at
tip; pappogenus together with pappus detaching from achene, detaching
from pappophorus, or pappus bristles firmly attached to achene, and
then pappophores absent and nectary developing on pappogenus itself.
Achenes obpyramidal, four-angled, less often somewhat compressed,
more or less oblong, with straight or slightly oblique areole, structurally
more simple than in Serratula, glabrous along ridges, smooth,
longitudinally striate or smooth, or transversely tuberculate-alveolate with
folds, rugose above with bands of teeth below corona, often with sessile
glandular hairs. Perennial herbs or semishrubs: often stemless or scape-
like; leaves simple and pinnatipartite.

Genus widely distributed in Eurasia from the Atlantic up to eastern
Siberia, in Western and Central Asia, and partly in Soviet Central Asia.

'Treatment by M.M. Iljin.
2Named after Louis Jurine, Professor of Medicine in Geneva at the end of 18th century.

534

It comprises about 300 species, of which more than 150 species are
found in the USSR.

Note. The diagnosis of the genus and its sections includes some
morphological structures that have been mentioned already in brief. In
species with a deciduous pappus, a conical or dome-shaped projection,
the so-called pappophorus, develops at the tip of the achene, from which
the pappus easily detaches at the maturation of the fruit (Plate XL). The
pappophorus does not develop in species with a persistent pappus, whose
bristles are firmly appressed to the achene by the pappogenus and the
nectary is shifted to the center of the pappogenus. The nectary usually
sits on a more or less short pedicel, the so-called manubrium. The bristles
supporting the pappogenus (2-5) and projecting above the others have a
unique structure. These bristles are always sturdier, broader and flatter;
they may differ sharply from all the other bristles, as for example, in
Jurinea xeranthemoides Ijin, which recalls the genus Xeranthemum. The
inner subtending bristles in this species are narrowly ensiform, hardly
serrate and with numerous fine hairs along the border; the rest are short-
barbate, with the barbules bilateral.

Type species: J. alata Cass.

KEY TO SECTIONS

1. Corolla gradually narrowed toward base, peripheral corollas abruptly

pendulouspved...cterhe..ceesben...eaarae .Seeee.. 2m. maeeer. we 2.
+A iitcorollasynevers pendulous e-crsseee. 2h pet e222. 228 Cee. teasers 3.
2. Involucre cylindrical or narrow-cupuliform; all leaves linear, entire,
sessiletiw.. toed... eeteeck. suo eechertaet ,.t6e Section 8. Olgaeae Iljin

+ Involucre cupuliform; leaves pinnatipartite or lyrately parted, often
entire, but then oblong-lanceolate or oval; cauline leaves decurrent
(ck, however ucephalopodag lyin) bere... tiserace5..2o-eteeheeeneesteee aoe:
Hits. Jediatinn. Aes. £35). .0eee Section 7. Penduliflorae [ljin

3. Plants thistle-like, exclusively Caucasian; pappus entirely deciduous,
pappophorus developed, marginal ribs of achene white-cartilaginous
Sloe lk Stakes wlogset/nettiet. 34. .cecneeee. Section 14. Jurinea.

4. Achenes with longitudinal, discontinuously alveolate ribs ........ 5
+ Achenes glabrous, smooth, scabrous along length or cylindrical,
sometimes with band of teeth or barbules below corona.......... 6.

5. Ribs of achenes only transversely discontinuous, often modified
into teeth above, less often into bristles, connate below; pappus
persistent on achene or weakly connate with it, less often entirely

540

10.

11.

14.

deciduous. Kazakhstan-Soviet Central Asian sections ...................
ee oN AR Ee cr ee Section 15. Chaetocarpae (Korsh.) Ijin
Ribs of achene sharply scabrous, so that pits alternate with cochleate
tubercles, teeth pointing upward, always densely glandular; pappus
always deciduous, pappophorus persistent. European-Caucasian

Section iw».).). dedeo).ceno..duces Section 13. Molles (Korsh.) Iljin
. Section from the European Part of the USSR, Caucasus and
SRY E) CUR 1 1 ors 8 0 cen ee ne a es ee Ti
sections from. Sovieh:CentralAsia.l:amwe.. eeunrien.caveek:. 11.
. Plants with scapelike stem; pappus not deciduous; achenes small;
Gam Cassis, Memento. sclera denciiten Bt Section 2. Bellae I[ljin
Sfemsalwiays leafy ses cect n:.wactet cttamenct. geanerhw ines. catedenleasem mes eg 8.

. Achenes regularly broadly four-angled, narrowed below; Pappus

not deciduous; European Part of the USSR, Northern Caucasus,
Western” Sibena!/ Kazakhstan! )0s, cot ee OE a REN BEE
PASTE d 83) ARS cst act a Section 1. Cyanoides (Korsh.) Iljin
Achenes subobtuse below, narrower ...........ccc0cccccccseeesssccseeeceesseees 9.

. Capitula cylindrical or narrow cupuliform, numerous in corymbose

inflorescence; pappus persistent, with serrate or serrate-barbed
bristles. European Part of the USSR, Caucasus, Western Siberia,
Kazakhstan .............. Section 3. Stenocephalae Benth. and Hook.
Capitula solitary, terminal on main stem and branches; pappus
décidhous” Catcastisgte- el, JBOMAaIN GOO, COON e) eae ee 10.
Plants with virgate branches; leaves pinnatisect or sinuate-toothed;
pappus barbed....................c Section 6. Floccosae (Sosn.) [lyin
Plants weakly branched, but not virgate, leaves undivided and
entire; pappus plumose (only species J. auscheriana DC.) ..........
Vt DeOIViD gay 8 Section 10. Derderia (Jaub. and Spach) Boiss.
Corolla gradually narrowed below, usually 27-35 mm long,
sometimes at least 25 mm; pappus plumose or barbate-plumose.

Corolla less than 25 mm long (cf., however, J. maxima Winkl.);
pappus serrate or barbate (cf. however, J. propinqua Iljin, J.
macranthodia Iljin, J. kokanica \ljin and J. serratuloides Ijin) ..

. Leaves auriculate; pappus always persistent on achene.................

He Nok: eens SANE af Section 10. Derderia (Jaub. and Spach) Boiss.
Leaves not auriculate; pappus always deciduous ................::c0:000
ESE NL ON a neat PINE Mas ee Section 11. Stechmannia Boiss.

. Leaves linear and entire or with linear lateral lobes, in exceptional

CASS MME Wig LOWS FOLAO Valente. ay crete Pee orcs encssntenense se hese Urn 14.
Leaves variable, but always broader, not linear....................- 15s
Pulvinate semishrubs, with sessile capitula, or plants with short

541

536

stem, less often up to 25 cm high, pappus always deciduous......
ELA RBA, 8 Jc EOS. osc Section 12. Nanae Iljin
+ Semishrubs of usual type, with linear always undivided and entire
leaves; pappus always persistent on achenes............0..:c:ceceeeeees
ULE oto RELEcE. n ARCOM Nos thes Med PA Ok Section 5. Suffrutices [jin
15. Leaves undivided, in exceptional case forked (VJ. trifurcata I\jin),
usually decurrent with undivided wing. Tien Shan.................0.
til aonlemandebMebehMeninct eee’ Bao lt asd cuts Aly he Section 4. Integrae Iljin
+ Leaves variously incised, decurrent, row of serrate appendages
(fimbria) or toothed wing (Pamiro-Alai Mountain System, less often
SURV HAT) ICI. ce Beccerecvnsdssueumeechwaveces Section 9. Insulptae Iljin

Section 1. Cyanoides (Korsh.) Iljin in Addenda XXVI, 715.—Stirps
Cyanoides Korsh. in Izv. Akad. Nauk 1, 2 (1894) 8; Iljin in Izv. Bot.
Sada Akad. Nauk SSSR, XXX, 34, 348; Group Cyanoides Klok. in
Bot. Zhurn. Akad. Nauk URSR, VIII, 1, 47.—Sect. §2. Longifolia DC.
Prodr. VI (1837) 674 p. p—§3. Deccurentes DC. Ibid. 675 p. p.—§4.
Pinnatilobeae DC. Ibid. 675 p. p.—Platycephalae Benth. and Hook.
Gen. pl. II, 1 (1873) 473 p. p—§ Pinnatae Boiss. Fl. or. II] (1875) 572
p. p.; Pflanzenfam. IV, 5, 321 p. p.—Sect. Platycephalae Benth. and
Hook. subs. Cyanoides Sosn. in Zhurn. Russk. Bot. Obshch. 11, 1-2
(1926) 196.—Pappus bristles not always connate into ring, serrate or
serrate-barbate. Achenes obpyramidal, four-angled, narrowed toward base,
glabrous, almost smooth or with longitudinal parallel stripes, simply
serrate above, not forming true corona, with pappogenus of different
shape in center at apex, pappus bristles appressed below it; pappophorus
undeveloped, pappogenus terminating in five-toothed nectary. Plants
always perennial with leafy stem, often winged; leaves divided to various
degree, less often entire, almost always gray or white-tomentose beneath.
This section comprises about 25 species; 22 of them occur in the USSR.

Type of section: J. cyanoides (L.) DC.

1. Stem wingless; leaves nondecurrent ...............- 5 eieat oa Bec Bae re Ds:
+ Stem winged; leaves decurrent in wing or border ..................... 2.
2. Outer and middle involucral bracts sharply decurved................ 3.
+ Outer and middle involucral bracts squarrose, slightly entangled

(rutiled) or all appressed ye sey ccc cee sas ean eee seers access eeeeseceenses 6.

3. Capitula medium; involucre always strongly arachnoid-hairy or
white-tomentose, decurved part of involucral bracts purple or dark
green, weakly arachnoid-hairy like base of inner bracts; leaves
stiffly scabrous, usually bi- or almost tripinnatisect into narrow
linear or almost filiform segments; cauline leaves sessile,
semiamplexicaul with well-developed auricles at base (Donbass)
Zu ae R Senet cele Nas MCC NUON ooiteh ORE hae Cae nas 7. J. granitica Klok.

542

544

33.7

Involucre glabrous, arachnoid-hairy or white-tomentose; leaves
pinnatisect or bipinnatisect, less often mixture of undivided,
scabrous or glabrous, with or without auricles (European Part,
Westernrsiberia:y Kazakiastam))/ ees iin... acc I AB... 4,

. Capitula small, less often medium; involucre white-tomentose; outer

and middle involucral bracts appressed, only slightly and short-
uncately exserted from tomentum, dark green or violet-purpie at

apex, (Dniepen Repion)avtiess Sain: 8. J. salicifolia Grun.
Capitula large, outer and middle involucral bracts long and deflexed
ye rer ot Roteus. neteoels meets Piraeus sinew | peli Or Wit... >.

. Leaves stiffly scabrous above due to fine cartilaginous Supillae,

semiamplexicaul, usually with distinct auricles; involucre usually
glabrous, less often arachnoid-hairy (plants of semidesert and
southern, steppes))<:..stsist..21.. Hii geess...taadl 5. J. eversmanni Bge.
Leaves smooth above, less often weakly scabrous, usually without,
less often with poorly developed auricles; involucre usually
arachnoid-hairy, less often almost glabrous (plants of forest-steppe
andenorthern, Steppes)............-...csescesecees 6. J. charcoviensis Klok.

. All involucral bracts appressed (eastern part of Western Siberia

and Northeastern Kazakhstan)............. 3. J. schischkiniana [\jin.
Outer and often also middle involucral bracts separated to various
CPST ET S| SINT AMR | GRAIN ee fa lo cela a Pe Rs EO ee q

. Stem usually flexuous and strongly branched above base; leaves

more or less coriaceous, green on both sides or slightly arachnoid-
hairy beneath. Capitula medium or small; outer involucral bracts
separate or almost appressed (central Kazakhstan) .......................
ete Ra See tees: lla ene RR NE 4. J. xerophytica Ijin
Stem upright, simple or weakly branched, usually with fewer
branches above middle or only in upper part, spreading upward;
leaves thin, herbaceous, white-tomentose or gray arachnoid-hairy
beneath. Capitula large or medium; outer and middle involucral
bractspasiif yruffled «sd seed. aliens elas ee 8.

. Basal leaves gradually narrowed toward base, pinnatisect, sessile

or with poorly developed petiole, smooth or arachnoid-hairy above;
cauline leaves except lowermost, sessile, semiamplexicaul and with
well developed auricles; involucre always arachnoid-hairy
(Belorussia, Lithuania, Western Ukraine) ................c:cccsccesseeeseeeees
simlsttennsthenmitem caste collab bal. bey 2. J. pseudocyanoides Klok.
Basal and lower cauline leaves with well- -developed petiole and
pinnatisect or bipinnatisect lamina, scabrous above; cauline leaves
not amplexicaul and without auricles.......0.......:csccseccesecenecereeeeeeeeees
nddncopaterth Are wows aks, etic lens len ass ome Apacs, 1. J. cyanoides (L.) Rchb.

. Plants robust, stem mostly thick, 3-7 mm; thick at base; basal

545

538

12

13:

14.

15.

16.

leaves 20-30 cm long; cauline leaves long- and broadly decurrent
(sands of Crimea, Dnieper and Bug rivers)................c:ccc0cee 10.
Plants less robust, stem thinner; leaves smaller ....................... lis

. Leaves undivided, entire, lanceolate-oblong; lower and middle

involucral bracts somewhat ruffled, like J. cyanoides (Aleshi sands)
» AER: SU RRE hat, nRRO RES 12. J. laxa Fisch. ex Ijin
Leaves pinnatisect; lower and middle involucral bracts sharply
deflexed (sands between lower reaches of the Dnieper and southern
Bip) magnet ead WAe eas SOE tae 19. J. paczoskiana [lyin

. All involucral bracts appressed, from shorter outer to more elongate

innenuGmbricate):.224..0)-Rd. BY OAR POTORON Ve! ee 12.
Outer, and often also middle involucral bracts ruffled to various
degrecivixe. Vir RAS) | VAN ene Aa oot. eeI Ole 13.
Stem always branched, usually in upper one-third or even upper
half. Capitula usually 15-25; involucre at anthesis almost
cupuliform; inner involucral bracts long and white-membranous,
acuminate, distinctly exserted (valley of the Don and its tributaries)
LPAPREDE EROSION. CED a cveadeanstaie tassesenexasede Aum 21. J. tanaitica Klok.
Stem simple or branched only at extreme tip. Capitula usually
fewer; involucre cupuliform, inner bracts although membranous
above, but attenuate into acuminate tip (Trans-Caspian deserts)...
eau wohaslsa saa mriedoc ncaa edsamaautediancnanesinaeheccadderads 20. J. tenuiloba Bge.
Outer and middle involucral bracts squarrose and as if ruffled (like
J. cyanoides); leaves quite narrowly decurrent (cf. however, J.

transuralensis {jin and J. pineticola Iljin) ...............c0ccccccceeee 14.
Outer and middle involucral bracts variously deflexed, but not like
VMCVOROGES 30.290... soacassrrnaeedosa PRs iaseCINeeENnte tii oRens ON IINGRIS DERN ES RES, 16.

Plants strongly white-tomentose; leaves undivided, linear, or
pinnatisect, light green above. Capitula large or medium; involucre
also white-tomentose (Baksan, Zaisan) ....... 9. J. albicaulis Bge.
Plants not white-tomentose, finely arachnoid-hairy, only lower leaf
surface whitish arachnoid-hairy; involucre weakly arachnoid-hairy
EA MORRILL SOR, AR DOV OTIRE: MIPIM ROVE MERE, £5.
Leaves usually always undivided, occasionally with mixture of
weakly divided laminas (Kurgan and Yalutorov districts) ............
ORR RET, ORME STROVE. SROs 10. J. gorodkovii Iljin
Leaves except upper, always finely pinnatisect in linear-filiform
segments (chalky deposits of Middle Volga Region) ...................
HIN. PRR OUVODALI OYE AW PAV BOS OF ERD, 11. J. creticola [jin
Outer and middle involucral bracts in upper half sharply deflexed
OF. AL CUALE. ..os.conianeioionivowcte err seberddenet eee eda eons vate AbDRGOR eee Mas aaa acaeet a lv:
Outer and middle involucral bracts slightly deflexed from involucre
orsonly fat apex shortly -uncatets...U70.0.. A.. aces See. 9.

543

|
Plate XL.

Structural details of pappus in Jurinea species. 1, 2—J. arachnoidea Bge., pappogenus
with “manubrium” and nectary in dorsal view; 3, 4, S—J. armeniaca Sosn. pappus base,
pappogenus and pappophorus; 6, 7—J. poacea Iljin, pappogenus with nectary, pappophorus;
8—J. psammophila Iljin, schematic section through upper part of achene with pappogenus
and nectary; 9, 10—J. olgae Rgl. and Schm., pappogenus with nectary, pappophorus; 11—
J. cyanoides (L.) DC., schematic section through upper part of achene; 12—J. rhizomatoidea
Iljin, pappogenus with nectary; 13, 14, 15, 16—J. tapetodes Iljin, pappophorus, in lower,

upper and lateral view, and upper part of achene with dentate border and pappophorus.

540

20.

Die

. Outer and middle involucral bracts sharply deflexed .............. 18.

Outer and middle involucral bracts arcuate in different directions
(western Kazakhstan) ............ csc eeeeeeees 15. J. transuralensis Iljin

. Deflexed part of involucral bracts glabrous or subglabrous, dark

violet or dark green (semideserts and southern steppes of the
European Part and bordering Kazakhstan) ..................sceseeseeeseeees
Sethosetiegs idee acca dou i Bbais 13. J. amplexicaulis (S.G. Gmel.) Bobr.
Deflexed part of involucre always gray arachnoid-hairy (Altai
Termitory;/Loktey Distt) 0 :..ii2%.cSh-<e-e 14. J. altaica [lin

. Outer and middle involucral bracts finely arachnoid-hairy, not

uncate but slightly deflexed at apex, especially at fruiting,
sometimes with membranous somewhat more deflexed tip (Ukraine)
bth a tae A ocd, socabatnst sake oataac cutee! oh cee. tase ot: 22. J. thyrsiflora Klok.
Plants with different characters (Ciscaucasia or Kazakhstan) .......

Involucre cupuliform or nafrow-cupuliform, narrow or roundish at
base, but in any case not depressed when attached to stem, always
white-tomentose; outer and middle involucral bracts with short
dark-purple or dark green uncate tip, exserted from tomentum
(Aktyubinsk and northern Kyzyl-Orda regions)...............:::ceceee
bos RIGOR ado a ae 17. J. kazachstanica [lyin
Involucre broadly cupuliform or cupuliform, depressed at base when
attached to stem, usually arachnoid-hairy, less often almost white-
tomentose (plants of Ciscaucasia and eastern Kazakhstan) ..... 21.
Involucre finely arachnoid-hairy, broadly cupuliform, always with
distinct depression at base when attached to stem; outer and middle
involucral bracts short-uncate at tip, arachnoid-hairy, not purple
(CiscaicaSia) ss. iciks-dur.escr. eae 16. J. ciscaucasica (Sosn.) Iljin
Involucre as high cup with weak depression at attachment to stem;
outer and middle involucral bracts not shortly uncate at tip, but
originating from involucre sometimes as arcuate tips, sometimes
slightly drooping, sometimes almost appressed. Plants with
decurrent or nondecurrent leaves (eastern Kazakhstan).................
PLAN i er: Se ee eT Oe Sear Bike hee enn ge 18. J. pineticola [ljin

Series 1. Cyanoides Iljin—Eucyanoides Klok. in Bot. Zhurn. Akad.

Nauk URSR, VII, 1 (1951) 48 p. p—Stem wingless.

Subseries |. Intricatae [ljin. Ibid——AlII involucral bracts appressed

or loosely ruffled.

546

1. J. cyanoides (L.). Rchb. Fl. Exc. (1831) 290; DC. Prodr. VI,

676; Boiss. Fl. or. III, 573 p. p.; Grossh. Fl. Kavk. IV, 172; Schmalh.

541

Fl. II, 110 p. p.; Fl. Yugo-Vost. VI, 390; Kryl. Fl. Zap. Sib. XI, 2931 p.
p.; Vizn. Rosl. URSR, 563, s. str.; O. and B. Fedtsch. Perech. Rast.
Turk. IV, 300.—Carduus cyanoides a. monoclonos L. Sp. pi. (1753)
822; MB. FI. taur.-cauc. II (1808) 272; Korsh. in Izv. Akad. Nauk, 1, 2,
117 p. p.; Tent. Fl. Ross. or., 251; Sosn. in Zhurn. Russk. Bot. Obshch.
11, 1-2, 196; Iljin, Tr. Turk. Nauchn. Obshch. 2, 20.—Serratula
cyanoides DC. Diss. comp. III (1813) 30; MB. FI. taur.-cauc. III, 550.—
Acarna cyanoides Bess. Enum. pl. Volh. (1822) 32.

Perennial. Plants 25—60(70) cm high; stem erect, simple sulcate,
whitish due to more or less dense arachnoid pubescence, especially in
lower part and on petioles of lower leaves, less often greenish, with
weak arachnoid-hairy coating, often branched, usually with few long
or more or less short branches. Basal and lower cauline leaves distinctly
petiolate, not decurrent, pinnatisect in 2—20 pairs of oblong or even
linear segments, sometimes bipinnatisect (f. bipinnata \ljin) or almost
lyrate-divided, less often all or only lower leaves undivided (f.
integrifolia Iljin), latter often long (f. /ongifolia Iljin) with decurrent
lobes, gradually acuminate upward or subobtuse; uppermost leaves
entire, linear, sessile, not reaching capitulum; all leaves very finely
arachnoid-hairy above, especially along veins, scabrous due to rather
small papillae, slightly rugose, with sparse, sessile, glandular hairs,
whitish beneath due to arachnoid pubescence. Capitula medium or
large, solitary, with many florets. Involucre broadly cupuliform, with
depression at base, 15—18(20) mm long, outer bracts linear-lanceolate,
all bracts almost equal in length, acuminate, lax, somewhat ruffled
and weakly deflexed, herbaceous, usually significantly arachnoid-hairy,
usually three-rowed; inner bracts one- or two-rowed, straight, acuminate,
only at extreme tip slightly recurved, reddish, less often yellowish,
with short bristles, ciliate and with scattered sessile glandular hairs.
Receptacle covered with glabrous and smooth scales, scales fused below,
reaching one-third height of involucre or slightly more. Corolla pink-
purple, 16-20 mm long, narrowed over 7-8 mm, with lobes of limb
4—6 mm long, with sessile glandular hairs; apical appendages of anthers
acuminate, about 2.5 mm long, basal appendages 2—2.5 mm long;
stigma 2 mm long; style glabrous; filaments glabrous. Pappus whitish,
divaricate, two to two and one-half times as long as achene, comprising
serrate-barbate bristles, not connate into ring; achenes obpyramidai,
four-angled, distinctly acuminate below, glabrous, smooth, with
longitudinal parallel stripes, somewhat serrate above, and with patellate
pappogenus at apex. Flowering VI-—VIII; fruiting VII-IX. (Plate XL,
Fig: 11.)

In open and dry pine forests, pine-forest sands, sandy steppes, chee

547 grass-sheep’s fescue and chee-grass-covered, gravelly stony steppes, on

542

limestone deposits and chalky slopes.—European Part: Upper Dnieper,
Upper Volga; Middle Dnieper; Volga-Don; Volga-Kama; Bessarabia,
Lower Don, Trans-Volga Region, Lower Volga; Western Siberia: Ob,
Upper Tobol, Irtysh, Altai; Caucasus: Ciscaucasia. Endemic? Described
from the Belgorod District and Crimea according to the report of Gerber.
Type in London.

Note. The closely related species, J. pollichii Koch. (1837) = J.
cyanoides f. pollichiana DC., grows in Central Evrope, which is
well distinguished from J. cyanoides (L.) DC. by having predominantly
simple stems and almost sessile leaves with usually linear segments
gradually decurrent to their base. Besides, both the lamina and the
narrowly winged rachis of the basal leaves are usually circinate, not
scabrous above.

2. J. pseudocyanoides Klok. in Vizn. Rosl. URSR (1950) 564
(nomen); Klok. in Bot. Zhurn. URSR, VIII, 1, 51.—J. pollichii Ldb. FI.
Ross. II (1844-1846) 764 specim. lithuan. and podolic. non DC.—J/.
cyanoides Schmalh. Fl. IH (1897) 110, specim. bjeloruss. non DC.;
Fedtsch. and Flerow Fl. 1005 specim. occid. and polon.—Exs.: Fl. polon.
exs.: No. 189.—Ie.: Klok. op. cit. (Bot. Zhurn.) Fig. 8.

Perennial. Plants 27-75 cm high; stems solitary or few, straight,
often somewhat thick, flocculose-arachnoid-hairy with longitudinally
striate, simple or less often weakly branched in upper half. Basal leaves
numerous, with poorly developed petiole or sessile, lamina often
undivided (f. integrifolia Iljin) or oblong, gradually decurrent toward
base, pinnatipartite or pinnatisect, with two to eight lateral segments on
each side, oblong or oblong-lanceolate, like undivided leaf acute or
cuspidate, less often acuminate, also decurrent toward base; cauline
leaves, except lowermost like basal, sessile, undivided or simple if
divided, amplexicaul or auriculate, less often without one or another,
all leaves weakly arachnoid-hairy above, smooth, with scattered sessile
glandular hairs, white-tomentose beneath, with prominent midrib and
lateral veins and numerous glands under tomentum. Capitula large,
usually few, with many florets. Involucre always arachnoid-hairy,
cupuliform, 10-18 mm long and 15—25(30) mm wide; outer involucral
bracts narrow, linear, irregularly deflexed in upper half (divaricate as in
J. cyanoides); inner bracts erect, narrowly lanceolate, acuminate, not
arachnoid-hairy, with sessile glandular hairs and fine appressed bristles,
ciliolate above, white-membranous and pubescent at tip. Corolla pink
or pink-purple, with sessile glandular hairs, 15-19 mm long, narrowed
over 6—8 mm length, with lobes of limb (4)5—6 mm long. Pappus snow-
white, 10-16 mm long, pappus bristles not connate into ring, serrate-
barbate, with barbs equaling thickness of hairs or shorter. Achenes 4—5

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543

mm long, oblong, obpyramidal, four-angled, subacute, glabrous, smooth,
whitish, weakly striate along length, serrate above, and with a pitcher-
shaped pappogenus in center of tip. Flowering VI-VIII; fruiting VIII-
IX.

On forest sands.—European Part: Bessarbia, Middle Dnieper, Upper
Dnieper. General distribution: Possibly grows in Central Europe.
Described from the region of the village of Zvonkovyi, Vasilkov District,
Kiev Region. Type in Kiev.

3. J. schischkiniana I[ljin sp. n. in Addenda XXVI, 716.—J.
cyanoides Schischk. (p. p.; excl. specim. europ. USSR and Asia media,
Kazah. occid) in Fl. Zap. Sib. XI (1949) 2931 non DC.; Ijin in Tr.
Turk. Nauchn. Obshch. 2, 20 p. p.—J. pollichii Ldb. FI. Ross. II (1844—
1846) 764 p. p. non DC.—Serratula cyanoides Bge. in Ldb. FI. alt. IV
(1833) 42 p. p. non DC.—J. albicaulis x J. cyanoides Schischk. op. cit.
2933.—Ie.: Ijin, op. cit. tabl.

Perennial. Plants 30-75 cm high; stems solitary or few, straight,
green or dirty purple, longitudinally striate, simple or sparsely branched
above middle, less often below, flocculose- or finely arachnoid-hairy,
often white-tomentose below. Basal and lower cauline leaves petiolate,
with white-tomentose, usually semicylindrical petioles, pinnatisect in
oblong-lanceolate or linear, acuminate or acute, entire, less often scarcely
toothed segments, decurrent along rachis, less often all entire; other
cauline leaves sessile, not decurrent, occasionally auriculate, uppermost
leaves simple, linear or even filiform; all leaves green and finely-
arachnoid-hairy above, finely rugose, covered with fine sessile glandular
hairs, often narrowly revolute, white-tomentose beneath and with
numerous sessile glandular hairs under tomentum. Capitula few (one to
five), with many florets; florets 2—3.5 cm long. Involucre 15-25 mm
long, broadly cupuliform, always distinctly arachnoid-hairy, with
numerous bracts, almost equal, linear acute; outer bracts appressed, firmly
enclosing involucre, slightly above yellowish, coriaceous short base
terminating into herbaceous apex, not deflexed, inner row of bracts erect
even with cusps, slightly curved only at apex, densely covered with
sessile glandular hairs and short appressed bristles, somewhat ciliate
and pubescent at apex. Corolla pink-purple, with scattered sessile
glandular hairs, 14-21 mm long, narrowed over 8-10 mm length, with
lobes of limb 5—7 mm long. Pappus snow-white, 12-15 mm long, pappus
bristles not connate into ring, serrate-barbate; achenes oblong, 5—6
mm long, four-angled, obpyramidal, glabrous, smooth with prominent
longitudinal stripes along angles, toothed above, teeth usually slightly
bent, pappogenus in center of tip. Flowering V—VIII; fruiting VII-
IX.

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544

Dunes in river valleys, hummocky sands; pine forests on sands, in
dryland meadows with loamy soil, in scrubs and chee-grass-covered
steppes on loamy soil mostly in forest-steppe zones——Western Siberia:
Irtysh, Altai(?); Soviet Central Asia: Balkhash Region. General
distribution: Possibly, in Dzhungaria-Kashgaria. Described from the
Barnaul District, between Solonovka on Kasmal and the Malyshevyi
Ravine. Type in Leningrad.

Note. The range of J. schischkiniana I\jin can be demarcated as
follows: in the north the border is at 54°N, in the west at 70°E; in the
east the range borders with the mountain-taiga region, and in the south
with the semidesert zone of Kazakhstan. The phylogenetically close
species J. cyanoides (L.) DC. occurs west of this range.

4. J. xerophytica [ljin sp. n. in Addenda XXVI,. 716.

Perennial. Plants 20-60 cm high, with long or short, often branched,
rhizome covered with dark brown scales, with woolly axillary buds.
Stems solitary or numerous, like petioles of lower leaves, densely white-
tomentose at base, flocculose-arachnoid-hairy above or almost glabrous
(f. glabrescens Iljin), finely striate-sulcate, streaks of yellowish or purple,
furrows greenish, usually more or less flexuous or straight, branched
from base or somewhat above, sometimes only in upper part. Leaves
somewhat stiff; basal and lower cauline leaves petiolate, pinnatipartite
or pinnatisect; middle and upper leaves sessile, segments oblong or linear,
acuminate, entire, sometimes lyrate, occasionally segments with lobes
or teeth at base, often mixed with entire and undivided leaves; uppermost
leaves reduced and undivided; all leaves greenish above, finely arachnoid-
hairy or glabrous, somewhat rugose, scabrous, with sessile glandular
hairs, narrowly involute, sometimes up to midrib, white-tomentose
beneath, glandular under tomentum. Capitula with many florets, 15—25
cm wide. Involucre 10-15 mm long, more or less arachnoid-hairy, with
outer bracts like J. cyanoides. Corolla pink-purple, sparsely glandular,
(11—12)16—20 mm long, narrowed over (4)6—9 mm and with (4)5—6 mm
long lobes of limb. Pappus snow-white, (10)12—15 mm long, with short-
barbate bristles, more or less dense; achenes 4—5(6) mm long, rusty,
dull, longitudinally striate, small-toothed above, and with apical
pappogenus.

In semideserts.—Soviet Central Asia: Aralo-Caspian Region, Balkash
Region. Endemic. Described from the Altai River, in the Sarysu River
system. Type in Leningrad.

Subseries 2. Reclinatae Iljin—Series Eucyanoides Klok. in Bot.
Zhurn. Akad. Nauk URSR, VIII, 1 (1951) 48 nom. nud. p. p.—Involucral
bracts deflexed.

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545

5. J. eversmannii Bge. in Flora, XXIV, 1 (1841) 155; Ldb. FI.
Ross. II, 2, 765 (excl. specim. ural.); Schmalh. Fl. I, 110 p. p.; Ijin Fl.
Yugo-Vost. VI, 391 p. p.; Grossh. Fl. Kavk. IV, 173; O. and B. Fedtsch.
Perech. Rast. Turk. IV, 301; Korzh. in Izv. Akad. Nauk. I, 1, 118 p. p.;
Sosn. in Zhurn. Russk. Bot. Obshch. 11, 1—2, 196; Iljin in Tr. Turk.
Nauchn. Obshch. 2, 20 p. p.—Ic.: Fl. Yugo-Vost. Fig. 699.

Perennial. Plants 20-75 cm high; stems solitary or few, erect, finely
striate-lineate, usually not strongly arachnoid-hairy, green or dirty purple,
simple or branched in upper half. Basal and lower cauline leaves
petiolate, petioles white-tomentose, shorter than lamina, leaves reduced
upward, pinnatisect (f. pinnata Iljin) or bipinnatisect (f. bipinnata I\jin),
less often undivided (f. integrifolia Iljin), segments oblong-lanceolate,
lanceolate-linear or even narrow-linear, decurrent along rachis, entire or
with occasional teeth on lobes, acute, small up to filiform in upper half
of stem, simple, undivided and entire; all leaves, except lowermost,
sessile, auriculate, semiamplexicaul, green above, somewhat stiffly
scabrous, glabrous or with fine tomentum, especially along midrib,
narrowly involute, white-tomentose beneath, with scattered sessile
glandular hairs under tomentum. Capitula solitary, medium or large,
with many florets. Involucre cupuliform, 5—20 mm long, glabrous or
arachnoid-hairy; bracts lanceolate-linear or linear, acuminate, equal, outer
almost coriaceous, strongly recurved, dark green or purple; inner bracts
glabrous, straight. Receptacle densely covered with smooth scales, more
than half as long as involucre. Corolla pink-purple, with sessile glandular
hairs, 16-20 mm, narrowed over 6-8 mm, with 4.5—6 mm long lobes of
limb. Pappus yellowish, 10-15 mm long, pappus bristles not connate
into ring, barbellate, with barbules shorter than thickness of hair or.almost
as long; achenes 5—6 mm long, four-angled, obpyramidal, glabrous,
smooth or with parallel stripes, toothed above, and with cupuliform
pappogenus at tip. Flowering V—VIII; fruiting VIJ-IX.

In the region of semideserts, on sands, sandy steppes, chalky areas —
European Part: Trans-Volga Region; Lower Volga, Lower Don; Western
Siberia: Upper Tobol (extreme west); Soviet Central Asia: Aralo-Caspian
Region (north). Endemic. Described from western Kazakhstan from the
Bolshoye [Great] Bogdo Mountains. Type in Leningrad.

Note. It must be remembered that the auricles at the base of
the leaves are not always well developed. Cf. the note to J. charcoviensis
Klok.

6. J. charcoviensis Klok. in Vizn. Rosl. URSR (1950) 564; Bot.
Zhurn. URSR, VIII, 1, 49.—J. eversmanni Korsh. in Izv. Akad. Nauk,
1, 2 (1894) 119 p. p. non Bge. (excl. specim. Saratow, Astrachan, Uralsk,
Orenburg); Iljin in Fl. Yugo-Vost. VI, 391; Schmalh. Fl. II, 110 p. p.;

5)

—

546

Korsh. Tent. Fl. Ross. or. 252.—J. cyanoides < eversmannii Korsh. Ibid.
252.—Ie.: Klok. op. cit. Fig. 7.

Perennial. Plants 15-60 cm high; stems solitary or few, straight,
finely-striated, finely flocculose-arachnoid-hairy, simple or with fewer
branches usually above middle or slightly below. Basal and lower cauline
leaves petiolate, with white-tomentose petioles, undivided (J. integrifolia
Iljin), pinnatisect or less often bipinnatisect into oblong or oblong-
lanceolate, less often linear, acuminate, acute or short-cuspidate, entire
or somewhat toothed segments, decurrent along rachis, revolute, green
or grayish-green above due to fine arachnoid-hairy, pubescence especially
dense along midrib, usually soft, less often scabrous due to short and
stiff papillae, white-tomentose beneath, with sessile glandular hairs under
tomentum; middle cauline leaves short-petiolate or sessile, divided, like
all other cauline leaves, uppermost short, simple, all without or with
poorly developed auricles, less often auricles well developed. Capitula
few, variable in size, terminal on stem and branches, usually medium,
less often large or smaller, with many florets. Involucre cupuliform,
arachnoid-hairy, less often glabrous; outer involucral bracts narrow-linear
or linear-lanceolate, acuminate, deflexed, or less often regularly arranged
so. Corolla 14-20 mm long, with scattered sessile glandular hairs, pink-
purple, narrowed over 6-8 mm, with 4-6 mm long lobes of limb. Pappus
10-15 mm long, usually yellowish, pappus bristles not connate into ring,
serrate-barbate, with barbules mostly shorter than thickness of hairs,
less often as long, densely arranged; achenes oblong, four-angled,
obpyramidal, 4-5 mm long, but glabrous, smooth or striate, brownish,
narrowed toward base, toothed above, and with prominent pappogenus
in center at tip. Flowering V—IX; fruiting VI—X.

Sandy river terraces, steppe sands, pine-forest sands.—European
Part: Lower Don, Middle Dnieper, Volga-Kama. Endemic. Described
from the vicinity of Merefa in Kharkov Region. Type in Kiev.

Note. This undoubtedly very young species is characterized by a
large amplitude of morphological features fluctuating between J.
eversmannii Bge. and J. cyanoides (L.) DC. According to M.V. Klokov,
the main features distinguishing J. charcoviensis Klok. from J.
eversmannii Bge. are: leaves not stiffly scabrous, pinnate, without auricles
at the base, semiamplexicaul; involucre arachnoid, the bracts sharply
deflexed as in J. eversmannii Bge. Even S. Korshinsky in 1894 and
1898 reported that to the north and west plants of the latter species
develop the characters that were later reported by Klokov for J.
charcoviensis Klok. These plants were tentatively assigned by Korshinsky
to the hybrid J. cyanoides x J. eversmannii Bge. However, he suggested
in his conclusion the possibility of climatic factors influencing the
manifestation of this character combination, i.e., in essence he suggests

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a racial character for these features. Following M.V. Klokov, we consider
it appropriate to treat these plants as separate species, albeit in the
formative stage.

7. J. granitica Klok. in Vizn. Rosl. URSR (1950) 564 (nomen);
Klok. Botan. Zhurn. Akad. Nauk URSR, VIII, 1 (1951) 48.—Ie.: Klok.
op..cit. (1951) Fig»,6.

Perennial. Plants 20-75 cm high; stems solitary or few, straight,
finely arachnoid-hairy, sulcate-striate, dirty purple, or striations dirty —
purple, and furrow green, rarely stems simple, usually branched above,
less often from base. Basal and lowermost cauline leaves short-petiolate,
bipinnatisect or even almost tripinnatisect, occasionally with mixture of
pinnatisect or even entire leaves, with narrow-linear or almost filiform,
acuminate, .entire lobes, narrowly involute, erect-spreading or almost
horizontal; cauline leaves sessile, especially middle leaves
semiamplexicaul or with well developed auricles; all leaves green and
stiffly scabrous due to well developed cartilaginous papillae on upper
surface, with scattered sessile glandular hairs, finely rugose, lower surface
gray arachnoid-hairy or white-tomentose. Capitula medium, with many
florets, solitary. Involucre cupuliform, always distinctly arachnoid-hairy
or even white-tomentose, especially its outer and middle bracts, near
middle or slightly above deflexed, dark green or purple in deflexed
part, gradually acuminate, inner row bracts straight, arachnoid-hairy only
below, toward apex brownish, membranous, acuminate and strigose on
outerside with scattered sessile glandular hairs, finely ciliate. Corolla
purple, with scattered sessile hairs, (14)15—16 mm long, narrowed over
6-7 mm, with (4.5)5-6 mm long lobes of limb. Pappus snow-white,
pappus bristles not connate into ring, serrate-barbate; achenes 4.5-5 mm
long, obpyramidial, four-angled, glabrous, smooth, longitudinally striate,
whitish or whitish-brown, toothed above, with patellate or smooth dome-
shaped pappogenus in center of tip. Flowering VI—VIII; fruiting VII-IX.

On gneiss-granites, stony steppes, sands.—European Part: Black Sea
Region (Donetsk, Zaporozhe and Dnepropetrovsk regions). Endemic.
Described from the village of Starodubovka in the Pervomaisk District
of Donetsk Region. Type in Kiev, isotype in Leningrad.

Subseries 3. Hamulatae [ljin—Series Salicifolia Klok. nom. Bot.
Zhurn. Akad. Nauk URSR, VIII, 1 (1951) 47.—Involucral bracts slightly
uncately bent upward.

8. J. salicifolia Gruner in Bull. Soc. Nat. Mosc. XLI, 2 (1868) 431
non Boiss.; Schmalh. Fl. II, 111 nom.; Iljin in Izv. Bot. Sada Akad. Nauk
SSSR, XXX, 3-4, 347 (emend.); Klok. in Vizn. Rosl. URSR, 564.—Ie.:
Klok. Ibid. Fig. 5.

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548

‘Perennial. Plants 20-70 cm: high; stems solitary or few, straight,
striate-sulcate, arachnoid-hairy, in upper half with few upward-spreading
branches. Basal leaves numerous, imparipinnate, with two to six pairs
of long, broad or narrow, acuminate or cuspidate lobes, often with
mixture of entire leaves or with solitary lobes, sometimes leaves
undivided predominate (f. integrifolia Iljin), narrowed into more or less
long petiole with wings, gradually reducing toward base; cauline leaves
sessile, except lowermost, always undivided and entire, narrow-lanceolate
or linear, uppermost leaves filiform, never decurrent; all leaves narrowly
involute and stiffly scabrous; above finely rugose due to short papillae,
with sessile glandular hairs in pits, glabrous, green, sometimes weakly
arachnoid-hairy, white-tomnetose beneath with scattered sessile glandular
hairs under tomentum. Capitula solitary, 20-30 mm wide, with many
florets. Involucre cupuliform, 12-15 mm long; outer and middle bracts
oblong-ovate, distinctly arachnoid-hairy, with short, uncate, glabrous,
purple or dark green, recurved tip; innermost bracts oblong-lanceolate,
straight, acuminate, purple, with sessile glandular hairs and slightly visible
appressed bristles on outer side, ciliate at apex. Receptacle covered with
smooth, acuminate scales almost up to half length. Corolla pink-purple,
with scattered sessile glandular hairs, 14-17 mm long, narrowed over
5.5—-8 mm length, with 4-5 mm long lobes of limb; apical appendages
of anthers gradually narrowed upward, about 2 mm long, basal
appendages caudate-setaceous, 2—2.5 mm long, filaments glabrous; stigma
about 2 mm long; style glabrous. Pappus snow-white, two times, and
with inner bristles three times, as long as achenes, pappus bristles serrate-
barbate, with teeth slightly shorter than thickness of hairs; achenes
oblong, obpyramidal, about 5 mm long, four-angled, narrowed below,
light colored, slightly brownish, toothed above, with ovoid dome-shaped
pappogenus at extreme tip and with stylar remains at end, with weak,
oblong-parallel ridges. Flowering VI-—VIII; fruiting VIII-IX.

Limestone deposits, steppe slopes, sands, especially along the banks
of Dnieper—European Part: Black Sea Region, Middle Dnieper, Volga-
Don (southwest). Endemic. Described by Gruner from the banks of the
Orel River above Dnepropetrovsk.

Note. Owing to the loss of the type specimens, I propose to select
as the neotype the specimen collected by S. Postrigan, 16 VII 1923, in
Cherkassy Region, near village Yablunivtsi. Type in Leningrad.

Series 2. Plerocladae Iljin——Series Albicaules Klok. |.c. 52, nom.
nud. (p. p.).—Stem winged; leaves decurrent or flanges decurrent.

Subseries 1. Albicaules Iljin.—Involucral bracts loosely and
entangled, deflexed as in J. cyanoides.

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549

9. J. albicaulis Bge. in Flora, XXIV, 1 (1841) 156; Ldb. FI. Ross.
II, 2, 765 (specim. sibir. alt.); [jim in Tr. Turk. Nauchn. Obshch. II, 20;
Kryl. Fl. Zap. Sib. XI, 293 (p. p.)—J. polyclonos, Kryl. Fl. Alt. III
(1904) 713 non DC.—J. cyanoides = polyclonos Korsh. in Bull. Acad.
Sc. Pétersb. V. ser. I (1894) 128.—Serratula cyanoides Ldb. FI. alt. IV
(1833) 42 non DC., specimina sibirica, synon. excl—J. angustifolia
Bge. in Flora, XXIV, 1 (1841) 156.

Perennial. Plants 15-40 cm high; stem densely white-tomentose,
straight, branched mainly in inflorescence. Leaves pinnatisect into linear
or linear-lanceolate, entire, long and deflexed segments, less often all
leaves undivided and entire (f. integrifolia) or with short solitary lobes,
acuminate, with callose tip; basal and lower cauline leaves petiolate,
with white-tomentose petiole, slightly broadened at base; cauline leaves
sessile, weakly semiamplexicaul and narrowly decurrent; uppermost
leaves linear, always undivided and entire, all leaves distinctly convolute,
rugose-alveolate above and scabrous due to small papillae, with sessile
glandular hairs in pits, grayish-green due to: weakly arachnoid-hairy
pubescence, strongly arachnoid-hairy along midrib, white-tomentose
beneath. Capitula medium, with many florets. Involucre broadly
cupuliform; outer involucral bracts densely arachnoid-hairy tomentose,
linear, almost equal, irregularly divided, almost as in J. cyanoides L.,
innermost row of narrow-lanceolate erect bracts, acuminate, usually
recurved, more or less densely covered on outer side with sessile
glandular hairs, slightly toothed along margin. Corolla pink-purple, 15—
22 mm long, narrowed over 7-10 mm length, with 5-6 mm long lobes
of limb, with scattered sessile glandular hairs, apical appendages of
anthers about 3 mm long, basal appendages caudate-setaceous, about
2.5—3 mm long. Pappus exceeding achene, pappus hairs serrate-barbate,
with the dense barbs, as long as or shorter than thickness of hairs, not
connate into ring; achenes 4—5.5 mm long, four-angled, obpyramidal,
distinctly narrowed below, glabrous with visible parallel stripes at angles,
toothed above, with broadly dome-shaped pappogenus at apex. Flowering
V-VII; fruiting VII-VII.

On sands, especially forest sands——Western Siberia: Irtysh, Altai;
Soviet Central Asia: Balkhash Region(?). Endemic. Described from the
region of the Bekun River, a tributary of Irtysh. Type in Leningrad.

10. J. gorodkovii Iljin sp. n. in addenda XXVI, 717.—J. cyanoides
Kryl. (non DC.) Fl. Alt. III (1904) 1904 (specim. Tobol).

Perennial. Plants 20-50 cm high; stems solitary or few, erect, ciliate,
finely arachnoid-hairy; simple or with fewer branches; branches slightly
deflexed. Basal leaves numerous, like cauline leaves, narrow, oblong-
linear, acuminate, simple or with isolated segments, entire, green and

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finely rugose above with scattered sessile glandular hairs, usually weakly
arachnoid-hairy, especially along somewhat depressed midrib, white-
tomentose beneath, with numerous sessile glandular hairs under
tomentum, gradually narrowed toward base; cauline leaves few, simple,
sessile, semiamplexicaul and decurrent with narrow wing; uppermost
leaves almost filiform, also decurrent or only with auricles at base.
Capitula 1-10, 10-15 mm wide, less often up to 20 mm. Involucre
cupuliform, 10-15 mm long, finely arachnoid-hairy, all or outermost
involucral bracts with herbaceous tip, thin-linear, not appressed or
appressed only at base, with flexuous erect-spreading apices, inner row
erect, of oblong, acuminate bracts with more or less sessile glandular
hairs, short-setose at apex. Corolla pink or pink-purple, 13-15 mm long,
narrowed over 5—6 mm length, with 4—4.5 mm long lobes of limb.
Pappus snow-white, about 10 mm long, with bristles not connate into
ring, with barbules as long as or usually shorter than hair thickness;
achenes 4.5—-5 mm long, oblong, obpyramidal, glabrous and smooth at
the faces or striate throughout, brownish, with small teeth above, teeth
slightly incurved, and with somewhat prominent cupuliform pappogenus
at tip. Flowering VI—VIII; fruiting VIII-IX.

In sandy pine forests and on sands.—Western Siberia: Upper
Tobol. Endemic. Described from the vicinity of Yalutorovsk. Type in
Leningrad.

Note. So far known only from the vicinity of Kurgan and
Yalutorovsk. This is the northermost species of the series Pleroclodae.

11. J. creticola Iljin sp. n. in Addenda XXVI, 717.

Perennial. Plants 15—30 cm high; stems erect, occasionally ascending,
flocculose-arachnoid-hairy, striate, branched, less often simple. Basal as
well as lower cauline leaves petiolate, white-tomentose, with pinnatisect
lamina, less often bipinnatisect, with narrow-linear segments, acuminate
or acute, deflexed, involute almost up to midrib, less numerous; middle
and upper cauline leaves sessile, decurrent forming narrow ridges; all
leaves somewhat rugose above, with sessile glandular hairs scattered
between wrinkles, slightly scabrous, white-tomentose beneath with similar
glandular hairs under tomentum; uppermost leaves undivided and entire,
distinctly reduced, linear or filiform. Capitula solitary, 20-30 mm wide,
with many florets, with involucre about 12-15 mm long, outer and middle
involucral bracts arachnoid-hairy, acuminate, loosely entangled, deflexed,
as in J. cyanoides; inner bracts straight, membranous in upper part,
purple, acuminate, ciliate, somewhat strigillose dorsally. Corolla pink-
purple, 16-18 mm long, narrowed over 7-8 mm length, with 5—6 mm
long lobes of limb. Pappus snow-white, with short-barbate bristles and
dense barbules; achenes immature, striate throughout the length.

531

Plate XLI.
Jurinea laxa (Fisch.) Ijin.

557

558

552

Sands of chalky exposures.—European Part: Lower Don (northeast).
Endemic. Described from the vicinity of Khvalinsk from the Peschanaya
Gora Mountain. Type in Leningrad.

12. J. laxa Fisch. ex Ijin in Izv. Glavn. Bot. Sada, XVII, 1 (1928)
84; Korsh. in Izv. Akad. Nauk 1, 2 (1894) 122, nomen; Klok. in Vizn.
Rosl. URSR, 565.—J. ambigua auct. non DC.: Lavr. and Pryan. in Mat.
do Dosl. Grunt. Ukr. 1 (1926) 199.—J. polyclonos Schmalh. F1. II (1897)
110 p. p.; Pachaskii in Zap. Novoross. Obshch. Est. XX VI, 101, XXXIV,
178.—J. albicaulis B. longifolia Ldb. Fl. Ross. II (1844-1846) 765.—J/.
longifolia Ldb. Ibid. 765, non DC. Iljin, op. cit. 85——Carduus laxus
Fisch. ex Iljin, op. cit. 84.

Perennial. Plants strong, usually 40-75 cm high, with strong woody
root; stem straight, 4-7 mm thick, ribbed-sulcate in lower part, strongly
arachnoid-hairy, branched in upper half, less often simple, solitary or
few. Basal or lowermost cauline leaves also 20-30 cm long, lanceolate-
oblong, undivided and entire, acuminate, repand, narrowed into relatively
short petiole slightly broadened at base, other cauline leaves of same
shape but reduced upward, sessile, semiamplexicaul and decurrent, not
reaching capitula; all leaves grayish-greenish above due to fine arachnoid-
hairy pubescence, with prominent whitish midrib, more densely
arachnoid-tomentose beneath, with scattered sessile glandular hairs on
both sides. Capitula large, with many florets, solitary on long leafless
peduncles. Involucre broadly cupuliform, four-rowed; outer involucral
bracts linear-lanceolate, quite loosely arranged, long acuminate, densely
arachnoid-hairy at fruiting rather sharply deflexed, dark purple on inner
side, glabrous, rugose, inner row of bracts erect, long acuminate,
sometimes with slightly deflexed apex, dark purple, densely covered
with short bristles and scattered sessile glandular hairs, ciliolate.
Receptacle covered with smooth, glabrous scales, divided in upper half.
Corolla pinkish-purple, covered with sessile glandular hairs, 17—22 mm
long, narrowed over 7-10 mm length, with 4.5—6 mm long lobes; apical
appendages of anthers acuminate, 2-3 mm long, basal appendages
densely caudate, usually about 2 mm long; filaments glabrous; stigma
1.5—2 mm long; style glabrous. Pappus of numerous hairs, with two
particularly prominent hairs, two times as long as achene, hairs
serrate-barbate, with barbels shorter than thickness of hair, not connate
into ring, divaricate at fruiting; achenes about 6 mm long, obpyramidal,
four-angled, acutely narrowed towards base, toothed at apex, glabrous,
smooth, with parallel stripes on edges, and with pappogenus at tip.
Flowering VI-VIII; fruiting VI-IX. (Plate XLI.)

Sands, sandy “hummocks.”—European Part: Black Sea Region.
Endemic. Described from the Aleshi Sands. Type in Leningrad.

F353

Subseries 2. Amplexicaules [ljin.—Involucral bracts abruptly
recurved from middle or above.

13. J. amplexicaulis (S.G. Gmel.) Bobr. in Bot. Zhurn. XLIII, 11
(1958) 1544.—Carduus inermis ...J.G. Gmel. Fl. Sib. (1749) 44, tab.
XVI.—C. cyanoides . polyclonos L. Sp. pl. (1763) 1152.—C. cyanoides
Pall. Reise, III (1776) 594.—C. polyclonos MB. FI. taur.-cauc. II (1808)
271.—Centaurea amplexicaulis S.G. Gmel. Reise I (1770) 136, tab.
XXIV.—Serratula polyclonos MB. Fi. taur.-cauc. HI (1819) 550.—
Jurinea polyclonos DC. Prodr. VI (1837) 675; Bge. in Flora, XXIV,
155; Ldb. Fl. Ross. II, 2, 762; Korzh. in Izv. Akad. Nauk V, 1, 119 p.
p.; Fedtsch. and Fler. Fl. Evrop. Ross. 1004; Ijin in Fl. Yugo-Vost. VI,
362; in Majevsk. Fl. Izd. 8, 598.—Jurinea polyclonos Claus in Goebel,
Reise, II (1838) 280.

Perennial. Plants 25—60(75) cm high; stems solitary or few, straight,
branched, especially in upper half, green or dirty-purple, usually
arachnoid-hairy, finely sulcate at base, winged. Basal and lowermost
cauline leaves petiolate, petiole densely white arachnoid-hairy, usually
pinnatisect (f. pinnatisecta), less often bipinnatisect (f. bipinnatisecta)
or linear-filiform, less often oblong-lanceolate segments entire, less often
all leaves entire (f. integrifolia); other cauline leaves amplexicaul,
decurrent; uppermost leaves usually undivided, all leaves bright green,
lustrous, and finely rugose above, with scattered sessile glandular hairs
or covered with fine arachnoid-hairy pubescence, with one vein, whitish-
tomentose below involute. Capitula comparatively small, numerous, with
many florets, aggregated in corymbose panicles. Involucre cupuliform,
10-15 mm long and 7-10 mm wide, grayish arachnoid-hairy; outer and
middle involucral bracts linear, acuminate, sharply recurved in middle
or slightly above, with glabrous or subglabrous dark violet or dark
green recurved part; innermost bracts linear-lanceolate, acuminate,
straight, whitish in upper half, with purple tinge in lower half.
Receptacle densely covered with yellowish acuminate scales reaching
almost to half of involucre length. Corolla pale-pink or pink purple,
with scattered sessile, glandular hairs, usually 12-15 mm long, narrowed
over 4-5 mm length, with 4—5 mm long lobes of limb; apical appendages
of anthers mostly about 2 mm long, somewhat pale, gradually acuminate;
basal appendages caudate, 2—-2.5 mm long; filaments glabrous. Pappus
snow-white, 12-15 mm long; pappus bristles not connate into ring,
serrate-barbate, with barbels always shorter than thickness of hair;
achene oblong, obpyramidal, gradually narrowed toward base, pale,
with parailel stripes on edges, glabrous, toothed above, and with
broad cupuliform, pappogenus at tip. Flowering V-—IX; fruiting VII-
IX.

560

554

In the southern steppe and semidesert zones, on sandy steppes and
hummocky sands, on chalky debris——European Part: Volga-Don, Lower
Don, Lower Volga, Trans-Volga Region; Western Siberia: Upper Tobol
(western part); Soviet Central Asia: Aralo-Caspian Region (north).
Endemic. Described from the Don Region in Voronezh District.

14. J. altaica Iljin sp. n. in Addenda XXVI, 717.

Perennial. Plants 25—40 cm high; stems solitary or few, branched
almost from base, branches spreading upward, more or less arachnoid-
hairy, finely striate. Basal leaves gradually narrowed into petiole, like
lower cauline leaves, pinnatisect into oblong-linear segments as broad
as leaf rachis, three to five pairs, obliquely opposite and directed upward
at an angle, acuminate, cuspidate, or subobtuse, relatively sparse; all
cauline leaves rather broadly long-decurrent, upper leaves grayish-green
due to finely arachnoid-hairy pubescence, more dense along depressed
midrib, finely rugose, with sessile glandular hairs, involute, white-
tomentose beneath with prominent midrib. Capitula solitary, medium,
aggregated in corymbose or corymbose-racemose inflorescence, with
many florets. Involucre cupuliform, arachnoid-hairy; outer involucral
bracts gray arachnoid-hairy, uncate, recurved; inner bracts erect,
membranous at apex, with somewhat ferruginous cusps, on outer side
with sessile glandular hairs or short appressed bristles, less often some
involucres with all appressed bracts. Pappus persistent, pappus bristles
not connate into ring, serrate-barbate. Corolla purple, with sessile
glandular hairs, 14-15 mm long, narrowed over 4—5 mm length, with

about 4 mm long lobes of limb. Achene four-angled, obpyramidal, with

fine longitudinal ribs, somewhat toothed above, pappogenus flat, dome-
shaped. Flowering VIJ—VIII; fruiting VII-IX(?).

Forest sands.—Western Siberia: Irtysh. Endemic. Described from
northeastern Kazakhstan in the vicinity of Krasnyi Aul. Type in
Leningrad.

Note. Close to J. polyclonos (L.) DC. Easily distinguished from the
latter species by arachnoid limbs of the outer involucral bracts.

15. J. transuralensis Iljin sp. n. in Addenda XXVI, 718.—J/.
polyclonos Ldb. FI. Ross. II (1844-1846) 763 (specim. alt. soong.-kirgh.)
non DC.

Perennial. Plants 15—70 cm high; stem erect, striate throughout,
flocculose-arachnoid or even densely white-tomentose, solitary or few,
less often simple, usually branched in upper half or upper one-fourth to
one-third, branches upward spreading, winged. Basal leaves numerous,
petiolate like sometimes lowermost cauline leaves, less often undivided
(f. integrifolia Iljin), usually pinnatisect in long, linear, remote lobes,

561

3)2)5)

acuminate, entire, narrowly involute, other: cauline leaves sessile,
auriculate, semiamplexicaul and decurrent along stem forming broad or
narrow, long or short wing, their laminas like basal leaves, becoming
progressively simple upward; uppermost leaves undivided, linear and
decurrent; all leaves green and glabrous above or usually with finely-
arachnoid-hairy pubescence, especially along midrib, smooth or scabrous
due to small cartilaginous papillae, finely rugose, with scattered sessile
glandular hairs, white-tomentose beneath with prominent midrib, and
with numerous sessile glandular hairs under tomentum. Capitula relatively
small to medium, aggregated in pyramidal panicle. Involucre cupuliform,
10-15 mm long, with many bracts, almost glabrous to strongly arachnoid-
hairy; outer involucral bracts acuminate, green in upper half with very
smoothly arcuate cusp and directed sideward, inner row erect, acuminate,
purple, with whitish membranous apices, ciliolate, with slightly visible
short, appressed bristles and sessile glandular hairs on outer side. Corolla
pink or pink-purple, with scattered sessile glandular hairs, 12-15 mm
long, narrowed over 5—6 mm length, with 4-5 mm long lobes of limb.
Pappus snow-white, (10)12—15 mm long, pappus bristles not connate
into ring, serrate-barbate, barbules usually shorter than thickness of hair,
achenes brown, glabrous, oblong, obpyramidal, four-angled, weakly
ribbed at edges, toothed on upper margin, with pappogenus of various
forms at tip. Flowering V-—IX; fruiting VI-X.

Feather-grass meadow steppes on sandy-loam, hummocky sands,
forested dunes, in pine forests, birch aspen forests on sandy soil, chalk
and chalky rubble-——European Part: Lower Volga (extreme east), Trans-
Volga Region (extreme southeast); Western Siberia: Upper Tobol (west);
Soviet Central Asia: Aralo-Caspian Region (northwest). Endemic.
Described from the Uila River Basin in Aktyubinsk Region. Type in
Leningrad.

Note. The western boundary of this species is approximately the
right bank of the Ural River (known from Urda and Orenburg and
stretching up to the Mugodzharsk Mountains and up to the large sandy
massifs of Taisugany, Kok-Dzhida, Kuguzyuk-Kum, Bolshie Barsuki);
in the latter, this species is replaced by J. kazachstanica I\jin. It may be
noted that J. polyclonos (L.) DC. s. str. is also found in this area, with
which J. transuralensis possibly hybridizes, as with J. cyanoides (L.)
DC. and J. eversmannii Bge.

Subseries 3. Hamatae I[ljin—Outer and middle involucral bracts
uncate only at extreme apex.

16. J. ciscaucasica (Sosn.) Iljin comb. n. hoc loco.—J. polyclonos
ssp. ciscaucasica Sosn. in Vestn. Tifl. Bot. Sada, 49 (1920); in Zhurn.

556

Russk. Bot. Obshch. 11, 1, 2, 196; Grossh. Fl. Kavk. IV, 171.—J.

‘polyclonos auct. p. p.: DC. Prodr. VI, 675; Ldb. Fl. Ross. II, 763; Boiss.

Fl. or. II, 576.—Carduus polyclonos MB. FI. taur.-cauc. II (1808) 271
p. p.—Serratula polyclonos MB. Ibid. Supplementum (1819) 550, syn.
excl.

Perennial. Plants 25-60 cm high, distinctly white-tomentose; stems
few or even solitary, erect, sulcate-striate, white-tomentose, less often
simple, usually branched above middle or in upper one-third. Basal and
lower cauline leaves gradually narrowed into petiole, white-tomentose,
as if bordering decurrent lamina, pinnatisect (f. pinnatisecta Iljin f. n.)
into narrow-linear or lanceolate-oblong, entire or dentate-sinuate,
acuminate segments, narrowly involute; all leaves undivided (f.
integrifolia Iljin) or almost entire, with isolated lateral segments; other
cauline leaves also pinnatisect or undivided, always decurrent, then stem
appearing angular; uppermost leaves linear-filiform, sessile; all leaves
usually finely arachnoid-hairy above, more densely along midrib, with
small sessile glandular hairs, densely white-tomentose beneath, with
prominent midrib. Capitula aggregated in corymbose, less often
corymbose-paniculate inflorescence, comprising 5—10, less often up to
20 capitula, occasionally solitary. Involucre broadly cupuliform, 10-12
mm long, up to 20 mm wide, with many bracts at fruiting imbricate,
outer rows of bracts powdery-arachnoid-hairy, with scattered sessile
glandular hairs, obtusely carinate, dark green or purple at tip; outer
bracts shortly-arcuate, subsequent bracts slightly recurved, other bracts
straight; innermost bracts prominent, acuminate, with purple tinge, tip
ciliolate, whitish-membranous, setulose. Receptacle densely covered with
smooth, pointed scales, half as long as involucre or slightly more. Corolla
pink-purple, with scattered sessile glandular hairs, 12.5—14.5 mm long,
abruptly narrowed over 5-6 mm, with 4-5 mm long lobes of limb,
apical appendages of anthers acuminate, 2.5—3 mm long; basal
appendages caudate, about 2—2.5 mm long; filaments glabrous. Pappus
snow-white, 8-12 mm long, pappus bristles not connate into ring, with
barbules shorter than thickness of hair axes, or almost as long; achene
whitish, obpyramidal, four-angled, narrowed toward base, with faint
parallel stripes on edges, toothed on upper margin, with cupuliform or
obtuse dome-shaped pappogenus in center of tip, 3.5—5 mm long.
Flowering V—VIII; fruiting VII-X.

Hummocky sands and dunes at sea coast, sandy-stony and gravelly
places of river valleys in foothills—Caucasus: Ciscaucasia, Dagestan.
Endemic. Described from Nalchik. Type in Leningrad.

Note. D.I. Sosnowsky described this plant as a subspecies (1920);
actually it is a good species, endemic to the foothills of the main
Caucasian Range. I consider the specimens collected by V.I. Lipsky

563

SDT

from the vicinity of Nalchik on July 25, 1893, as the types of the
species.

The range of J. ciscaucasica (Sosn.) Iljin and J. polyclonos (L.)
DC. adjoin at the boundary of the lowland and the Ciscaucasian foothills.

17. J. kazachstanica Iljin sp. n. in Addenda XXVI, 718.—
J. polyclonos B. Fedtsch. (non DC.). Perech. Rast. Turk. IV (1911)
30 (specim. turg. non DC.); Ijin in Tr. Turk. Nauchn. Obshch. 2, 21
p. p.
Perennial. Plants 20-60 cm high, with long rhizomes covered with
brown scales; stems solitary or several, erect, white-tomentose or less
often flocculose-arachnoid-hairy, discontinuously winged, less often
simple, usually branched above middle; branches upward spreading,
weakly deflexed. Basal leaves numerous, petiolate like lower cauline
leaves, petiole white-tomentose, shorter than lamina, pinnatisect into
linear, oblong, remote, acuminate, entire segments in three to five pairs,
obliquely directed upward, less often undivided, oblong, or oblong-linear
(f. integrifolia lljin); other cauline leaves sessile, long-decurrent,
becoming simple upward to undivided linear or filiform, also decurrent;
all leaves green or gray-green above, weakly arachnoid-hairy, especially
along midrib or even flocculose-arachnoid-hairy, smooth or rough due
to small papillae, finely rugose, revolute, white-tomentose beneath, with
numerous sessile glandular hairs under tomentum. Capitula about 10—12
mm wide, aggregated in pyramidal-paniculate or almost corymbose
inflorescence, with many florets. Involucre narrow cupuliform or
cyathiform-cupuliform, 10-15 mm long, usually white-tomentose, less
often arachnoid hairy-flocculose; outer as well as middle involucral bracts
with slightly floccose, recurved, dark green or purple tips, exserted from
tomentum; bracts of innermost row erect, acuminate, purple, covered on
outer side with sessile glandular hairs or rather short, almost punctate
bristles, with membranous border, ciliolate, narrowed into pale
membranous apex. Receptacle covered with smooth acuminate scales,
reaching more than half of involucral height. Corolla pink or pink-purple,
with sessile glandular hairs, (10)12—16 mm long, narrowed over 4—5
mm length, with 3-6 mm long lobes of limb. Pappus snow-white, 10—
12 mm long, pappus bristles not connate into ring, serrate-barbate, with
dense barbels almost as long as thickness of bristle; achenes 5-6 mm
long, four-angled, obpyramidal, dark brown, finely longitudinally
ribbed, with small teeth above, with somewhat flat pappogenus at tip,
slightly dentate along margins and with nectary above. Flowering V—
VIU; fruiting VI-IX.

Sandy steppes, hummocky sands of large sandy massifs of
semidesert.—Soviet Central Asia: Aralo-Caspian Region. Endemic.

564

558

Described from the Taisuchan Sands (western Kazakhstan). Type in
Leningrad.

18. J. pineticola [Ijin sp. n. in Addenda XXVI, 719.—J. pollichii
Ldb. Fl. Ross. II (1844-1846) 764 p. p. non DC.—Serratula cyanoides
Bge. in Lab. FI. alt. [V (1833) 41 p. p. non DC.—J/. polyclonos Korsh.
in Izv. Akad. Nauk, 1, 2 (1894) 119.—J. cyanoides x J. polyclonos
Korsh. Ibid. 121 (specim. e Loktjewsk. Irtysch, Semipalatinsk); O. and
B. Fedtsch. Perech. Rast. Turk. IV, 301, specim. semipal.—J. albicaulis
Iljin in Tr. Turk. Nauchn. Obshch. II (1925) 20 p. p. non Bge.—J.
albicaulis x J. cyanoides Schischk. in Kryl. Fl. Zap. Sib. XI (1949)
2933.

Perennial. Plants 15-35 cm high; roots strong, tap roots, vertical,
passing over to vertical or horizontal rhizomes, covered with brown
scales and shaggy remains of old leaves. Stems usually several, erect or
ascending, white-tomentose, less often flocculose-arachnoid, branched
above middle or near apex. Basal leaves numerous, usually undivided
and entire, narrow-oblong, acuminate, gradually narrowed toward base,
almost sessile, less often with some mixture of pinnatipartite or
pinnatisect into lanceolate or linear-oblong lobes, two or three_on each
side on short white-tomentose petioles; cauline leaves usually undivided,
gradually narrowed toward base, but sessile, not decurrent, auriculate or
slightly decurrent, reduced above, to small, filiform, white-tomentose,
sessile, nondecurrent leaves, all leaves green on grayish-green above
due to the fine arachnoid-hairy pubescence, finely rugose, with large
sessile glandular hairs in pits, smooth, less often weakly scabrous, with
white-tomentose midrib, more or less narrowly involute, white-tomentose
beneath, with prominent midrib and sessile glandular hairs under
tomentum. Capitula usually numerous, small, solitary or in groups of
two or three in corymb or panicle, with many florets, 10-12 mm long.
Involucre cupuliform, 10—12 mm wide; outer involucral bracts arachnoid-
hairy, appressed, sometimes slightly recurved, with flat tip, sometimes
purple or gray-arachnoid-hairy; inner bracts erect, purple, white-
membranous and acuminate, with sessile glandular hairs. Receptacle
covered with smooth acuminate scales, reaching almost to half of
involucral length. Corolla pink, pink-purple, or sometimes cream-colored,
with scattered sessile (10)13—15 mm long glandular hairs, narrowed over
3-5 mm, with 3-5 mm long lobes. Pappus snow-white, up to 10-12
mm long, bristles not connate into ring, barbate, with dense barbules, as
long as or slightly exceeding thickness of bristle; achenes (4.5)5—6 mm
long, four-angled, obpyramidal, whitish or brownish, glabrous, smooth,
striate along edges, finely toothed above, with flat, scarcely serrate
pappogenus and nectary at tip. Flowering VI-—VIII; fruiting VII-IX.

565

559

Forest bands, fringes of pine forests, sand dunes.—Western Siberia:
Irtysh (southeast). Endemic. Described from Shulbinsk Forest in
Semipalata Region. Type in Leningrad.

Note. An extremely polymorphic species, variable in the cutting of
the leaf blade, degree of decurrency of the leaves, and nature of the
involucral bracts. An increase of speciation is observed at the eastern
edge of the range of the species of section Cyanoides.

19. J. paczoskiana Iljin in Izv. Bot. Sada Akad. Nauk SSSR, XXX,
3-4 (1932) 345; Klok. in Vizn. Rosl. URSR, 564; Bot. Zhurn. URSR,
VII, 1, 53.—J. polyclonos Schm. FI. II (1897) 110 p. p.

Perennial. Plants 30-60 cm high; stems sulcate-ribbed, straight,
always arachnoid hairy-tomentose, especially at base, with scattered
sessile glandular hairs, less often simple, usually branched, especially in
upper half. Lower leaves often up to 25-30 cm long; basal and lowermost
leaves on petioles winged almost up to base, wing gradually narrowed
toward base, lamina imparipinnate, with few (two to six pairs), rather
long, remote, broadly lanceolate or oblong, less often lobes narrower,
acuminate or subobtuse, obliquely spreading, entire, sometimes in
combination with undivided leaves; middle cauline leaves distinctly
decurrent, also divided, but with fewer lobes or abruptly terminating
from amplexicaul part into entire, linear or oblong lamina with entire
margin; uppermost leaves strongly reduced, filiform, amplexicaul; all
leaves green or greenish above, weakly arachnoid-hairy, more distinct
along midrib, slightly rugose-punctate, with sessile glandular hairs in
pits, canescent beneath due to more dense arachnoid-hairy pubescence
with scattered sessile glandular hairs. Capitula medium or large, with
many florets. Involucre 15-20 mm long; outer involucral bracts
arachnoid-hairy, sharply recurved, with long, glabrous, recurved part,
usually exceeding appressed part, and gradually narrowed at tip;
outermost row of erect, lanceolate, acuminate, distinctly purple bracts,
with barely noticeable appressed bristles and scattered sessile glandular
hairs on outer side, ciliolate. Receptacle with smooth acuminate scales,
almost half as long as involucre. Corolla pink-purple, with scattered
sessile glandular hairs, 16-20 mm long, abruptly narrowed over 7-10
mm length, with 4-6 mm long lobes of limb; pappus 10-15 mm long
(with longer bristles), snow-white, with serrate barbules, not connate
into ring, serration or barbules shorter or almost as long as thickness of
bristle, achenes oblong-obpyramidal, four-angled, cuspidate and usually
with oblique hilum, not strongly toothed above, with slightly prominent
longitudinal ridges, glabrous, at apex with dome-shaped pappogenus
inside corona. Flowering and fruiting: VII-IX.

Compacted sands.—European Part: Black Sea Region (vicinity of

566

560

Nikolaev). Endemic. Described from the vicinity of Nikolaev. Type in
Leningrad.

Subseries 4. Apressae Iljin |.c——Series Tenuilobae Klok. in Bot.
Zhurn. Akad. Nauk URSR, VIII (1951) 53 n.n—aAll involucral bracts
appressed and slightly lax, but not recurved or uncate.

20. J. tenuiloba Bge. in Flora, XXIV, 1 (1841) 155; Ldb. Fl. Ross.
II, 765 p. p.; Iljin in Fl. Yugo-Vost. IV, 393; Ijin in Tr. Turk. Nauchn.
Obshch. 2, 21 (specim. turcom).—Ie.: Fl. Yugo-Vost., Fig. 700.

Perennial. Plants 25-40 cm high; stems solitary or few, erect, slender,
with few branches only at extreme tip or even simple, finely striate-
sulcate, finely arachnoid-hairy, white-tomentose only at extreme base,
winged. Basal leaves pinnatisect into narrow-linear, acuminate, far
recurved lobes, usually three to five on each side and, of course,
particularly narrowed gradually into petiole, shorter than lamina; all
cauline leaves, except uppermost, sessile, divided in same manner, but
with fewer lobes above; all leaves abruptly broadened at base,
semiamplexicaul and decurrent; uppermost leaves short, simple, filiform;
all leaves green, finely arachnoid-hairy above, more densely arachnoid-
hairy along depressed midrib, lustrous, fmely rugose, entire or weakly
repand, sometimes involute up to midrib, white-tomentose beneath.
Capitula one to five, with many florets. Involucre somewhat broadly
cupuliform, about 10-12 mm long; all involucral bracts appressed;
outermost bracts weakly arachnoid-hairy, oblong, cuspidate, with dark
green and purple tips; inner bracts purple, acuminate, membranous above,
with sessile glandular hairs and colorless cusps, setulose, indistinctly
ciliate. Corolla pink, with sparse sessile glandular hairs, 12-13 mm long,
abruptly narrowed over 4 mm length, with about 4 mm long lobes of
limb. Pappus snow-white, about 10 mm long, pappus bristles not connate
into ring, serrate-barbate, with squarrose barbules, as long as or slightly
shorter than thickness of bristle; achenes oblong, obpyramidal, four-
angled, pale, glabrous, smooth, with scarcely visible longitudinal, parallel
stripes, serrate above, about 5—5.5 mm long, with high pappogenus at
apex, terminating in nectary.

Soviet Central Asia: Aralo-Caspian Region (western). Endemic.
Described from the vicinity of the former Shevchenko Fort (Novo-
Aleksandrovsk). Type in Leningrad.

Note. This species was described by Bunge on the basis of a specimen
from the Herbarium of Escholtz with the label “Carduus cyanoides e
Tataria.” Since the original type specimen is lost, I consider the specimen
from the vicinity of Shevchenko Fort, collected by Karelin, as the type
of the species. In the description of J. tenuiloba Bge., Ledebour reported

561

the following localities: 1) Penza, based on collections of Jacquet; this
specimen does not conform to the original description of Bunge; 2)
Sibirica altaica (apparently, these are collections of Gebler from the
Altai); this plant conforms to the description, but it is doubtful if it was
collected from the Altai; 3) des. caspio—cited from Pallas; I did not
see Pallas’ specimens; however, Karelin’s specimens from Mangyshalk
fully agree with Bunge’s description; Karelin’s label from Novo-
Aleksandrovsk (Shevchenko Fort) bears the inscription: “Jurineo
polyclonos aff. forsan sp. nov.”.

21. J. tanaitica Klok. sp. n. in Addenda XXVI, 719.—J/. polyclonos
auct. specim. fanaitica non DC.—J. tenuiloba auct. sp. tanaitica non
Bge.

Plants 25—60 cm high. Stem solitary or few, erect, arachnoid-hairy,
finely striate, branched, usually discontinuously winged in upper one-
third part. Basal and lowermost cauline leaves petiolate, pinnatisect into
linear or oblong-lanceolate, entire, acuminate segments, petioles whitish
arachnoid-hairy; other cauline leaves, except uppermost, undivided, small
and short-filiform, pinnatisect or even undivided, sessile with
semiamplexicaul auricles, decurrent; all leaves green, lustrous, and
glabrous above, with depressed midrib, arachnoid-hairy, or lamina
entirely finely arachnoid-hairy, densely white-tomentose beneath. Capitula
numerous, in corymbose-paniculate inflorescence, small. Involucre
cupuliform, 10—12(15) mm long, usually, except innermost row,
arachnoid-hairy, less often almost glabrous, all bracts imbricate and
compactly appressed; outer bracts short, oblong, acuminate, with dark
green tips; inner bracts prominent, linear-lanceolate, acuminate, with
scattered sessile glandular hairs on outer surface or with appressed,
scarcely visible short bristles, or with both, light purple, terminating in
white membranous cusp, slightly ciliate along margin. Receptacle densely
squamose with smooth acuminate scales, up to half as long as involucre
or slightly longer. Corolla pink-purple, with scattered sessile glandular
hairs, 12-14 mm long, narrowed over 4—5 mm length and with 3.5—-5
mm long lobes of limb; apical appendages of anthers about 2—2.5 mm
long; basal appendages caudate, about 2 mm long. Pappus up to 10 mm
long, snow-white, with serrate-barbate bristles, not connate into ring,
with barbules as long as diameter of bristle or slightly shorter; achenes
pale, glabrous, four-angled, obpyramidal, 3.5—5 mm long, with parallel
striations on edges, somewhat serrate, with patellate or smooth dome-
shaped pappogenus at tip. Flowering VI—VI]J; fruiting VII-IX.

Sands of the Don River Valley in the lower and middle reaches
and along its tributaries—European Part: Lower Don. Endemic.
Described from the Chir River Valley near the village of Oblivskaya.
Type in Leningrad.

567

562

22. J. thyrsiflora Klok. in Bot. Zhurn. URSR, VIII, 1 (1951) 53;
Vizn. Rosl. URSR, 555.—J. polyclonos auct. mult. ucr.—Ic.: Klok. op.
cit. Fig. 9.

Perennial. Plants 25—80 cm high, with solitary or few stems. Stems
grayish-arachnoid, finely striate, strongly branched from middle or even
below. Basal and lower cauline leaves numerous, petiolate, usually
pinnatisect into linear or oblong-linear, entire, acuminate, often somewhat
arcuate segments; other cauline leaves also pinnatisect, long-decurrent
on stem; uppermost leaves, especially in inflorescence, linear, undivided,
reduced; all leaves green, glabrous, lustrous above, often with arachnoid-
hairy depressed midrib or entirely arachnoid hairy-pubescent and with
scattered sessile glandular hairs, white-tomentose beneath. Capitula very
small, usually numerous, with many florets, in corymbose-paniculate
inflorescence. Involucre cupuliform, about 10 mm long; involucral bracts
imbricate, grayish arachnoid-hairy; outer bracts shorter, oblong-lanceolate,
with dark purple or dark green apices, slightly deflexed (not appressed);
innermost bracts linear-lanceolate, acuminate with membranous apex,
and sessile glandular hairs on outer side. Receptacle squamose with
numerous, smooth, acuminate scales, up to half as long as involucre or
longer. Corolla pink-purple, with scattered sessile 11-13 mm long
glandular hairs, abruptly narrowed over 4—5 mm length, with 3-4 mm
long lobes of limb, apical appendages of anthers about 2 mm long,
basal appendages caudate, about 2 mm long, filaments glabrous; style
smooth. Pappus snow-white, about 10 mm long, pappus bristles serrate-
barbate, barbules dense, shorter than thickness of bristle, connate or not
into ring; achenes 3—4.5 mm long, pale, glabrous, smooth, elongate,
obpyramidal, gradually narrowed toward base, toothed above, with flat,
slightly dome-shaped pappogenus at extreme tip. Flowering VI-VIII;
fruiting VIII—X.

Sands in the valleys of the Dnieper, Oskol, northern Donets and
Don, mainly restricted to the left bank of Ukraine——European Part:
Lower Don (northwestern part); Black Sea Region; Volga-Don (western),
middle Dnieper. Described from the vicinity of Dnepropetrovsk from
the Chepelskn Island.

Section 2. Bellae Iljin sect. n. in Addenda XXVI, 720.—Section
Subacaules Boiss. Fl. or. II] (1875) 582 p. p. non Benth.: Sosn. in Zhurn.
Russk. Bot. Obshch. 11, 1-2, 198.—Perennial plants. Stem scapose, or
capitula sessile. Leaves rosulate; pappus persistent, serrate or serrate-
barbate, and bristles not connate into ring. Achenes 2.5—3(4—5) mm long,
glabrous, smooth or usually with parallel longitudinal striations, toothed
above, not forming a true corona. Pappogenus developed, pappophorus
absent. A plant of alpine and subalpine mountain zone of Caucasia.
Type of section: J. cadmea Boiss.

568

569

&

563

. Leaves undivided and entire, narrowly lanceolate, acuminate,

gradually narrowed toward base into petiole, sometimes almost
RAR IIMOUS!. Nia AL AEROS MES eet eetee cen dene snnentenonea 2.
Leaves usually pinnatisect, pinnate, lobate, or lyrate, if undivided
and entire, then of different shape and not acuminate.............. ay

. Leaves glabrous above, white-tomentose beneath ..............::eee

RDA ER OEE (Ney ROD GOLIST | ARIE: Es 2ay's. pumila Alb.
Leaves glabrous on both sides, green, scabrous above due to
papillacerene. atond..Jomuleval, 290... 24. J. woronowii Iljin

. Leaves undivided and entire, oval, oblong or oblong-obovate;

roundish at apex, obtuse or with short cusp. Pappus almost as
long as achene. Achenes about 2.5-3 mm long...........c:ceseeeeee
nthe AA OTT SOE! ONL Eh Lae REE, Lis 25. J. bellidioides Boiss.
eavesdvaniausly divided. 22008 Lee OM Oona PEO eed 4.

piMeavessareenvoneboth sides igs Lh oe. eae el cree cece 52

Leaves grayish arachnoid-hairy beneath or usually white-tomentose

. Leaves pinnatilobate, with wide, ovate, obtuse lobes almost as band

as winged leaf rachis, and almost as long as sinuses between lobes.
Pappus with fine bristles equaling human hair in thickness, sparsely
serrate. Achenes about 3 mm long..... 28. J. sosnovskyi Grossh.
Leaves pinnatipartite into oblong-linear, remote segments. Pappus
bristles two times as thick. Achenes about 5 mm long............ 6.
Rosettes reaching to half of scape or more, sometimes exceeding
capitula. Leaves 5-15 mm wide; segments of lamina 3-10 mm
long, all entire, usually acute or acuminate, especially terminal
segments, distinctly obliquely spreading. Involucral bracts
appressed, sometimes slightly decurved...... 26. J. iljinii Grossh.
Rosettes not reaching to half of scape. Leaves usually 20-40 mm
wide, segments 10-20 mm long, mostly with solitary teeth or short
lobes at base, obtuse or subobtuse, sometimes with short cusp,
obliquely spreading or almost horizontal. Involucral bracts contorted
HEPSAMS, (a Cer EE LAM ina. ALLS RGR IGUi CamREaT RATAN AMG AEE 27. J. venusta Iljin

. Leaves pinnatipartite, with obtuse lobes, usually broader than

sinuses between them. Capitula sessile, occasionally on short
peduncle wlarse ..Au00Rk. RR 32a. J. filicifolia Boiss.
Leaves of different type. Scape longer than leaves .................. 8.
Leaves pinnatisect in thin, linear or oblong-linear segments as broad
as narrowly bordered scape and narrower than sinuses between
them, acuminate, acute, cuspidate, or less often obtuse. Pappus
fourttimes! as fonetas achenien iii p ARO OSD bet ae
ith tC ute MERA Rain | GaN 31. J. coronopifolia Somm. and Lev.
Leaves lyrate or pinnatipartite, with oblong-obovate, always obtuse

570

564

lobes, usually as broad as sinuses or broader, or sinuate-cristate,
with less divided apex, less often almost undivided ................. oe
9. Leaves sparsely arachnoid-hairy above, weakly scabrous. Pappus
two to two and one-half times as long as achene, with sparse
SEMAN ON es asco. beott oeee. steep. ener 32. J. cartaliniana Boiss.
+ Leaves always glabrous and smooth above ...............:ceeeeeeeeeee 10.
10. Leaves with deeply depressed veins above, midrib as well as lateral
veins, distinctly punctate under high magnification, with more or —
less approximate lobes. Involucral bracts straight, erect, not
recurved, somewhat lax. Corolla 18-20 mm long. Pappus several
times as long as achene with sparse barbules .............0....eeeeeeeeee

+ Leaves with somewhat depressed midrib, not punctate, lobes never
approximate. Involucral bracts arcuate and somewhat differently
curved. Corolla about 15 mm long. Pappus almost as long as achene
or slightly longer, densely barbate .................. 30. J. levieri Alb.

Series 1. Integrae Iljin—All leaves undivided and entire.

23. J. pumila Alb. in Bull. Herb. Boiss. I (1893) 243; Tr. Bot.
Sada, XVI, 268; Grossh. Fl. Kavk. IV, 169; Sosn. Zhurn. Russk. Bot.
Obshch. 11, 1-2, 198.—J. pumila Alb. var. major Alb. Prodr. Fl. Colh.
(1895) 745.—Ie.: Fl. Gruzii, VIII, Plate 408; Alb. op. cit. (1893) t. IX,
3, 4 and 5—Exs.: GRF No. 3249.

Perennial. Plants 10-40 cm high, caespitose, usually with short
vertical rhizome, covered with dark brown scales, producing roots. Stems
scape-like, solitary, few, erect, cylindrical, sulcate, arachnoid-hairy, with
fairly scatered sessile glandular hairs, usually exceeding leaves. All leaves
in basal rosette, numerous, erect, narrow-lanceolate or linear, acuminate,
undivided and entire, less often, usually in large forms, largely and
sparsely retrorsely serrate in upper part, glabrous, lustrous and weakly
rugose above with depressed midrib, very narrowly involute, white-
tomentose beneath with numerous sessile glandular hairs and prominent
brownish midrib. Capitula solitary, with many florets. Involucre
cupuliform, 2—3.5 cm wide; involucral bracts four- or five-rowed; outer
row weakly arachnoid-hairy, linear, herbaceous, acuminate, arcuate in
upper part, usually slightly contorted and lax; innermost bracts more or
less erect, broader, three- to seven-veined, narrowed apically into long,
often aristate cusp; all bracts with scattered sessile glandular hairs.
Corolla pink-purple, usually glabrous, 14-18 mm long, narrowed over
5-7 mm length from base, with lobes of limb 4-6 mm long, apical
appendages of anthers 2-3 mm long, basal appendages 2.25—3 mm long,
flexuous-caudate; stigma about 2 mm long, with short papillae on outer

565

side, style glabrous. Pappus of numerous unequal bristles, 12-15 mm
long, not connate into ring, very short-barbate, almost serrate-barbate;
achenes 4—5 mm long, four-angled, glabrous, dark brown, light colored
at base, with corona of long simple teeth above, modified along edges
in long parallel ridges. Flowering and fruiting VII-VIII.

On rocks, especially limestone rocks in alpine belt.—Caucasus:
Ciscaucasia, western Transcaucasia. Endemic. Described from Abkhazia,
from the Akhalibokh Mountains. Type in Florence.

24. J. voronowii [Ijin in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 1
(1933) 185; Grossh. Fl. Kavk. IV, 169.

Perennial. Entire plant green, 12-20 cm high, with branched dark
caudex, densely covered with narrow blackish remains of dead leaves,
and rosette of basal leaves above. Stems leafless, erect, simple, distinctly
finely sulcate, terminating in solitary capitula, almost two times as long
as leaves, with arachnoid-hairy pubescence throughout. Leaves rosulate,
numerous, 5—10 cm long, narrowly-lanceolate, sometimes almost linear,
undivided and entire, or occasionally with isolated teeth, acuminate,
convolute, gradually narrowed toward base into petiole, green on both
sides, with single vein, finely rugose above due to distinct papillae,
densely glandular-hairy beneath. Capitula 20-25 mm across. Involucre
cupuliform, in upper part 1.5-2.0 mm long, glabrous or with only
scarcely visible, weakly arachnoid-hairy pubescence, purple, yellowish-
green at base; bracts linear, almost equal with long, thin, contorted,
smooth apical appendage. Corolla pink-purple, 16—17 mm long, narrowed
over 7-8 mm length, with 5-6 mm long limb; basal appendages of
anthers 3 mm long, setose, divided at apex; filaments smooth, inserted
at junction of broad and narrowed parts of corolla. Pappus with relatively
long-serrate, snow-white bristles with teeth shorter than thickness of
bristles, dense; achenes (immature) four-angled, glabrous and smooth.
Flowering VII—VIII; fruiting VIII-IX(?).

Alpine zone of mountains.—Caucasus: Ciscaucasia. Endemic.
Described from Abshier-Akhub. Type in Leningrad.

Series 2. Micrachaneniae [ljin—Achenes small; pappus equal or
shroter than achene.

25. J. bellidioides Boiss. Fl. or. III (1875) 583; Grossh. Fl. Kavk.
IV, 169; Sosn. in Zhurn. Russk. Bot. Obshch. 11, 1-2, 198.

Perennial. Plants about 10 cm high, caespitose. Rhizomes densely
covered with hardened remains of old leaves; stems scape-like,
cylindrical, erect, sulcate, arachnoid-hairy, terminating in single
capitulum. Leaves rosulate, petiolate, broadened at attachment, lamina

572

566

oblong-ovate, undivided and entire, less often very first leaves lyrate or
incised-pinnatipartite, obtuse or with very short cusp, less often subacute,
green with fine arachnoid-hairy coating and whitish midrib above, finely
rugose with sessile glandular hairs in pits, white-tomentose beneath, with
numerous sessile glandular hairs and brownish midrib, scarcely shorter
than scape. Capitula relatively small. Involucre cupuliform, 10-15 mm
wide; outer involucral bracts arachnoid-hairy, narrowly lanceolate or
almost linear, squarrose in upper half, smoothly recurved, acuminate,
innermost bracts erect, broader, three- to five-veined, narrowed apically
in membranous cusp. Pappus as long as achenes or slightly shorter,
pappus bristles serrate-barbate, with dense serrations, not connate into
ring; achenes usually 2.5 mm long, pale, glabrous, toothed above. Fruiting
VIII-IX. (Plate XLII, Fig. 2.)

On slopes in alpine belt—Caucasus: Central Caucasia. Endemic.
Described from Alagir near the Ardon River. Type in Leningrad.

Series 3. Tenuisectae Iljin Leaves with linearly divided segments.

26. J. iljinii Grossh. in Dokl. Akad. Nauk AzerbSSR, III, 9 (1947)
166.

Perennial. Plants 8—25 cm high, with branched rhizome; stems
simple, scape-like, leafless, erect, cylindrical, sulcate and finely ribbed,
finely and sparsely arachnoid-hairy, with scattered whitish papillae,
especially below. Leaves rosulate, numerous, narrowly oblong, first leaves
often undivided or almost undivided, others pinnatipartite, with entire
lobes remote and obliquely spreading, with broad stripe, bordering leaf
rachis, narrowly convolute, graudally narrowed toward base into usually
dirty-purple petiole, light green on both sides, with pale midrib, scabrous
due to short scattered papillae, with fine, sparse, sessile glandular hairs,
glabrous, with tufts of fine hairs at isolated places along midrib. Capitula
solitary, with many florets. Involucre cupuliform, usually 25-30 mm
wide, glabrous or weakly arachnoid-hairy below; outer involucral bracts
somewhat decurved above, herbaceous, slightly purple in remaining part,
more or less appressed; bracts of innermost row erect, weakly purple.
Corolla pink-purple, glabrous 14-18 mm long, narrowed over 6—9 mm
length, with 5-7 mm long lobes of limb; apical appendages of anthers
about 2.5 mm long, basal appendages flexuous-caudate, of same length;
style glabrous, stigma 1.5-2 mm long, with tuft of basal bristles, and
more or less large papillae outside. Pappus smooth, 12-15 mm long,
with few thicker and longer bristles, pappus bristles barbate, barbules
dense, shorter than thickness of bristles, not connate firmly into ring;
achenes immature, about 5 mm long, smooth, glabrous, with long-toothed
apical corona. Flowering VII—VIII; fruiting VIII.

567

SS

\
ap
ys

Vl

Ag

N

\

J

Plate XLII.
2—J. bellidioides Boiss; 3—J. cartaliniana Boiss.

573

Jurinea sosnovskyi Grossh.;

1

574

568

Subalpine meadows.—Caucasus: Western Transcaucasia.. Endemic.
Described from the Urup River Basin, and from Atsgara Mountain. Type
in Leningrad.

27. J. venusta Ijin sp. n. in Addenda XXVI, 720.

Perennial. Plants 25-30 cm high. Stem solitary, finely striate, with
scattered sessile glandular hairs, finely and sparsely arachnoid-hairy at
some places, with small filiform leaflet in middle. Leaves rosulate,
gradually narrowed into petiole, pinnatisect into oblong-linear, subobtuse,
entire, 2-3 mm wide segments, occasionally with small cusp, often with
solitary teeth near base or short obtuse lobes, convolute, as wide as
rachis and separated from each other by three to three and one-half
times their width, obliquely spreading or almost horizontal; all leaves
pale green on both sides, glabrous, scabrous due to short papillae, with
depressed veins and few sessile glandular hairs above, often glandular
beneath with prominent pale midrib. Capitula about 30-35 mm wide,
with many florets. Involucre broadly cupuliform, 18-22 mm long, at
base somewhat arachnoid-hairy or completely glabrous, with narrow-
linear, almost equal, contorted lax bracts; inner bracts usually erect.
Corolla pink-purple, 12-13 mm long, abruptly narrowed over 5-6 mm,
with 4-5 mm long lobes of limb. Pappus about 10-12 mm long. smoky,
pappus bristles not connate into ring, serrate-barbate, with sparse barbules
as long as thickness of bristle or shorter; achenes about 5 mm long,
pale, glabrous, oblong, compressed, four-angled, with longitudinal parallel
striations on edge, nonuniformly toothed above, with pappogen in center
of tip, and nectary on it. Flowering VII—VIII; fruiting VIII.

Stony places in meadows in the subalpine zone.—Caucasus:
Ciscaucasia, western Transcaucasia (northwest). Endemic to the
mountains of Krasnopolyansk Region. Described from the upper reaches
of the Malaya Laba River. Type in Leningrad.

Note. Three species of the section, J. sosnovskyi Grossh., J. iljinii
Grossh. and J. venusta Iljin, with glabrous incised leaves, are distributed
in the adjoining mountainous regions; the species described from the
upper reaches of the Malaya Laba River and from the Aigbi Mountains,
J. sosnovskyi, is found in the region of the Rocky Range, along the
Bolshaya Laba River, and J. iljinii Grossh. along the Urup River; all
the species are narrow endemics.

Series 4. Pinnatilobae Iljin—Leaves with broad, usually ovate lobes.

28. J. sosnovskyi Grossh. in Dokl. Akad. Nauk AzerbSSR, III, 4
(1947) 165.—Ie.: Grossh. Ibid. 166 (only leaf).

575

569

Perennial. Plants up to 20-25 cm high, with branched rhizomes.
Stem ‘solitary, scape-like, almost two times as long as leaves, erect,
brownish, cylindrical, sulcate-ribbed, densely covered with numerous
hairs, eglandular. Leaves 5—14 cm long, oblong-lanceolate, pinnatipartite,
with ovate or oblong-ovate, obtuse lobes, usually with scarcely visible
one or two lobules at base, with broadly winged axis, usually somewhat
obliquely spreading, green, with pale midrib, and smilar hairs on both
sides, as on stem, at isolated places with sparse groups of sessile glandular
hairs beneath. Capitula solitary. Involucre cupuliform, 15-20 mm wide;
involucral bracts four- or five-rowed, linear-lanceolate, acuminate, lax,
squarrose; inner row of bracts erect, with long cusps and scattered sessile
glandular hairs. Corolla about 15 mm long, narrowed over 6 mm, with
lobes of limb about 5 mm long. Pappus smoky, with few bristles, bristles
serrate, not connate into ring; achenes 3-4 mm long, smooth, glabrous,
brownish in upper half, whitish-yellowish in lower, with distinct
longitudinal ribs and corona of long teeth above. Fruiting IX. (Plate
XLII, Fig. 1.)

Rocks of high mountain belt.—Caucasus: Ciscaucasia. Endemic.
Described from the Bolshaya Laba River Basin from the Rocky Range.
Type in Leningrad.

29. J. annae Sosn. in Zam. Sist. Geogr. Rast. Tbil. 13 (1947) 61;
Shkhiyan in Fl. Gruzii, VIII, 455.—Ic.: Sosn. op. cit. Fig. on page 64.

Perennial. Plants 12-15 cm high. Rhizome vertical, covered with
broad remains of old leaves; stem scapose, densely white-tomentose,
usually with single filiform, small, white-tomentose leaf slightly below
capitulum. Leaves only basal, distinctly shorter than stem, lyrate or
almost pinnatipartite, oblong-obovate or oblong-elliptical, former with
large obtuse terminal and usually short-cuspidate and semicircular lateral
lobes, narrowed toward petiole; latter with similar but usually oblong
and also obtuse or boldly-sinuate lateral lobes, approximate in upper
one-third, reduced toward base and separated from each other, green,
glabrous, and punctate above, sometimes arachnoid-hairy basally along
midrib, with depressed veins, densely white-tomentose beneath, with
sessile glandular hairs hidden under tomentum, short petiolate, petiole
white-tomentose, almost one-third as long as lamina. Capitula solitary
terminal, with many florets, 20-25 mm wide. Involucre broadly
cupuliform, 17-20 mm long, floccose-arachnoid-hairy at base, involucral
bracts erect, but slightly divaricate, linear-subulate; outer bracts slightly
shorter, all acuminate, innermost bracts long and almost aristate. Corolla
pale pink, 18—20 mm wide, abruptly narrowed over 8—9 mm, with lobes
of limb about 6 mm long; apical appendages of anthers about 3 mm
long, acuminate. Pappus snow-white, 13-15 mm long, 8-10 times as

576

570

long as immature achenes, pappus bristles not connate into ring, serrate-
barbate, with barbules as long as or shorter than thickness of bristle;
achenes immature, pale, glabrous, four-angled.

On limestone rocks.—Caucasus: Eastern Transcaucasia. Endemic.
Described from Georgia, from the vicinity of the village of Chmi, from
the Didneua Kom Ravine. Type in Tbilisi.

30. J. levieri Alb. in Bull. Herb. Boiss. II (1894) 257; Prodr. Fl. ,
Colch. (1895) 145; Shkhiyan in Fl. Gruzii, VIII, 455; Sosn. in Zhurn.
Russk. Bot. Obshch. 11, 1-2, 199; Grossh. Fl. Kavk. IV, 171.—Ie.: Fl.
Gruzii, VIII, 410; Iljin, Kauchuk 1 Kauchukonosy, II (1953) 501, Fig. 1;
Alb. op. cit. (1894) tab. IX.

Perennial. Plants usually 12—25 cm high, with rather long rhizome,
covered with hardened dark brown remains of old leaves. Stem solitary,
scape-like, erect, cylindrical, sulcate-cylindrical, weakly arachnoid-hairy,
brownish. Leaves rosulate, gradually narrowed into petiole, shorter than
stem, lyrate, with large, oblong or oblong-ovate, short-cuspidate terminal
lobe, narrowed towards base in ever decreasing, obtuse, ovate lobes or
lobes equally long in middle part of lamina and oblong-obovate and
reduced only at base, or lamina at extreme tip undivided, obtuse, not
incised below into large teeth or oblong lobes, sometimes, in turn,
somewhat toothed; all leaves green, glabrous, lustrous, and finely rugose
above, white-tomentose beneath, with numerous sessile glandular hairs
under tomentum, and prominent ferruginous midrib. Capitula solitary,
with many florets. Involucre cupuliform; outer bracts herbaceous, usually
arachnoid-hairy, acuminate, arcuate and somewhat irregularly curved;
innermost bracts erect, gradually narrowed into long membranous cusp,
almost glabrous. Receptacle covered with smooth acuminate scales up
to one-third as long. Corolla pink-purple, with sessile glandular hairs
about 15 mm long, narrowed over 6 mm, with 44.5 mm long lobes of
limb. Pappus distinctly exceeding achene, smoky, with serrate-barbate
hairs and dense barbules, not connate into ring; achenes about 4 mm
long, light-colored, toothed above, wtih teeth modified into weakly
developed parallel ridges along borders. Flowering VII; fruiting VIII.

Limestone rocks in alpine belt——Caucasus: Ciscaucasia. Endemic.
Described from Akhzkhcha in Karachaevo-Cherkesskaya Autonomous
Region. Type in Leningrad.

Series 5. Humilae Iljin——Plants short, sometimes stemless; leaves
divided into more or less approximate segments; outer involucral bracts
arcuately recurved.

31. J. coronopifolia Somm. and Lev. in Tr. Peterb. Bot. Sada, XIII
(1893) 48; XV (1900) 270; Sosn. in Zhurn. Russk. Bot. Obshch. XI,

Si,

Sal

1-2, 199; Grossh. Fl. Kavk. IV, 171; Fl. Gruzii, VII, 456.—J. filicifolia
var. akinfiewii Winkl. in sched. ex Sosn. l.c.—Ic.: Tr. Peterb. Bot. Sada,
XVI, Plate XXVIII.

Perennial. Plants 5—20 cm, less often up te 25-30 cm high, with
long, creeping, reddish-brown rhizomes, covered with large dark brown
scales. Stems scape-like, erect, sulcate-ribbed, sparsely and finely
arachnoid-hairy, glabrescent, with scattered sessile glandular hairs. Leaves
rosulate, gradually narrowed into petiole, usually sparsely pinnatisect,
fairly narrowly winged, with narrow linear or oblong segments, entire
or in lower part toothed or with short lobes, somewhat antrorse, less
often horizontal, narrowly convolute, green, usually glabrous above, less
often weakly arachnoid-hairy, lustrous, usually weakly rugose, with
depressed veins and scattered sessile glandular hairs in pits, white-
tomentose beneath or canescent, with prominent veins, densely covered
with sessile glandular hairs. Capitula solitary, with many florets. Involucre
cupuliform, 20-30 mm wide, weakly arachnoid-hairy or almost glabrous,
with or without sessile scattered glandular hairs, involucral bracts
imbricate, four- or five-rowed; outer bracts with long herbaceous apices,
arcuately recurved; bracts of inner row narrowed from broader
membranous-frill and purple lower half into thin, usually aristate and
often flexuous, membranous apex. Receptacle densely covered with
smooth, long, acuminate, scales. Corolla pink, glabrous, 14-17 mm long,
narrowed over 5—7 mm, with 4—7 mm long lobes of limb. Pappus smoky
or smoky-pink, 12-13 mm long, with unequal bristles, serrate-barbate,
with dense barbules, not connate into ring and not divided; achenes up
to 3 mm long, obpyramidal, four-angled, pale, glabrous and smooth,
with corona of elongate teeth above, modified at border as indistinct
ridges, pappogen at apex cupuliform. Flowering VI—VIII; fruiting VIJ-—
IX.

Alpine and subalpine belts, on rocks, limestone deposits—Caucasus:
Ciscaucasia, western Transcaucasia. Endemic. Described from Kuban.
Type in Florence; cotype in Leningrad.

Note. This species occurs in two forms: pappus and achenes with
few pale bristles (f. paucisetacea Iljin), or with smoky-pink, dense bristles
on the pappus (f. multisetacea Iljin). The latter plants have thicker
achenes.

32. J. cartaiiniana Boiss. Fl. or. Suppl. (1889) 311; Grossh. FI.
Kavk. IV, 170; Fl. Gruzii, VIII, 452; Sosn. in Zhurn. Russk. Bot. Obshch.
11, 1-2, 199.

Perennial. Plants 5—25 cm high, caespitose, with strong rhizomes,
covered with rather hardened remains of old leaves. Stem scape-like,
cylindrical, usually erect, sulcate-cylindrical, slightly arachnoid-hairy,

578

TAZ

with scattered sessile glandular: hairs, always longer than leaves,

sometimes several times as long. All leaves rosulate, gradually narrowed

into petiole, with oblong-lanceolate, oblong-obspatulate, or oval lamina,
pinnatipartite or pinnatilobate, to almost undivided, with oblong or
roundish lobes, coalescing near leaf rachis forming winged axes, green
or greenish, finely arachnoid-hairy, with white arachnoid-hairy midrib
above, finely rugose, with sessile glandular hairs in pits, white-tomentose
beneath, with numerous sessile glandular hairs and ferruginous midrib.
Capitula relatively small, with many florets. Involucre cupuliform, 20—
25 mm wide, weakly arachnoid-hairy in lower half, with scattered
glandular hairs; outer bracts lax, herbaceous-green, arcuately deflexed,
middle bracts acute, inner erect, acuminate, purple. Receptacle covered
with smooth acuminate scales, almost up to half as long as involucre.
Corolla pink-purple, with sessile glandular hairs, about 14 mm long,
narrowed over 6 mm length, with about 4 mm long lobes of limb;
apical appendages of anthers 2.5—-3 mm long, acuminate, basal
appendages caudate, 2.5—3 mm long. Pappus two times as long as achenes
(including longest bristles up to apex), yellowish-whitish, bristles sparsely
serrate, not connate into ring; achenes 4.5—-5' mm long, glabrous, light
brownish, lighter below with toothed border, teeth of corona continued
along border as parallel striations. Flowering V—VI; fruiting VI-VIII.
(Plate XLII, Fig. 3.)

Rocks in high mountains.—Caucasus: Ciscaucasia. Main range,
eastern and southern Transcaucasia. Endemic. Described from Kartalinia,
between Akhaltsikh and Borzhomi. Type in Finland; cotype in
Leningrad.

32a. J. filicifolia Boiss. Fl. or III (1875) 582; Sosn. in Zhurn. Russk.
Bot. Obshch. 11, 1-2, 198; Grossh. Fl. Kavk. IV, 170; Fl. Gruzii, VIII,
451.—Ie.: Fl. Gruzii, VIII, Plate 409.

Perennial. Plants 3—5 cm, less often up to 10 mm high, with branched
rhizomes, caespitose, stemless or with slightly developed, scape-like,
arachnoid-hairy stems. Leaves rosulate, exceeding stem, or less often as
long, oblong, more or less short-petiolate, pinnatisect or pinnatipartite,
with oblong-ovate or ovate, less often entire lobes, usually obtusely
toothed or sinuate, reduced both upward and downward on lamina,
approximate on rachis with sinuses between them smaller than width of
lobes, coalesced at base forming: winged rachis, green, weakly arachnoid-
hairy, glabrescent, somewhat rugose with depressed veins and scattered,
very small, sessile glandular hairs above, white-tomentose beneath, also
with fine sessile glandular hairs under tomentum. Capitula medium, with
many florets. Involucre cupuliform; involucral bracts five-rowed, linear,
outer bracts arachnoid-hairy, herbaceous-green in upper half, acute and

579

575

arcuately recurved; inner bracts erect, narrowed into membranous finely
arcuate apex. Pappus 12-15 mm long, smoky with numerous serrate-
barbate bristles, not connate into ring, and not shedding off at fruiting;
achenes 3 mm long, grayish, obpyramidal, four-angled, glabrous, toothed,
each tooth with two or three denticles, modified as ridge. Flowering
VI-VII; fruiting VITI-IX.

Rocks in alpine belt—Caucasus: Ciscaucasia (main range). Endemic.
Described from Alagir near Ardon River. Type in Leningrad.

Section 3. Stenocephalae Benth. and Hook. Gen. pl. III (1873) 473;
Pflanzenfam. IV, 3, 321—§1. Linearifoliae Boiss. Fl. or. II] (1875).—
§3. Pinnatae |.c. 568 (quoad J. pulchellam DC.) p. p.—Plants branched
at anthesis; leaves undivided and entire (excluding J. pulchella DC.);
involucre narrow, with nonprickly bracts; pappus comprising serrate or
serrate-barbate bristles, not connate into ring (see, however, J. kapelkini
O.Fedtsch.), teeth approximate. About six species, of which four species
in the USSR.

lj Acowemlleaves pinnatisect=.¢s:..Nenis.te.tpeitinse. come. ete
CN ogtah subst meated. die 35. J. pulchella (Fisch. and Mey.) DC.
at All leavesundividedvand entire. 2 ance le. ee 2:
2. Capitula solitary; pappus deciduous; pappophorus present............
we I ME es Sh J PRI Ue NAT ee oe 36. J. kapelkinii O. Fedtsch.
+ Capitula in corymbose inflorescence; pappus persistent; pappophorus
absenityes wee his. bee Aaa. es ona, Alp Seat! ana eteies 3:
3. Capitula in simple corymb; leaves filiform or linearly filiform,
sessile, nondecurrent; involucre white arachnoid hairy-flocculose;
corolla usually 16-18 mm long; pappus 15—20 mm long; achenes
1=71.5-mm long .glabrous; Smoothicd..tieh sc tetelas. aca
Fae ease ae aeveh Lake vas, Risse, eevee aaonae sige! 34. J. stoechadifolia (MB.) DC
+ Capitula in compound corymbs; leaves linear or lanceolate,
sometimes even oblong-ovate, sessile, usually semiamplexicaul and
slightly decurrent; involucre glabrous or only slightly farinose-
arachnoid-hairy, 10-15 mm long; corolla 13-17 mm long; pappus
far exceeding achene; achenes 3.5—5 mm long, dark brown, often
with white-cartilaginous ribs, transversely rugose ..............0:c0e
bE aetiaids 3 a Listssstssseteseeessesseeeeeee 33. J. multiflora (L.) B. Fedtsch.

Note. Besides the species treated above, the Iranian species J.
heterophylla (Jaub. and Spach.) Boiss. is also included in this section.
Boissier assigns J. pulchella DC. to section Pinnatae Boiss., which,
furthermore, includes extremely unlike species.

Series 1. Linearifoliae Iljin—Leaves undivided with entire margin.

580

574

33. J. multiflora (L.) B. Fedtsch. in O. and B. Fedtsch. Perech.
Rast. Turk. IV (1911) 295; Ijin in Fl. Yugo-Vost. VI, 389; Vizn. Rosl.
URSR 564; Krylov, Fl. Zap. Sib. XI, 2930.—J. linearifolia DC. Prodr.
VI (1837) 675; Ldb. Fl. Ross. II, 761; Boiss. Fl. or. II], 571; Schmalh.
Fl. II, 109; Grossh. Fl. Kavk. IV, 169; Iljin in Tr. Turk. Nauchn. Obshch.
II, 16; Sosn. in Zhurn. Russk. Bot. Obshch. 11, 1-2, 194—J. ambigua
DC. Prodr. VI (1837) 675 non al.—Serratula multiflora L. Sp. pl. (1753)
1145: MB. FI. taur.-cauc. II, 265; II], 547; Ldb. Fl. alt. IV, 41.—S.
salicina Pall. It. II (1776) 682.—S. salicifolia Lepech. It. 1 (1795) 262.—
Serratula ambigua DC. in Ann. Mus. Hist. Nat. XVI (1810) 186.—S.
tincta Cass. in Dict. sc. nat. XLVI (1826) 476.

Perennial. Plants 10-50 cm high. Rhizomes slender, long, usually
branched; stems several or solitary, erect or bent, ribbed-sulcate, green
or dirty purple, whitish-gray throughout or only below due to arachnoid
hairy-flocculose pubescence, with scattered sessile glandular hairs,
branched in inflorescence. Leaves lanceolate, oblong, or linear,
acuminate, green, less often finely arachnoid-hairy, and scabrous above
due to numerous short papillae, less often without papillae, slightly
rugose, with scattered, very small sunken glandular hairs, ash-gray or
whitish beneath due to fine arachnoid-hairy tomentum, less often green
on both sides (f. subglabra Serg.), with prominent midrib, undivided
and entire, occasionally lower and middle leaves sparsely serrate (f.
serrata I\jin); cauline leaves sessile, often amplexicaul, sometimes
auriculate (f. auriculata Iljin), shortly or even strongly decurrent,
sometimes lateral (f. ambigua I\jin); basal and lowermost cauline leaves
gradually narrowed in petiole, all involute often up to midrib. Capitula
most numerous, in corymbose inflorescence, small. Involucre cylindrical,
slightly narrowed at base, usually 10-15 mm long and 3-5 mm wide,
glabrous or slightly farinose-arachnoid hairy, pale or purple; involucral
bracts with dark green or purple prominent midrib, membranous
along margin; outer bracts oblong-ovate, subobtuse, middle bracts
oblong, inner lanceolate-linear, acuminate; all bracts appressed, with
scattered sessile glandular hairs. Receptacle covered with long, smooth,
acuminate scales, reaching to one-third of involucre or even to its tip.
Corolla pale pink or purple, less often creamish, 13-17 mm long, more
or less abruptly narrowed into 5—7.5 mm long tube, with 4-5 mm long
lobes of limb, having scattered sessile glandular hairs; apical appendages
of anthers abruptly narrowed, about 2 mm long; basal appendages
caudate, about 1.25-1.5 mm long; stigma 1.25—1.5 mm long; style
glabrous; filaments glabrous, smooth. Pappus snow-white, 8-12 mm long,
with serrate bristles, dense, shorter than its thickness, not connate into
ring, persistent; achenes 3.5-5 mm long; obpyramidal, four-angled,
glabrous, smooth, dark brown, often with white cartilaginous ribs, at

581

Te)

edges mildly transversely rugate-rugose, with sharp-toothed corona having
filiform pappogenus in center, and nectary at tip. Flowering VI-VIII;
fruiting VII-IX.

Forest-steppe, steppe and semidesert zones, where it grows in feather-
grass covered, stony, saline steppes and saline meadows, on wastelands
and among crops.—European Part: Bessarabia, Black Sea Region,
Crimea, Middle Dnieper, Volga-Don, Lower Don, Volga-Kama, Trans-
Volga Region, Lower Volga; Caucasus: Ciscaucasia; Western Siberia:
Upper Tobol, Irtysh, Altai; Eastern Siberia: Angara-Sayans (south).
General distribution: Western China (Sinkiang). Mongolia. Type in
London.

34. J. stoechadifolia (MB.) DC. Prodr. VI, 674; Ldb. Fl. Ross. II,
762; Boiss. Fl. or. II, 571; Schmalh. Fl. II, 109; Vizn. Rosl. USSR,
564; Grossh. Fl. Kavk. IV,°169; Sosn. in Zhurn. Russk. Bot. Obshch.
11, 1-2, 194.—Serratula stoechadifolia MB. Fl. taur.-cauc. II (1808)
266.—S. salicifolia Willd. in herb. no. 14853.—S. affinis Pall. Ind. Taur.

(1801).—Iec.: MB. Cent. pl. rar. tab. 49; Prodan, Descr. Pl. Romaniae,

113(1923); tab:38:

Perennial. Plants 15-40 cm high, developing numerous stems; stems
erect, whitish-grayish arachnoid-hairy, sulcate-cylindrical, with scattered
small sessile glandular hairs, branched only in inflorescence. Leaves
linear, undivided and entire, cuspidate, with callose thickening at apex,
scabrous above due to dense papillae, with numerous small glandular
hairs in depressions, with depressed midrib, green, glabrous or weakly
arachnoid-hairy, usually strongly involute, white-tomentose beneath, with
prominent greenish or brownish midrib; basal leaves in dense rosettes,
up to 7 cm long; cauline leaves reduced above up to capitula. Capitula
in corymbs, with many florets. Involucre cylindrical, 15-20 mm long,
whitish arachnoid-hairy-floccose; outer bracts oblong-ovate, middle more
oblong, with obtuse, prominent, or entirely dark green or dirty purple
midrib, subobtuse or subacute, all bracts imbricately appressed, five- or
six-rowed; innermost bracts oblong-lanceolate, membranous in upper half,
silvery white or purple, covered with small appressed bristles and sessile
glandular hairs, along margin and often also at apex ciliate-fimbriate.
Receptacle covered with smooth, glabrous, curly scales, up to half as
long as involucre. Corolla pink-purple, with scattered sessile glandular
hairs, (14)16—-18 mm long, sharply narrowed over 7—9 mm, with 4-6
mm long lobes of limb; apical appendages of anthers 2-3 mm long,
gradually narrowed above, often purple, basal appendages caudate,
flexuous, 1.5—2.5 mm long; stigmas with very small papillae on outer
side; style glabrous. Pappus almost as long as achene, with few prominent
bristles, up to 10-12 mm long, with very dense, serrate bristles not

582

576

connate into ring, divaricate during fruit-maturation; achenes 7—7.5 mm
long, oblong, four-angled, with small teeth above, milky-white along
teeth, upper part, ribs, and along edges below, dark brown in upper half
of edges, glabrous, smooth, with somewhat prominent, longitudinal,
parallel striations, with pappogenus in the center of tip, terminating in
five-toothed nectary. Flowering VI—VII; fruiting VII-IX.

Stony places and limestone deposits, chalks, steppes and dry pine
forests—European Part: Crimea, Black Sea Region(?), Lower Don,
Volga-Don (south); Caucasus: Ciscaucasia, western Transcaucasia (south).
General distribution: Bulgaria, Romania. Described from Karasu-Bazar.
Type in Leningrad.

Note. Very recently, the Lower Don populations of this species have
been described as a separate race, J. brachycephala Klok. (Tr. Sp. Rosl.
Gerb. FI. URSR, Tsent. Il, 1962, 32, No. 197).

Series 2. Pinnatae Iljin—Lower leaves pinnatisect.

35. J. pulchella (Fisch. and Mey.) DC. Prodr. VI (1837) 676; Boiss.
Fl. or. III, 572; Trautv. in Tr. Peterb. Bot. Sada, II, 456; Grossh. FI.
Kavk. IV, 169; Sosn. in Zhurn. Russk. Bot. Obshch. 11, 1-2, 194.—
Serratula pulchella Fisch, and Mey. ex DC. Prodr. l.c.

Perennial. Almost semishrub; plants 25-60 cm high. Stems few,
erect, virgate, ribbed-sulcate, whitish-grayish-arachnoid-hairy throughout,
covering numerous sessile glandular hairs, less often simple, usually
branched, with upward spreading branches. Basal and lower cauline
leaves pinnatisect into long, linear, distant, usually obliquely opposite
segments, which, in turn, sometimes with short sparse appendages, short-
petiolate, sharply erect; upper leaves linear, undivided and entire, sessile,
densely appressed, reduced upward, not reaching capitula, with callose
thickening at apex; all leaves grayish due to fine arachnoid-hairy
pubescence, involute, reticulate-rugose above, with numerous, deeply
sunken sessile glandular hairs in pits, with prominent lustrous midrib,
more dense pubescence and less distinct glandular hairs beneath. Capitula
numerous, in corymbs. Involucre narrow-cupuliform, usually 15—18(20)
mm long, grayish due to fine flocculose pubescence; involucral bracts
compactly appressed, with scattered sessile glandular hairs and prominent
obtuse midrib, greenish or purple in outer bracts, and dirty purple in
middle and inner bracts, with two less prominent lateral veins; outermost
bracts small, oblong, gradually broader inward, innermost acuminate,
membranous above; all bracts finely divided along margin. Receptacle
covered with smooth acuminate scales, almost up to half as long as
involucre. Corolla pink, 17-20 mm long, narrowed over 7—10 mm length,
with 4-6 mm long lobes of limb, and scattered sessile glandular hairs;
apical appendages of anthers 3—-3.5 mm long, abruptly narrowed at apex

583

577

in a short cusp; basal appendages membranous, divided below, 2—2.5
mm long; stigma finely papillose on outer side, not divided up to base,
with tuft of short, 2-3 mm long bristles at base; style glabrous; filaments
light colored, glabrous. Pappus persistent, about 10 mm long, of serrate-
barbate bristles, barbules often shorter than thickness of bristles,
especially in more prominent and somewhat flat bristles; bristles not
connate into ring, only outermost short bristles persisting on achene;
achenes elongate, gradually narrowed only toward base, four-angled,
about 6 mm long, glabrous, smooth, dark brown, with corona of short
teeth, pappogenus short-cylindrical, with five-toothed short nectary in
the center of tip. Flowering V—VIII; fruiting VIII-IX.

Stony slopes of mountains, limestone rocks.—Caucasus: Southern
Transcaucasia. General distribution: Iran. Described from Azerbaidzhan,
in the vicinity of Koi from the collections of Szovits. Type in Leningrad.

36. J. kapelkinii O. Fedtsch. in Tr. Pochv.-Bot. Eksp. Pereselench.
Upravl. II, 5 (1910) 44; O. and B. Fedtsch. Perech. Rast. Turk. IV, 297;
Iljin in Tr. Turk. Nauchn. Obshch. 2, 4.—Ic.: O. Fedtsch. op. cit.

Perennial. Plants 8-20 cm high, almost semishrubs, producing leafy
vegetative branches as well as erect, fertile, simple stems or even
arachnoid stems with solitary branches, leafy in lower two-thirds. Leaves
narrow-linear, entire, with callose thickening at apex, strongly revolute,
green, slightly rugose, and weakly arachnoid-hairy above, white-
tomentose beneath, sessile, reduced toward upper side; basal leaves
numerous. Capitula solitary. Involucre cupuliform, 10-12 mm long and
8-10 mm wide at base, young involucre somewhat arachnoid-hairy,
roundish at base; outer bracts appressed, lanceolate, brown or dirty-
purple, densely covered with short appressed bristles and sessile glandular
hairs. Receptacular scales half as long as involucre or slightly longer;
corolla pale pink, 12-15 mm long, narrowed over 5—6 mm of its length,
with 3.5—4 mm long lobes of limb, with scattered sessile glandular hairs;
apical appendages of anthers 2—2.5 mm long, abruptly narrowed in cusp.
Pappus deciduous, 6-8 mm long, white, barbate, with dense barbules,
almost as long as thickness of bristles; bristles connate into ring, with
entire membranous base, bearing cupulate pappogenus with small finely
toothed nectary in center; achenes elongate, four-angled, 4.5—-5 cm long,
glabrous, with small-toothed corona, distinctly rugose on edges, with
dome-shaped pappophorus. Flowering at end of VI-—VII; fruiting VI-
VII.

Gravelly slopes of hills, saline and sheep’s fescue-feather grass
steppes, exposures of white clay—Western Siberia: Irtysh (southwest);
Soviet Central Asia: Balkhash Region. Endemic. Described from Borla-
Chabra Hill in the Atbasar Region. Type in Leningrad.

Note. Hybrids with J. multiflora (L.) B. Fedtsch. occur.

584

578

Section 4. Integrae Ijin sect. n. in Addenda XVI, 721.—Stem more

or less winged; leaves undivided and entire, occasionally ternate,
decurrent leaves found between undivided leaves; achenes glabrous,
smooth, often scabrous or with smooth striations and ribs, usually with
ring of teeth under corona; pappus usually shedding entirely together
with pappogenus (except in J. kokanica Iljin, J. rhizomatoidea Iljin,

and

J. trifurcata \\jin, in which the pappus is persistent). Plants

predominantly of the Tien Shan Mountain system. About 12 species in
the USSR.

nt & +t Ww

Type of section: J. abolinii Ijin.

. Involucral bracts equal, except innermost, apically terminating in

long, filiform, herbaceous, flexuous appendage (western Tien Shan)
BN 2 GS ES se Ne, Ae eh hes | Gans et 43. J. tortisquamea I[ljin
All involucral bracts without herbaceous flexuous appendage......

vested usu cage tes cuspees ecaseet acess itrae trarteuese nee Meera Seen eee ae 2
7 all involucral bracts, APPRESSEd coc sea eas cccaecscteean cee ie eee eee 3.
Outermost involucral bracts slightly recurved or lax and deflexed
RE GRE SE rT NRE Meat ne SARE Oe SANA SIEM gil 10.
IwGAVESpm@t sp CCUITC I Pc ci aR A Me can aga ea 4
All. caubine, leaves eCuntertit.: sds sccm eee, bt becn cn cuetbebe: borage tara 7
Stem with more or less evenly distributed leaves..................... 5p
béaveswmamlyangrosettesats.fverutiors a lieeee: Ualtg och mel bye alien 6.

. Plants strongly branched; leaves undivided, sessile, with roundish

base, narrowly linear, green on both sides; involucre cyathiform,
achenes very small, gradually narrowed below, glabrous, smooth,
less than» “mmlong(PashaatasRawine): Statins. uhh. lees
i RE RE, SUIS: RY SOROS CS TPR 38 39. J. stenophylla [jin
Leaves undivided, white-tomentose beneath, green above; involucre
cup-shaped; achenes about 6 mm long, glabrous and rugose-

tuberculate "above eae eas a ee 40. J. caespitans [lyin
Achenes two times as long, convex below corona, leaves ternately-
CIVIGER cere et re ee 48. J. trifurcata [jin

. Leaves oblong or lanceolate, always entire; achenes below corona

with scattered teeth (Buam Ravine) .............. 37. J. abolinii [jin
Leaves obspatulate or oblong-obspatulate, sometimes mixed with
lanceolate, entire or with isolated teeth at base of lamina; achenes
toothed to two-thirds to three-fourths of their length (Alai Range)
beams Menem eh pe rca de ahlhey ehcan 37. J. schachimardanica [jin

. Leaves arachnoid-hairy beneath, sometimes toothed......................

pected utp ak 2 Riana ty aonb tn 42. J. capusii Franch.

585

579

8. Pappus persistent; achenes glabrous, with row of teeth only below
corona (Turkestan Range) ............:::cseeeeeee 46. J. kokanica I[ljin
+ Pappus deciduous, all bristles connate into ring ..............0.+ S.
9. Plants large, with 30-40 mm long capitula and large basal leaves;
pappus bristles plumose-barbate (Darbaza-Bel’ Pass in Pamiro-Alai)
Lbbi cep Eelam oe 55 4a ANSE SE Ea et Oi che 45. J. macranthodia []jin
+ Capitula half as long: pappus bristles serrate-barbate, with
dense barbules (western Tien Sham) ...............:ccccessceseeeseeseeseeeeeeees
Ne wins) heen LA oe ES eR Rees OAS alee 41. J. eduardi-regelii igh
10. Plants erect, with well developed fibrous roots; leaves oval to
lanceolate; outer involucral bracts somewhat recurved; pappus
shedding entirely; achenes of unclear structure (central Tien Shan)
PE CEE ATE TIE, PELARRELELC EE Lk ERD See Eee 44. J. androssovii [jin
+ Plants with long creeping rhizome, covered with membranous
scales; stem ascending; leaves linear, thin; involucral bracts light
green, translucent, divergent, outer bracts recurved; pappus
persistent; achenes about 4 mm long, toothed above below corona
(Rarataw)! 2.002 Pe 47. J. rhizomatoidea [jin

Series 1. Scapiformes I[ljin—Plants with short stem and rosulate
leaves.

37. J. abolinii Iljin in Izv. Glavn. Bot. Sada, XXVII, 1 (1928) 83.

Perennial. Plants 15-20 cm high, with strong woody tap root,
converted at soil surface in many-headed caudex bearing rather large
number of shoots. Stems woolly-tomentose at base, with remains of
fallen leaves, simple, erect or weakly ascending, flocculose-arachnoid,
leafless in upper part. Basal leaves numerous, oblong or lanceolate, acute,
undivided and entire, sometimes somewhat hairy, gradually narrowed in
somewhat short petiole, pale greenish above, at some places flocculose-
arachnoid-hairy, grayish beneath due to arachnoid-hairy pubescence; lower
cauline leaves like basal, upper sparse, linear, sessile. Capitula solitary.
Involucre up to 15 mm long, cupuliform, with glabrescent flocculose-
arachnoid-hairy pubescence at base, other parts glabrous, green; involucral
bracts lanceolate, acute; middle bracts lanceolate or linearly lanceolate,
gradually acuminate; inner bracts prominent, with prominent veins,
acuminate. Receptacle densely squamose, scales up to one-third as long
as involucre. Pappus snow-white, up to 5 mm long, usually of few bristles,
bristles short-barbate, barbules dense, approximately as long as thickness
of bristle; achenes 5—6 mm long, oblong-obpyramidal, with obtusely
roundish ribs above and here somewhat swollen, glabrous and small
toothed above along edges. Flowering VI(?); ne VII. (Plate XLV,
Fig.’ 2.)

586

580

Red gypsum sandstones.—Soviet Central Asia: Tien Shan. Endemic.
Described from the Buamsk Ravine. Type in Leningrad.

38. J. schachimardanica Iljin in Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XXI (1961) 386, Fig. 4.

Perennial. Plants about 20-25 cm high, caespitose, with strong tap
root and aerial stem, densely covered with brown scales and remains of
petiole. Stems few, simple or with isolated long, upward spreading
branches in the lowermost part, sparsely arachnoid-hairy, finely sulcate,
leafy below, leafless or with isolated small simple leaves above; basal
leaves forming dense, rosettes of many leaves, obspatulate or oblong- -
obspatulate, less often mixed with oval, entire leaves, laminas only at
base with single, less often two pairs of triangular or triangular-oblong
teeth, abruptly cuspidate, less often acute or acuminate, green and
glabrous above, whitish-grayish beneath due to arachnoid-hairy
tomentum, gradually narrowed in long petiole, almost as long as lamina,
somewhat broadened at base, with densely woolly bud in axil. Capitula
solitary, with many florets. Involucre cupuliform, 10—10.25 mm long,
three- or four-rowed, with rather loosely arranged bracts; involucral bracts
with three prominent veins, oblong-lanceolate, acuminate, thin-
membranous along margin, somewhat serrate, puberulent above (in -
inner bracts), sparsely arachnoid-hairy; innermost bracts gradually
narrowed into membranous reddish awn. Receptacle covered with short
(about 3 mm long), smooth, acuminate scales. Corolla 11.5—13.5
mm long, on 3.5—5 mm long stalk and 2.5-3.5 mm long lobes of
limb, slightly incurved above, pink, with few, scattered sessile glandular
hairs; apical appendages of anthers about 2.5 mm long, whitish, narrowly
attenuate at apex and slightly incurved; basal appendages setaceous,
slightly divided, about 2 mm long; filaments glabrous; stigma about 2
mm long, bifid up to base, covered with rather short papillae, basally
with a brush of short, erect bristles; style glabrous. Pappus colorless,
consisting of 4-5 mm long unequal bristles and 4-10 mm long bristles
with very dense barbules almost as long as or usually one and one-half
times as long as thickness of bristle, pappus shedding, leaving
inconspicuous annulus with fused bases of bristles, except outer, shortest
and persistent; achenes oblong-obpyramidal, 5—6 mm long, four-angled,
olive-brown, lighter along ribs, with uniformly toothed border at apex,
edges slightly convex, covered throughout with rows of one-toothed
scales connate adbasal to achene. Flowering VI; fruiting VII.

Sandstones, conglomerates and products of their weathering —Soviet
Central Asia: Pamiro-Alai. Endemic. Described from the Kara-Kazyk
River Basin, from the vicinity of the Shahimardan Resort. Type in
Tashkent.

587

581
Series 2. Sessiles Iljin—Leaves sessile, nondecurrent.

39. J. stenophylla Iljin in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1,
1 (1933) 185.

Perennial. Plants up to 25-30 cm high, bright green, many-stemmed.
Stems slender, in lower part up to 2 mm, in upper part up to 0.75 mm
thick, branched from base, with spreading branches, not branched above,
glabrous or slightly and sparsely arachnoid in upper part only in very
small patches, brownish basally with scaly recurved remains of leaves,
densely leafy in middle. Leaves linear, acuminate, undivided and entire,
strongly revolute, almost up to midrib, one-veined, concolorous, with
scattered, sunken, very fine glandular hairs, sessile, with slightly roundish
base. Capitula solitary, on long leafless peduncles. Involucre cyathiform,
distinctly narrowed toward base, 12-15 mm long, broadened up to 10—
12 mm in width at fruiting, yellowish-brownish, greenish with florets,
glabrous and only scarcely flocculose-arachnoid at base. Involucral bracts
oblong-lanceolate, acuminate, appressed, with membranous border;
innermost bracts usually pinkish-purple. Receptacle squamose; scales
finely pointed, yellowish, smooth, almost half as long as involucre.
Pappus about 15 mm long, whitish, barbate, with barbules one and one-
half times as long as thickness of bristles or almost as long, dense;
achenes (immature) very small, glabrous, smooth, four-angled, with small
teeth above along edge. Fruiting IX. (Plate XLIII, Fig. 2.)

Rocks.—Soviet Central Asia: Tien Shan. Endemic. Described from
the ravine of the Pashaata River, Namangan District. Type in Leningrad.

40. J. caespitans Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada, V, 11—
12 (1924) 167; Ijin in Tr. Turk. Nauchn. Obshch. 2, 24.

Perennial. Semishrub, 15-30 cm high, with many-headed caudex,
many stems. Root very thick, branched, woody. Stems many, divaricately
branched from base, almost virgate, angular, sulcate, somewhat
arachnoid-hairy, greenish. Leaves oblong or lanceolate, acute or
acuminate, less often subobtuse, gradually narrowed toward base, sessile;

‘basal leaves short-petiolate, with petioles broadened at base, undivided

and entire or sometimes with one to three distant teeth; upper leaves
linear or almost filiform, gradually reduced toward stem apex, green or
grayish-greenish, punctate and weakly arachnoid-hairy, whitish or gray-
tomentose beneath with distinct midrib and few indistinct lateral veins
on both sides. Capitula on long peduncles, often slightly divergent, about
20 mm wide including florets. Involucre 15-20 mm wide, slightly
arachnoid-hairy or subglabrous, dirty purple or green and purple above,
five- or six-rowed; bracts linear-lanceolate, acuminate,with distinct veins,
almost entire, appressed (sometimes outermost bracts weakly deflexed),

588

582

outer bracts shorter. Receptacle squamose; scales more or less dense,
smooth, short. Corolla red-purple, 15—17 mm long, its narrow part one-
third as long as broad part, with 4-5 mm long limb; anther appendages
setiform, 1-2 mm long; filaments smooth. Pappus deciduous, with very
dense barbules, shorter than thickness of bristles, or almost as long,
connate at base into ring; achenes obpyramidal, four-angled, about 6
mm long, glabrous and lustrous, rugose-tuberculate, with spinescent teeth
along edges and toothed corona enclosing flask-shaped pappophorus with
five-lobed nectary at its throat. Flowering VII—VIII; fruiting VIII-IX.
(Plate XLIII, Fig. 1.)

Subalpine belt, mostly on stony and gravelly slopes.—Soviet
Central Asia: Tien Shan. Endemic. Described from the rise towards
the pass between the Kok-Bulak and Ak-Kur rivers. Type in
Leningrad.

Series 3. Decurrentes [ljin—Cauline leaves decurrent.

41. J. euardi-regelii Iljin in Tr. Turk. Nauchn. Obshch. 2 (1925)
18.—J. heterophylla auct. non Jaub. and Sp.: O. and B. Fedtsch. Perech.
Rast. Turk. IV (1911) 296.—Ie.: Iljin, op. cit.

Perennial. Semishrub, 25—40 cm high, strongly branched in lower
part, green branches of current year glabrous or glabrescent with
arachnoid hairs, simple or with few virgate branches. Leaves oblong-
lanceolate or even linearly lanceolate, less often oblong, acute or
acuminate, entire or often with remote solitary teeth, narrowly recurved,
green and glandular hairy on both sides or with thin, indistinct arachnoid-
hairy indumentum beneath; upper leaves weakly scabrous, sessile,
narrowly decurrent; lowermost leaves gradually narrowed into petiole,
uppermost strongly reduced so that stem appears leafless above. Capitula
solitary. Involucre narrow-cupuliform, roundish at base, 10—15 mm long
and 7.5-12 mm wide, green, purple above, glabrous or weakly arachnoid
hairy-flocculose; involucral bracts appressed, lanceolate, acuminate, outer
smaller. Receptacle covered with membranous hairs, one-third to half
as long as involucre. Corolla pink, 12-15 mm long, narrowed below
over 5-6 mm length, with 3.5—-4 mm long lobes of limb. Pappus
deciduous, snow-white, 9-12 mm long, with barbules equal or slightly
longer than thickness of bristles; bristles connate into ring and shedding
entirely together with pappogenus and nectary; ring extended into
membranous base, with purple pappogenus, having pitcher-shaped nectary
in center; achenes roundish to four-angled, 5-6 mm long and 3-4 mm
thick, brownish admedially, smooth, with sharply toothed corona, convex
below corona.

Mountain slopes, mostly in juniper forests—Soviet Central Asia:

583

| he
=

sett,

A Y, i XY WY

ty
Pa

On",

iy

Ve,

Plate XLIII.
1—VJurinea caespitans lljin; 2—J. stenophylla Iljin.

589

590

584

Tien Shan. Endemic. Described from western Tien Shan, from mountains
between Ak-Tag-Tai and Ak-Sagaty. Type in Leningrad.

42. J. capusii Franch. in Ann. Soc. Nat. Bot. XVI (1883) 328.—
Exs.: H.F.AM. No. 490.

Perennial. Semishrub, 15—30 cm high. Stems several, usually with
long branches in lower half, and leafy. Basal and lower cauline leaves
oblong or more often oblong-obovate, entire, narrowed into petiole,
subobtuse, other leaves narrower, lanceolate, sometimes subacute or even
acute, sessile and weakly decurrent, all leaves green and scabrous
beneath, grayish due to finely arachnoid-hairy pubescence, occasionally
almost glabrous. Capitula solitary, on long leafless peduncles. Involucre
10-15 mm long and 6-10 mm wide, glabrous, green, purple above;
involucral bracts appressed, acuminate, lowermost often oblong-ovate,
reduced, others broadly lanceolate to linear-lanceolate. Receptacle densely
squamose; scales up to half as long as involucre or slightly longer.
Corolla pink-purple, 10-16 mm long, narrowed over 3—5.5 mm length,
with 2.5-4 mm long lobes of limb. Pappus entirely deciduous, weakly
yellowish, 7-10 mm long, densely barbate, with barbules as long as
thickness of bristles; achenes about 6 mm long, smooth, with isolated
spiny tubercles and toothed edge at apex, enclosing a short conical
pappophorus. Flowering VI-VII; fruiting VIJ—VII.

Stony slopes in upper mountain zone.—Soviet Central Asia: Tien
Shan (western). Endemic. Described from Ona-Ulgan in the Chirchik
Mountains. Type in Paris.

Note. The author of this species included it among the relatives of
J. trautvetteriana Rgl. and Schm. without any basis whatsoever. These
are species from different sections. The author made another mistake by
listing both of these species under section Derderiae. Our herbarium
does not have either the authentic specimen or the cotype of J. capusii
Franch.; however, neither its description nor its locality raises any doubt
that the specimens identified by us and other collectors belong to this
particular species.

43. J. tortisquamea Iljin in Tr. Turk. Nauchn. Obshch. 2 (1925)
18; in Bot. Mat. Gerb. Bot. Sada, V, 58—Exs.: H.F.A.M. No. 496.

Perennial. Plants 50—70(100) cm high, light green, almost glabrous.
Stems branched from middle; branches virgate, with scattered, soft -
arachnoid hairs. Leaves sessile, decurrent, oblong or oblong-lanceolate,
entire, acuminate, slightly scabrous above, with sessile glandular hairs
beneath, strongly reduced on branches, distant, narrowly linear or almost
filiform. Capitula solitary. Involucre broadly cylindrical, rounded at base,
15—20 mm long and 10-15 mm wide, green, almost glabrous, all bracts

59]

585

equal, outer linear-filiform, herbaceous, sinuate; inner bracts linear-
lanceolate, acuminate. Receptacle densely covered with long scales,
connate up to two-thirds, mostly in lower part. Corolla pink, 15-18 mm
long, narrowed below over 4.5—6 mm length, with 4—5 mm long lobes
of limb. Pappus deciduous, 5—8 mm long, dirty-whitish, barbate, with
dense, sessile barbules shorter than thickness of hair, hairs connate into
ring at base with fleshy patellate pappogenus and nectary in center;
achenes smooth, yellowish smoky, 7 mm long and 2.5—3 mm thick,
toothed along top edge, convex below corona, with weakly developed
and more or less tuberculate pappophorus in center of apex. Flowering
VI-VU; fruiting VII-VIII(?).

On grassy, clayey-stony soil, slopes, in thickets of shrubs.—Soviet
Central Asia: Tien Shan (western). Endemic. Described from Brich-
Mulla, near Chimgan. Type in Leningrad.

44. J. androssovii Iljin in Bot. Mat. Gerb. Bot. Sada, V, 11-12
(1924) 168; Iljin in Tr. Turk. Nauchn. Obshch. 2, 18.—Iljin, op. cit.
(1925), Plate III.

Perennial. Plants 40—70 cm high, light green. Root somewhat thick,
woody. Stems few, simple or branched in lowermost part, weakly sulcate,
almost glabrous or covered with short scattered hairs, slightly arachnoid-
hairy below capitulum and sometimes glandular-hairy. Leaves oblong
or lanceolate, acute or subobtuse, undivided and entire, not convolute,
glabrous and green on both sides, along veins usually with rather short,
sparse hairs; basal leaves narrowed into short arachnoid-hairy petiole;
lower cauline leaves sessile, slightly narrowed toward base, narrow-
decurrent; middle and upper leaves usually unilaterally decurrent, in
upper half or sometimes at the base of two halves of lamina with roundish
auricles; one or two uppermost leaves strongly reduced, remote. Capitula
solitary, 20-25 mm long. Involucre 15-20 mm wide above, five- or six-
rowed, almost glabrous or slightly sparsely arachnoid-hairy, with
lanceolate-linear bracts, narrowed toward base; outer bracts slightly
recurved, acuminate, innermost long attenuate, with distinctly projecting
veins, appressed, rather hispid on outer side, slightly toothed above.
Receptacle densely covered with smooth hairs. Corolla pale-purple, with
scattered glandular hairs especially below; 17-18 mm long, narrowed
over 5—6 mm; basal appendages of anthers setose, 2—2.5 mm long;
filaments smooth. Pappus deciduous, barbate, of unequal bristles, with
dense barbules, slightly exceeding diameter of hair, bristles connate into
ring with hollow base; achenes four-angled, obpyramidal, yellow, toothed
above along edges, glabrous and tuberculate-spiny, particularly in upper
part, smooth near base, with dome-shaped pappophorus at tip. Flowering
VI-VII; fruiting VI—-VIII(?).

592

586

Stony slopes of mountains.—Soviet Central Asia: Pamiro-Alai
(eastern). Endemic. Described from Uch-Kurgan. Type in Leningrad.

45. J..macranthodia I[ljin in Bot. Mat. Gerb. Glavn. Bot. Sada, V,
5 (1924) 66; Tr. Turk. Nauchn. Obshch. 2, 18.

Perennial. Plants 35-55 cm high. Stems few, simple or with fewer
branches, greenish, sulcate, scabrous, leafless from middle. Leaves oblong
or oblong-lanceolate, entire, or less often with few distant, large teeth
along margin, acuminate or acute, greenish on both sides, somewhat
stiffly scabrous, especially above due to very short crisped hairs and,
besides, with numerous sunken glandular hairs beneath; basal leaves
petiolate, cauline leaves sessile, usually weakly decurrent; uppermost
leaves very distant, linear or filiform. Capitula solitary, about 30 mm
long (including florets). Involucre, cupuliform, rounded at base, broader
above, up to 30 mm, glabrous, not arachnoid-hairy, five- or six-rowed;
inner involucral bracts linear-lanceolate, acuminate, gradually reduced
outward, increasingly hispid-scabrous on inner side, not arachnoid-hairy,
with small teeth. Receptacle covered with smooth hairs. Corolla purple,
with scattered glandular hairs or almost glabrous, 20-22 mm long, on
7-10 mm long stalk, with about 5 mm long limb; anther appendages
setose, almost undivided, 2-3 mm long; filaments glabrous. Pappus
deciduous, plumose, wtih plumes several times as long as thickness of
hair, bristles connate into ring with common base, and nectary in center;
achenes immature, striate-streaked, glabrous, with scattered spinules in

' upper part, serrate-spinose, with dome-shaped pappophorus at tip.

Flowering VI; fruiting VII.

Foothills—Soviet Central Asia: Pamiro-Alai (eastern). Endemic.
Described from the Osh Region, near the Derbaz-Bel Pass along slope
to the Tara Valley. Type in Leningrad.

Series 4. Persistentes Iljin—Pappus persistent.

46. J. kokanica Ijin in Bot. Mat. Gerb. Glavn. Bot. Sada, V, 5
(1924) 66; Tr. Turk. Nauchn. Obshch. 2, 18.

Perennial. Plants 25-50 cm high, usually with distinctly developed
caudices. Root thick, branched, woody. Stems simple or weakly branched,
greenish, slightly sulcate, woolly tomentose at base and covered with
scales, almost glabrous or somewhat scatteredly hairy over short distance,
sometimes finely arachnoid-hairy at isolated places, leafless above.
Leaves basal and cauline, oblong-lanceolate or lanceolate, entire or
usually large-toothed, acuminate, basal leaves gradually narrowed into
petiole, slightly broadened at base; cauline leaves sessile, usually narrow-
decurrent; uppermost leaves strongly reduced, linear or filiform; all leaves

593

587

green on both sides or sometimes weakly grayish arachnoid-hairy
beneath; besides with scattered, short, usually crisped hairs, on both
sides, especially along margin and veins, scabrous, lower surface also
with sunken and more or less numerous glandular hairs. Capitula solitary
terminal, 20-25 mm long (with florets slightly narrowed at base).
Involucre 15-20 mm wide in upper broader part, scaly, four- or five-
rowed, almost glabrous and weakly flocculose-arachnoid-hairy; outer
bracts oblong, middle and inner linear-lanceolate, acuminate, somewhat
small-toothed. Receptacle densely squamose; scales glabrous, unequal,
reaching one-third to half of involucral height. Corolla exserted from
involucre, purple, with scattered glandular hairs, 17-19 mm long, on 6—
8 mm long stalk and 4-5 mm long limb; basal appendages of anthers
setose, almost undivided, 3—3.5 mm long; filaments glabrous or with
scarcely visible tubercles above. Pappus barbate-plumose, with numerous
unequal bristles; mature achenes about 5 mm long and 2 mm wide,
four-angled, obpyramidal, glabrous, striate-streaked, with thickened edges,
spinose-toothed above along edges, edges with isolated spinules below,
with patellate thickening and pappophorus at tip, with pitcher-shaped
nectary in center.

Steppe mountain slopes.—Soviet Central Asia: Pamiro-Alat.
Endemic. Described from the Rabat area at the exit from the gorge of
the Karamuinak River. Type in Leningrad.

47. J. rhizomatoidea I\jin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XX (1960) 346.—Iec.: Ibid. p. 349, Fig. 2.

Perennial. Plants 8-15 cm high, with long-branched rhizomes,
covered with whitish, glabrous, lustrous sheaths of lower leaves and
with deep tap root. Stems simple, ascending, bright green, finely sulcate,
lustrous, with sessile glandular hairs and scattered pubescence, more
dense below capitulum. Leaves rosulate, like lower cauline leaves
narrowed toward base, slightly broadened at very base, amplexicaul;
cauline leaves sessile, narrowly decurrent, narrowly lanceolate, usually
5—8 mm long and 3—5 mm wide, acute, undivided and entire, one-veined,
usually revolute, glabrous beneath except midrib, and with sessile
glandular hairs, upper surface covered with short hairs, rather frequent
and mixed with short-stalked and sessile glandular hairs. Capitula solitary,
on short peduncles. Involucre wide, cupuliform, about 2 cm long,
glabrous, lustrous, arachnoid-hairy, only at base, glabrescent; involucral
bracts almost equal, innermost bracts narrowly lanceolate; all bracts
acuminate, with sessile, fine, scattered glandular hairs, green, with whitish
membranous border and small teeth here, usually three-veined. Corolla
18-20 mm long, on 6—7 mm long stalk, with about 5 mm long limb,
and very thin lobes, pinkish, with numerous sessile glandular hairs; apical '

594

588

appendags of anthers 2.5—-3 mm long, narrowed above; basal appendages
caudate, about 2 mm long, divided up to base, at least into two parts,
pubescent, stigma branches long. Pappus deciduous, 7-8 mm long, snow-
white, barbate, with barbules equal or much longer than thickness of
hair; achenes four-angled, obpyramidal, 3.5—4 mm long, brownish, corona
small-toothed at edge, longitudinally rugose along edges, with tubercles
modified into appressed spinules in upper half, apex with smooth conical
pappogenus, in center on stalk with membranous five-partite nectary. |
Flowering VII; fruiting VIII-IX. (Plate XL, Fig. 12.)

Stony and rocky habitats ——Soviet Central Asia: Tien Shan (Karatau).
Endemic. Described from the Utelshuya River Valley in the Karatau
Mountains. Type in Tashkent.

Note. The species is quite unique and stands by itself among all
other species of the section and may be split off into a separate
subsection.

48. J. trifurcata Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XXI (1961) 382, Fig. 1.

Perennial. Plants 8-12(15) cm high, with long, slender, branched
roots, forming perennating buds. Stems several, simple, grayish. Basal
leaves in compact rossetes, narrowly oblong to linear, about 2—3(4) cm
long, short-petiolate, like lower cauline leaves, usually three-partite at
apex, with entire margin, strongly convolute, almost up to midrib, on
both sides whitish-grayish due to dense indumentum, finely punctate;
upper cauline leaves remote, short, ‘sessile, always simple. Capitula
solitary. Involucre cupuliform, about 10—12 mm long, imbricate, four-
or five-rowed, sparsely arachnoid-hairy; involucral bracts oblong, with
prominent midrib; outer bracts appressed, yellowish-green, reddish only
above; middle bracts brighter, membranous along margin, cuspidate; inner
bracts entirely purple or only mottly-purple, with wide membranous
border, abruptly narrowed from gibbous tip into oblong-triangular, slightly
recurved, membranous cusp, with small teeth. Receptacle squamose with
short, smooth, acuminate scales. Florets relatively less numerous; corolla
pale pink, with sessile glandular hairs, about 12—13.5 mm long, narrowed
below over 5—5.5 mm length, and with 3.5-4 mm long lobes of limb,
tuberculate at tip; apical appendages of anthers 2—2.5 mm long, arcuate
at tip, basal appendages 2—2.5 mm long, caudate, curly; filaments
glabrous. Pappus persistent, 4-5 mm long, slightly shorter than mature
achene, barbate, with very dense barbules, almost as long as or usually
one and one-half times as long as thickness of bristles; achenes 5—5.5
mm long, oblong, obpyramidal, four-angled, glabrous, smooth, with
small-toothed border above. Fruiting VII.

Alpine zone, on rocks and screes.—Soviet Central Asia: Tien Shan

595

589

(western). Endemic. Described from the Chatkal Range, on the drainage-
divide between the Karagaily and Karasu rivers. Type in Tashkent.

Section 5. Suffrutices [ljin sect. n. in Addenda XXVI, 721.—
Semishrubs with undivided and entire, linear or filiform leaves; involcural
bracts appressed, sometimes only apices of cusps recurved; pappus serrate
or serrate-barbate, persistent (deciduous in J. sintenissii Bornm.); achenes
glabrous, smooth, often with longitudinal striations or glabrous folds;
pappogenus always present, pappophorus present only in J. sintenissii
Bornm.

Type of section: J. mugodsharica Iljin.

1. Semishrubs with long woody branches .............. cesses renee De
+ Semishrubs with strongly branched rhizome, forming many-headed
CAUGERMINMSOMN, , SEM) ay ee a nO RE, ON ee! 4.

“2. Capitula small; involucre not longer than 12 mm; pappus bristles
serrate; achenes glabrous, 1.75—2 mm long (Mugodzhary, Boktaubai
Mountam).122008.. SORT AES eee 51. J. mugodsharica [jin

+ Capitula larger, involucre 14—18 mm long; achenes 5—6.5 mm long,
with poorly developed longitudinal striations or streaks, smooth

3. Involucre cylindrical or oblong-narrow cyathiform; involucral enacts
abruptly narrowed into short cusp, finely arachnoid hairy-tomentose
on outer side, entire or inequidentate (Karatau)...........0.eceeeeeeee
ti EDN nL ae ney Re EC a oh rm 50. J. multiceps [jin

+ Involucre cupuliform; involucral bracts gradually acuminate, densely
covered on outer side with rather short, appressed bristles, finely
and densely uniformly ciliate (western Tien Shan and Karatau)..
PUR OE SOO ERE BE. Aer od a 49. J. suffruticosa Rgl.

4. Involucral bracts with prominent veins, narrowed into membranous,
long, recurved cusp; pappus serrate, 5—6 mm long, with inner
bristles up to 10 mm long; achenes 5 mm long (Fergana)...........
La A Le RE Hi Lot Eis Aa AL LE at lcd 53. J. orientalis [ljin

+ Involucral bracts with inconspicuous veins; cusps usually appressed,
and if recurved, then not membranous; pappus serrate-barbate, 7—
8 mm long, with inner bristles up to 20 mm long; achenes 5-6
mmeong* (Kopetdag) 2.552... ee 52. J. sintenisii Bornm.

Series 1. Ligneae Iljin——Stem basally and its lower branches
prostrate, ascending, sparse.

49. J. suffruticosa Rgl. in Bull. Soc. Nat. Mosc. XL, 2 (1867) 166;
Trautv. in Tr. Peterb. Bot. Sada, VIII, 1, 513; O. and B. Fedtsch. Perech.

596

590

Rast. Turk. IV, 299 p. p.—J. suffruticosa var. major Winkl. ex Ijin in
Tr. Turk. Nauchn. Obshch. 2 (1925) 16.—Exs.: H.F.A.M. No. 495.

Perennial. Semishrub, 30-50 cm high, with long, woody branches
below, 3—6 mm thick, producing numerous vegetative, densely leafy
shoot. Stems of current year elongate, simple, leafless above, arachnoid-
hairy. Leaves filiform-linear or linear, green and glabrous or finely
arachnoid-hairy above, grayish beneath due to fine tomentum, strongly
revolute, often almost covering lower surface, entire, sessile, strongly
reduced above and disappearing. Capitula solitary. Involucre cupuliform,
14-18 mm long and 10-15 mm wide, almost glabrous, young with
glabrescent arachnoid-hairy pubescence, green, purple above, somewhat
appressed at base; involucral bracts appressed, acuminate; outer bracts
oblong, others lanceolate, with scarcely visible, appressed bristles,
ciliolate along margin. Receptacle densely squamose; scales connate,
almost up to half as long as involucre. Corolla pinkish-purple, 14-17
mm long, narrowed over 5—7 mm length and with 3.5-4.5 mm long
lobes of limb. Pappus persistent, snow-white, 7-10 mm long; achenes
(5)6—6.5 mm long, yellowish-brown, glabrous, almost smooth, without
teeth along upper edge and ribs, toothed above along edge, with traces
of parallel ridges, apex with two-tiered pappogenus; upper petallate,
with nectary on small elevation in center, lower almost hemispherical.
Flowering V—VI; fruiting V—VII. (Plate XLVI, Fig. 3.)

Gravelly and tuffaceous mountain slopes, rocks, among juniper
forests—Soviet Central Asia: Tien Shan (western). Endemic. Described
from Karatau. Type in Leningrad.

50. J. multiceps Ijin in Bot. Mat. Gerb. Glavn. Bot. Sada, V, 4—5
(1924) 4; Tr. Turk. Nauchn. Obshch. 2, 16.

Perennial. Semishrub, 15—25(30) cm high, forming loose turf with
strong, usually vertical tap root, branched above, fibrous, producing
numerous short woody stems at soil surface, covered with remains of
old leaves. Stems numerous, simple or often with isolated short branches.
Leaves filiform, entire, strongly revolute usually up to midrib, sessile,
dense in lower half of stem, shorter and much remote above, leafless
below capitula, arranged in rosettes at the end of caudices, flocculose-
arachnoid-hairy, glandular-punctate. Capitula numerous or solitary.
Involucre oblong-cylindrical, usually 14-16 mm long and about 6-8
mm wide, often slightly narrowed at base, with appressed bracts; outer
bracts shorter, others oblong-oval, with somewhat prominent veins,
abruptly narrowed into relatively short cusp, finely arachnoid hairy-
tomentose on outer side, entire or with fine isolated, and irregular teeth.
Corolla pink-purple, 13-14 mm long, narrowed over 5—6 mm, with about
4 mm long lobes of limb. Pappus persistent, colorless, short-barbate,

597

591

with dense barbules, usually shorter than thickness of bristles; achenes
obpyramidal, obtusely four-angled, slightly broader below corona, mature
achenes 6—6.5 mm long, strongly narrowed at base, with toothed corona
at apex, edges and ribs without teeth, tuberculate, white-cartilaginous
below ribs, with parallel longitudinal ridges, with small patelliform
pappogenus at apex having nectary in center. Flowering VI—VIII; fruiting
VIII. (Plate XLVI, Fig. 1.)

Rocks, gravelly and loamy-gravelly mountain slopes—Soviet Central
Asia: Tien Shan (Karatau Range). Endemic. Described from Karaguz.
Type in Leningrad.

51. J. mugodsharica Iljin in Mat. Komiss. Eksp. Iss]. Akad. Nauk.
SSSR, 26 (1930) 277, Fig. 1; Pavlov, Fl. Tsentr. Kazakhst. II], 304.—
Ic.: Iljin, op. cit. Fig. 1.

Perennial. Semishrubs, 10—20 cm high, with prostrate woody stems,
producing short branches, covered with remains of dead leaves and
reducing rosette of leaves, from center of which arise simple, erect,
leafless, arachnoid-hairy branches in upper part. Leaves linear-filiform,
entire, strongly convolute, slightly. broadened at base; cauline leaves
sessile. Capitula solitary, with almost glabrous, 10-12 mm long and 5-—
8 mm wide involucre; outer involucral bracts oblong-triangular; middle
bracts lanceolate or oblong; inner bracts linear-lanceolate; all bracts
pointed. Pappus 2.5—3.5 mm long, serrate, with dense teeth shorter than
thickness of bristles; achenes 1.75—2 mm long, smooth, compressed,
four-angled.

Gravelly soils.—Soviet Central Asia: Aralo-Caspian Region.
Described from Mugodzhar-Botubai Mountain and vicinity of town of
Mugodzharskii. Endemic. Type in Leningrad.

Series 2. Grumosae Iljin—Rhizomes clustered; stems with dense
rosette of basal leaves.

52. J. sintenisii Bornm. in Beih. Bot. Centralbl. XX, 2 (1906) 165;
O. and B. Fedtsch. Perech. Rast. Turk. IV, 299; Iljin in Tr. Turk. Nauchn.
Obshch. 2, 16.—J. suffruticosa var. latifolia Winkl. in Tr. Peterb. Bot.
Sada, XI, 1 (1890) 145; O. and B. Fedtsch. op. cit. 299; spec.
kopetdag—J. arachnoidea f. aprica Bornm. 1. c—Ie.: Iljin, op. cit.
Plate 37; Fl. Turkm. VII, 231.—Exs.: Ed. H.B.P.M. No. 99.

Perennial. Semishrub, 20(15)—40(50) cm high, with fibrous root
branched above, and strongly branched rhizomes densely covered with
dark brown scales—remains of old leaves. Stems numerous, simple,
relatively slender, finely-sulcate, usually with scattered, sessile, fine
glandular hairs, grayish due to sparse arachnoid-hairy pubescence, leafy

598

ae

predominantly at base and up to middle, less often up to two-thirds in
dwarf forms, almost leafless above. Leaves narrow-oblong or oblong-
linear, tender, 3-7 mm long and 1—3 mm wide, gradually narrowed on
lower side and rather broadened at base, short-cuspidate, especially in
broad-leaved forms, reaching 4.5—10 cm in length and 5—6 mm width;
all leaves entire, convolute, less often almost flat or sparsely pinnatipartite
and pinnatilobate or only obtusely toothed, usually with fine sessile
glandular hairs on both sides, grayish-green above with depressed midrib,
more or less prominent on lower sides, whitish beneath due to tomentum;
cauline leaves gradually reduced above, solitary, usually indistinct.
Capitula solitary, usually narrow-cupuliform. Involucre imbricate, 10-
15 mm long, and 7-10 mm wide (at anthesis); outer involucral bracts
very short, lanceolate, oblong-ovate or oblong-lanceolate, grayish-green
due to arachnoid-hairy pubescence, with narrow dark green or brown-
greenish, appressed or recurved apices, carinate, with scattered sessile
glandular hairs; middle and especially inner bracts oblong, usually dirty
purple, but with many sessile glandular hairs, terminating into long,
appressed, brownish cusps, very finely and frequently toothed. Receptacle
squamose; scales smooth, glabrous, 3-6 mm long, often split above.
Corolla pink, with scattered, sessile glandular hairs, 14-16 mm, usually
15 mm long, narrowed over 5—6(6.5) mm length, with 4-5 mm long
lobes of limb; apical appendages of anthers 2—2.5 mm long, whitish,
oblong-acute, basal appendages setaceous, colorless, weakly divided.
Pappus deciduous, 7-8 mm long, with two hairs up to 20 mm long,
snow-white, with many bristles, densely barbate-serrate; bristles connate
into ring with common base, developing pappogenus at apex with
tumbler-shaped membranous nectary. Flowering [V—VI(VIJD); fruiting VI—
IX.

Steppe and juniper zone, in ravines, rocks, stony slopes.—Soviet
Central Asia: mountainous Turkmenia. Endemic. Described from
Sundosag. Cotype in Leningrad.

53. J. orientalis [ljin in Tr. Turk. Nauchn. Obshch. 2 (1925) 17
(pro subsp. orientalis)—J. suffruticosa Fedtsch. non Rgl.; O. and B.
Fedtsch. Perech. Rast. Turk. IV (1911) 299 (specimina ferganica).

Perennial. Plants 20-30 cm high, grayish; tap root straight, fibrous,
terminating into many-headed caudex with rosettes of usually. numerous
basal leaves. Stems few, simple, leafy only in lower one-third part, 1—
2 mm thick, finely sulcate-ribbed, with scattered sessile glandular hairs,
finely sparsely arachnoid-hairy. Basal leaves usually 5-8 mm long,
cauline leaves sessile; all leaves linear, entire, subobtuse or usually short-
cuspidate, punctate, with depressed midrib and greenish-grayish above
due to sparse arachnoid-hairy pubescence, whitish beneath due to more

2/3)

SSS

LZ

es Lg

Plate XLIV.
Jurinea robusta Schrenk.

538)

1

600

594

dense arachnoid tomentum, strongly convolute, upward spreading,
reducing up to one-third part of stem, absent above. Capitula solitary,
with many florets. Involucre cupuliform, 12-16 mm long, usually
five-rowed, imbricate; involucral bracts oblong-oval, outer shorter,
innermost narrowly lanceolate; all bracts appressed, terminating in long
membranous, recurved cusp, with prominent veins, especially midrib,
covered with short, crisped, arachnoid hairs and scarcely visible, more
or less densely appressed, golden bristles, ciliate along margin. Receptacle
squamose; scales smooth, pointed one-fourth to one-third as long as
involucre. Corolla pinkish-purple, 13—14 mm long, with scattered sessile
glandular hairs, narrowed over 5 mm length, with 4-5 mm long lobes
of limb; apical appendages of anthers 2.5—3 mm long, acuminate, basal
appendages setaceous, about 2—2.5 mm long, divided below; filaments
glabrous. Pappus snow-white, pappus bristles almost as long as achenes,
and four longer flat hairs two times as long, bristles serrate; achenes
about 5 mm long, four-angled, obpyramidal, light-colored, yellowish,
glabrous, smooth, weakly fine-striate longitudinally, with weak
longitudinal folds, and toothed corona, with scarcely visible lustrous
dots or glandular hairs above below corona, and dense thick annular
pappophorus bearing conical nectary. Flowering V; fruiting VI.

Stony wormwood-saltwort desert.—Central Asia: Syr-Darya.
Endemic. Described from the vicinity of the village of Kara-Kurgan in
Fergana Valley. Type in Leningrad.

Section 6. Floccosae (Sosn.) Iljin comb. nov.—Sect. II.
Platycephalae Benth. and Hook. Subsect. Floccosae Sosn. in Zhurn.
Russk. Bot. Obshch. XI, 1—2 (1926) 195.—§4. Pinnatilobatae DC. Prodr.
VI (1937) 697 p. p.—§1. Involucro cylindrico Ldb. Fl. Ross. II, 2 (1845—
1846) 761 p. p.—§3. Pinnatae Boiss. Fl. or. III (1875) 572 p. p.—
Stems virgate, branched; involucre cylindrical; corolla 25-35 mm long;
pappus_ entirely deciduous, pappus bristles connate into thick ring,
nectary exserted from ring; pappogenus developing inside at border of
ring; pappophorus conical or smoothly dome-shaped.

1. Leaves glabrous above, whitish only along midrib due to fine
arachnoid-hairy tomentum, somewhat arachnoid hairy beneath; basal
and lower cauline leaves sinuate-toothed, or margin entire; involucre
glabrous, outer bracts uncate along half length; pappus bristles
barbate, barbules as long as or shorter than thickness of bristles
seer cue t ak Mente aes Eisai: 54. J. armeniaca Sosn.

+ Leaves glabrous on both sides, slightly arachnoid-hairy above only
along midrib; basal and lower cauline leaves pinnatipartite or
pinnatisect; involucre arachnoid hairy-flocculose, outer bracts with

601

32)

herbaceous appendage, uncate; pappus bristles long-barbate,
barbules two to five times as long as thickness of bristles ........ Ms
Sey eae en, Sale as to nsonit cs seemuedemeae ease cs 55. J. elegans (Stev.) DC.

54. J. armeniaca Sosn. in Vestn. Tifl. Bot. Sada, No. 49 (1920)
31; Sosn. in Zhurn. Russk. Bot. Obshch. 11, 2 (1926) 195; Grossh. FI.
Kavk. IV, 172.

Perennial. Plants 15-30 cm high, with stiff caudex. Stems numerous,
flocculose-tomentose at base due to dense woolly buds in axils of
approximate petioles of dead leaves, angular-sulcate, erect, usually
densely leafy, less often simple, usually branched in lower half, with
long upward spreading branches. Basal and lower cauline leaves oblong
or lanceolate-linear, sinuate and sparsely toothed, less often almost
undivided, gradually narrowed into flat petiole, broadened at base, leaves
withering much before anthesis; upper leaves oblong-lanceolate or linear,
undivided and entire, narrowed at base; all leaves acute or cuspidate,
with callose thickening at apex, convolute, somewhat hard, arachnoid-
hairy in axils, green on both sides, with dense sessile glandular hairs,
weakly rugose and glabrous above, slightly arachnoid-hairy only along
midrib, slightly arachnoid-hairy beneath. Capitula 15-25 mm long,
solitary, with many florets. Involucre broadly turbinate or cupuliform,
glabrous, usually six-rowed; outer bracts one-veined, from oblong or
oblong-lanceolate base narrowed into uncate appendage at apex, almost
half as long as middle bracts, arachnoid-hairy on inner side; inner bracts
oblong-linear, membranous and gradually acuminate, purple, usually
pubescent on outer side, all bracts ciliolate along border. Receptacle
squamose; scales thin, long, acuminate, one-fourth to one-third as long
as involucre. Corolla 25—30(32) mm long, pale pink, with dense, sessile
glandular hairs, gradually narrowed over 6.5-10 mm length, with 6—7
mm long lobes of limb; apical appendages of anthers 6-7 mm long,
acuminate, basal appendages 3-4 mm long, setaceous, divided below;
stigma 3—4 mm long, bifid up to base, basally with separate, squarrose,
short bristles, gradually terminating into papillae; style glabrous; filaments
glabrous, ferruginous-brownish. Pappus 18-20 mm long, with less
numerous bristles connate into ring and shedding together with it, bristles
barbate-serrate, with barbules shorter or almost as long as thickness of
bristles; achenes immature, 6-7 mm long, oblong; four-angled, but not
pyramidal, slightly narrowed toward base, glabrous, smooth, whitish,
with corona of long teeth, modified along border as parallel striations;
pappophorus as white membranous, roundish base, with five-angled
reddish star and a nectary inside. Flowering V—VII; fruiting VII-VIII(?).
(Plate XL, Figs. 3-5.)

Rocks, stony slopes, especially with phryganoid vegetation; on stony
river banks.—Caucasus: Southern Transcaucasia. Endemic; possibly may

602

596

be found in Iran. Described on the basis of specimens from Migri in the
Nakhichevan Republic from the ravine of the Araksa River between
town of Ordubad and Karchevan post. Type in Tbilisi.

55. J. elegans (Stev.) DC. Prodr. VI (1837) 675; Boiss. Fl. or. III,
572; Ldb. Fl. Ross. II, 762; Grossh. Fl. Kavk. IV, 171; Sosn. in Zhurn.
Russk. Bot. Obshch. 11, 2, 195.—Serratula elegans Stev. in Mem. Soc.

Nat. Mosc. IV (1813) 101; Stev. in Trans. Linn. Soc. XI, 414; MB. FI.

taur.-cauc. III, 547.—Ic.: Trans. XI, tab. 37; Grossh. and Prilinko,
Geobot. Ocherk Kavk. Stepei (1929) Plate 15—Exs.: Pl. or. exs. No.
249.

Perennial. Plants 15-60 cm high, with stiff caudex at stem base,
covered with pubescence of dense woolly flufly mass. Stems several,
erect, branched in lower half; branches horizontal and also erect or
slightly arcuate, cylindrical, glabrous, not strongly sulcate, densely
covered with sessile glandular hairs, arachnoid-hairy only below
capitulum. Basal leaves pinnatisect, less often pinnatipartite, “oblong,
gradually narrowed into flat petiole broadened at base, with more or
less long and narrow, linear, acuminate segments; lower cauline leaves
short-petiolate or sessile, either mostly pinnatipartite or sinuate-toothed,
also pinnatisect; other leaves on branches more or less undivided, linear-
oblong or linear, with entire margin or weakly toothed, somewhat
strongly convolute, acute or acuminate, almost reaching capitulum; all
leaves hard, green and glabrous on both sides, whitish above and along
midrib due to fine arachnoid-hairy pubescence, rugose and with sessile
glandular hairs, more strongly glandular beneath. Capitula solitary, with
many florets. Involucre poculiform, grayish due to arachnoid-hairy
pubescence, usually seven-rowed, appressed, all bracts with single
prominent vein; outer bracts narrow-membranous and finely serrate,
besides innermost, with herbaceous apex, short uncate; innermost bracts
gradually long-acuminate, covered with appressed short bristles on outer
side, somewhat purple. Receptacle squamose; scales smooth, acuminate,
much shorter than involucre. Corolla (25)27—30(31) mm long, gradually
narrowed over 10-12 mm length, with 4-6 mm long lobes of limb,
covered with sessile glandular hairs; apical appendages of anthers 4—5
mm long, gradually narrowed above, basal appendages setaceous, 3-4
mm long, divided, smooth; stigma about 2 mm long, bifid up to base,
with short tuft of bristles at base, and fine papillae on outer side; style
glabrous; filaments brownish, glabrous, smooth. Pappus deciduous, about
17-20 mm long, pappus bristles connate into ring and easily shedding
at fruiting, with distinct sparse barbules, two to two and one-half times
as long as thickness of bristles, and barbules on two to five prominent
thicker bristles shorter, equal or one and one-half times as long as their

603

597

thickness, more dense and usually allover, thick ring forming inside
pappus at tip, through which columnar pappophorus penetrates; achene
four-angled, oblong, slightly narrowed toward base, 7—7.5 mm long,
with regularly toothed corona above, columnar pappophorus bearing a
nectary exserted from inside and middle glabrous, smooth, with indistinct ©
ridges along edges, hilum usually slightly lateral. Flowering VI—VIII;
fruiting VIII-IX.

Steppe slopes, gravelly and sandy deposits, limestone deposits.—
Caucasus: Eastern Transcaucasia, southern Transcaucasia (rarely).
Endemic. Described from the vicinity of the village of Dzhanak Bulak,
near the Gokchai River. Type in Leningrad.

Section 7. Penduliflorae Iljin sect. n. in Addenda XXVI, 721.—
Plants with strongly pendulous ray florets; corolla gradually narrowed
below, pink or purple, with scattered sessile glandular hairs. Pappus
persistent, except in J. comarovii Iljin, J. elegantissima Ijin (entirely
shedding) and J. kamelinii Iljin (partially shedding in central part).
Achenes large, glabrous, with more or less prominent or poorly visible,
longitudinal, parallel, smooth, ridges, mostly modified into sharp
tubercles, below corona. Pappophorus usually not developed (present
only in species with deciduous pappus: J. komarovii, J. elegantissima
and J. kamelinii), dome-shaped pappogenus present in all species as
thick, beet-red, broad cup, whitish on inner side, nectary relatively large
and on more or less short manubrium. A total of about 15 species in
this section, so far known only from the USSR.

Type of section: J. robusta Schrenk.

—

. Plants stemless or with short up to 10 cm long stem............... as
+ Plants witht wellodéveloped "Stemi: s0..22 22... 4¢c2.f0tesceecaeenee--- vbr ae eee Si
2. Stem short; leaves almost all basal, or spatulate, white-tomentose
beneath, green above, with depressed veins; involucral bracts
appressed; pappus bristles barbate (southern Tadzhikistan) ..........
sh ih cal Dl Ae Mee oe BD 71. J. impressinervis I]jin
+ Plant stemless or with weakly developed stem; leaves concolorous;
involucral bracts uncate (mountains of southern Tadzhikistan) ....
PNR YS «28. AG cesccanacecancevse SUS: ott ee ee etn totes 72. J. apoda [lyin

3.2 Allsleaves*undividedVandMentire 2229 122) Re 4.
+ All leaves usually pinnatilobate, pinnatipartite or pinnatisect, less
often..only ssmuate-toothed AH AS7SOnO0ne.2 Peleg SRE... Pte
4. Pappus deciduous entirely or partially; pappophorus and pappogenus
alWays+ present eeee AGE AVRO) OARS TE As 5
+ Pappus persistent; only pappogenus present ................ cece 6.

5. Robust plants with large, somewhat hard, decurrent leaves, reaching
capitula; involucral bracts, except innermost terminating into long,

598

++

12:

smoothly incurved, spinose tip. Pappus plumose; achenes up to 8—
9 mm long, linear, glabrous (Khan-Tau Mountains) (subsection
ROGUSIAE et or oie stoner aee cease erarce a eee 56. J. robusta Schrenk
Leaves herbaceous, decurrent, but absent in upper part of stem;
involucral bracts appressed, not terminating into strong spine.....

Leaves nondecurrent, linear, entire...... 59. J. elegantissima nie

. Plants entirely covered with glandular and flexuous granose hairs,

often reduced and terminating into papillae (Zeravshan, western
Pamir) (subsection Papilligerae) oo oe: tenn skp es pac dee eee vaccearees fie
Stem and leaves with only sessile glandular hairs and with or
without arachnoid-hairy pubescence, or always without granose
flexuous hairs. Pappogenus developed, Pappophorus absent
(subsection! Pensisterttes):. tierce. Ayu. aan. eAtee. op Ae 8.

. Plants densely covered with granose, vacuolated flexuous hairs;

pappus partially deciduous (western Pamir, upper Zeravshan) .....
fa A TLOLIED. ATS RPSL NG, OPEC EAL i 58. J. kamelinii [ljin
Plants densely covered with granose pappillae and scattered, granose
flexuous hairs, or without them. Pappus entirely deciduous (western
Bannrit)y o.ciove A ERD ote ce, EOE 57. J. komarovii []jin

. Leaves long, narrow, linear, narrowly and shortly decurrent,

convolute almost up to midrib, arachnoid-hairy beneath, like stems.
Capitula numerous, on long, leafless bent peduncles (Karatau, Syr-
Datya) sc. Ceea. geet Ee OD. BIG ad 63. J. cephalopoda [lyin
Leaves oblong-oval, usually laterally irregularly decurrent, not
convolute, narrowed towards both ends. Capitula on long, straight
érectopatentapedunclests.:cc sa sek... pee ea Leen ome era o

. Leaves distinctly gray-arachnoid hairy beneath (Uzbekistan, and

Madzhikistan))). 605.2029 60. J. trautvetteriana Rgl. and Schm.
Both sides of leaves green, not arachnoid-hairy ................... 10.

. Barbules of pappus bristles, three to four times as long as thickness

Of “Brastles:!05¢! JU i) as: Ss eect de Ue ee Rtarys chee M ad 62. J. popovii [lyin
Barbules of pappus bristles as long as, slightly shorter or longer
than thickness of bristles (Baisun)................. 61. J. gracilis Ijin

. Stem slender; all leaves petiolate, even uppermost leaves not

decurrent and not curly; capitula more or less narrow, on long
bent peduncles (Khodzha-Kazian mountains) ...............::ccccsee
| cue duak -Eyabih «naceptenbeatateate es. + SERS a tees. eee eats 64. J. bucharica Winkl.
Stem thicker; cauline leaves always sessile ...............:::c:eee v2
Leaves. distinctly) decurrent:s scicsie- re. ceqaes sc ees esbeet peepee HSE
Leaves not decurrent, sometimes only semiamplexicaul or with
leaf bases coalesced with corresponding branch .....................- les

605

512)5)

13. Plants grayish arachnoid-hairy; stem branched, mainly in lower
half; cauline leaves decurrent, distinctly sinuate; capitula large,
with 20-30 mm long involucre, also arachnoid, on straight upward
‘spreading branches (southern Kyzyl-Kum, northern: Kara-Kum) ..
Er gs ch 4O cases ft adi eb ta 65. J. psammophila [ljin

+ Stem branched up to the tip; branches more or less arcuately
incurved; involucre 15—20 mm [Ong .......... eee cee ceseeeeteeeeees 14.

14. Involucre not arachnoid-hairy; glandular and with short appressed
bristles; stem dark green, almost glabrous; leaves hard, linear,
convolute, finely sinuate-toothed (northwestern foothills of Karatau)
th erctas ta bale RA a a alts valet Reranch 67. J. karatavica [jin

+ Involucre, like entire plant, arachnoid-hairy; leaves oblong-
lanceolate, boldly sinuate-toothed, pinnatilobate or less often
pinnatipartite (Kopetdag and northwestern foothills of Pamiro-Alai)
ie Mh Lah aca hae ee one ahi ee 66. J. lasiopoda Trautv.

15. Leaves always sinuate; cauline leaves sessile, semiamplexicaul,
pinnatipartite or pinnatisect. Capitula large, on long, erect peduncles
argo lela panera La i pent l | ahece Mn 68. J. botschantzevii [jin

+ Leaves not sinuate, undivided, oval, oblong-oval or lyrate-pinnate,
with larger terminal lobe, toothed or lobate............. 16.

16. Entire plant with glabrescent, flocculose-arachnoid-hairy pubescence,
with less deeply depressed midrib, almost not rugose above, densely
covered with sessile glandular hairs, not sunken in pits; pappus
bristles somewhat thick, with more or less dense, long, three to
four times as long as thickness of bristles, arcuate and obliquely
amanged Datoules: (Pereana Valley)... ccstss..gncceresscosdacaapen tates scansegtnc?
ah anne Ua nat trae nepeadruat «Noe iliidead ts pall ea ee 69. J. winkleri [Ijin

+ Entire plant, especially stem, uniformly pubescent with crisped
arachnoid-hairy pubescence; leaves with distinctly depressed midrib,
strongly finely rugose, with scattered sessile glandular hairs, in
deep alveoles; pappus bristles sparse and short-barbate; barbules
as long as thickness of bristles, shorter, or more than one and one-
half to two times as long, semiappressed to axis (Buruyutau
Mountains in Vakhsh Valley) ........... 70. J. tadshikistanica [ljin

Subsection 1. Robustae I[ljin——Leaves reaching up to capitula;
involucral bracts terminating into long spines.

56. J. robusta Schrenk in Bull. Acad. Sc. Pétersb. III (1844) 109;
Trautv. in Tr. Petersb. Bot. Sada, VIII, 2, 515; O. and B. Fedtsch. Perech.
Rast. Turk. IV, 303; [jin in Tr. Turk. Nauchn. Obshch. 2, 18.

Perennial. Plants 20-40 cm high, glabrous. Rhizome strong,
terminating into turf of short, dense, woolly-tomentose caudices. Stems

606

600

very strong, thick, 5—8 mm thick at base, ribbed-sulcate, usually several,
simple or weakly branched, like branches densely leafy up to capitula.
Leaves somewhat hard, oblong or even almost oval, entire, often
somewhat sinuate, acute or cuspidate, densely covered with sessile
glandular hairs, with prominent veins on lower side; basal leaves
gradually narrowed into short and strong basally broadened petiole;
cauline leaves (except lower and uppermost) strongly decurrent,
uppermost reduced and strongly narrowed; all leaves green on both sides.
Capitula solitary, large. Involucre 30-40 mm long and 25-35 mm wide,
together with squarrose appendages somewhat turbinate at base, yellow-
green, sometimes arachnoid-hairy-flocculose at base; involucral bracts
numerous, gradually narrowed into 8-15 mm long, somewhat strong,
initially divergent cusps, later almost arcuately incurved, covered with
rather short bristles, innermost bracts erect. Receptacle covered with
yellowish hairs, several times shorter than involucre. Corolla pink, 28—
45 mm long, gradually narrowed over 10-20 mm, with 5-10 mm long
lobes of limb, covered with sessile glandular hairs; apical appendages
of anthers acuminate,5~7 mm long, basal appendages setaceous, about
5 mm long, slightly divided; filaments papillose; style glabrous, stigma
branches bifid almost up to base, densely papillose, flat, 4-5 mm long,
with short bristle below branches. Pappus deciduous, snow-white,. 20-
25 mm long, including two long bristles, many-rowed with unequal
bristles, plumose, easily detached from achene on maturation, with ring
at base and drawn out membranous base, thickened into patelliform
pappogenus, bearing pentagonal membranous nectary in center; achenes
linear, 8—9 mm long and about 1.5 mm thick, smooth, yellowish-brown,
with toothed border above, and oblong-conical whitish pappophorus
inside. Flowering VI—VIII; fruiting VII-VIII. (Plate XLIV.)

Varicolored clayey, stony slopes and rocks.—Soviet Central Asia:
Tien Shan. Endemic. Described from Khantau (Chu-Ili Mountains). Type
in Leningrad. |

Note. An extremely distinctive species, combining characters of this
section as well as of section Derderiae Boiss. We have placed it in a
separate subsection. The Khantau Mountains, where it grows, are
generally distinguished by sharp endemism. Besides this jurinea, the
most striking example to use would be Niedzwetzkia semireczenskia, a
species from the tropical family Pedaliaceae.

Subsection 2. Papilligerae Iljin—Pappophorus and pappogenus
developed.

57. J. komarovii I[ljin in Bot. Mat. Gerb. Glavn. Bot. Sada, V, 4
(1924) 59; Ijin, Tr. Turk. Nauchn. Obshch. 2, 19.

607

601

Perennial. Plants 25-80 cm high, many-stemmed, light green. Root
thick, strongly branched, woody. Stems densely woolly-arachnoid-hairy
near collar, finely sulcate, subglabrous with isolated scattered, rather
short hairs, slightly arachnoid-hairy below capitula, simple or usually
with few virgate branches, strongly leafy from base to middle. Leaves
oblong undivided and entire, acuminate or acute, mainly with appressed
glandular hairs above, especially in upper leaves, on both sides green
and almost glabrous or with scattered hairs, short along veins; basal
leaves less numerous, gradually narrowed into petiole; cauline leaves
sessile, strongly long-decurrent, upper ones small, linear. Capitula
solitary, 3-4 cm long. Involucre relatively broad, cupuliform, more or
less narrowed toward base, five- to seven-rowed; involucral bracts
linearly-lanceolate, reduced above, on outer side weakly and sparsely
arachnoid-hairy, on inner side glabrous. Receptacle covered with smooth,
unequal hairs, almost half as long as involucre. Corolla pink, up to 27—
30 mm long, narrowed over half, glabrous, almost two times as long
as involucre, slightly projecting, with about 5-6 mm long limb;
appendages of anthers setaceous, 3—3.5 mm long or weakly divided;
filaments glabrous. Pappus easily shedding, with many bristles,
with slightly yellowish tinge, bristles barbate-plumose, unequal, with
barbules almost three times as long as thickness of bristles, all bristles
connate into ring; achenes four-angled, glabrous with indistinct parallel
ridges, and isolated short tubercles or teeth above, below toothed corona,
pappophorus almost oblong, dome-shaped, whitish in middle of corona
with five-toothed membranous nectary in center. Flowering VII—VIII;
fruiting VIII-IX.

Rocks, stony slopes in alpine belt—Soviet Central Asia: Pamiro-
Alai (eastern part). Endemic. Described from Shugnan, from the valley
of the Garmchashma River. Type in Leningrad.

58. J. kamelinii [jin sp. n. in Addenda XXVI, 721.

Perennial. Plants 25-40 cm high. Stem solitary or several, sulcate-
angular, with whitish ribs and green furrows, densely covered with sessile
glandular hairs and long, vacuolate, granose, many-celled flexuous hairs.
Like leaves, stems also weakly arachnoid-hairy only in upper part,
especially below capitulum, usually with long branches in lower half.
Leaves usually crowded in lower half of stem; basal leaves gradually
narrowed into petiole, narrow-lanceolate like cauline leaves, only
uppermost leaves linear and strongly reduced; all leaves undivided and
entire, acuminate, green on both sides, densely covered with hairs as on
stem, especially younger, besides with numerous sessile glandular hairs,
midrib depressed above, prominent and whitish beneath; cauline leaves
always sessile, often asymmetrically decurrent, uppermost leaves

608

602

semiamplexicaul, with small auricles, all spreading. Capitula solitary,
with many florets, 25-35 mm wide at anthesis. Involucre 15-20 mm
long, all involucral bracts appressed, oblong to narrowly lanceolate; outer,
and often also middle bracts greenish, cuspidate or short-acuminate,
broadly white-membranous along margin, one- to three-veined, arachnoid-
hairy on outer side, and also with scattered glandular hairs; inner bracts
acuminate, with purple tinge, densely glandular, but not arachnoid-hairy,
like all other bracts ciliolate along border. Receptacle densely squamose,
scales smooth, acuminate, one-third to half as long as involucre. Corolla
24-27 mm long, pink or purple, with scattered sessile glandular hairs,
gradually narrowed over 10-13 mm length, with 5—6 mm long lobes of
limb. Pappus snow-white, partially deciduous, lateral’ bristles persistent,
bristles in middle with barbules one and one-half to three times as long
as thickness of bristles, bristles weakly connate into ring (except
outermost), with membranous base, bearing thick, brown, stalked
pappogenus in center, terminating with membranous, five-fid, flexuously
frilled nectary; mature achenes about 7 mm long, oblong, whitish, smoky
above, glabrous, with indistinct, longitudinal, smooth striations,
terminating into spinules only below corona; corona unequally toothed,
slightly recurved, pappophorus dome-shaped on small elevation inside
corona, with remains of manubrium above. Flowering VI; fruiting VI—
VHI(?).

Stony slopes.—Soviet Central Asia: Pamiro-Alai Region. Endemic.
Described from Zeravshan. Type in Leningrad.

59. J. elegantissima Iljin in Addenda XXVI, 722.

Perennial. Plants 50-60 cm high, branched right from base, entirely
green, weakly arachnoid-hairy pubescent and with numerous sessile
glandular hairs; branches long, arcuately spreading above, slender. Leaves
linear, undivided and entire, one-veined, gradually narrowed toward base,
sessile, not decurrent, concolorous, usually more or less horizontal, woolly
fluff in axils of lower leaves. Capitula on long peduncles, solitary.
Involucre almost poculiform, 20-25 mm long, involucral bracts appressed,
acuminate, green, almost glabrous, with prominent veins; outer bracts
strongly reduced, innermost gradually acuminate into awn. Corolla
pinkish, with scattered sessile glandular hairs, up to 25-27 mm long,
narrowed over 6—10 mm, with 6—7 mm long lobes of limb, outer corolla
lobes strongly pendulous. Pappus deciduous, with plumose bristles
connate into ring, and with pappogenus and nectary on its membranous
base; achenes oblong, 9-10 mm long, with smooth longitudinal ribs,

[sic]; recte outer—Sci. Editor.

603

with band of teeth only below corona, and with dome-shaped
pappophorus inside corona. Flowering VI-VII; fruiting VII-VIII.

On gypsum rocks.—Soviet Central Asia: Pamiro-Alai Region.
Endemic. Described from the upper reaches of the Yakkabak-Darya River
from the vicinity of the village of Tashkurgen. Type in Tashkent.

Subsection 3. Persistentes Iljin—Only pappogen present.
Series 1. Graciles Iljin—All leaves undivided and entire.

60. J. trautvetteriana Rgl. and Schmalh. in A.P. Fedtsch. Putesh.
v Turk. III, No. 18 (1882) 52; O..and B. Fedtsch. Perech. Rast. Turk.
IV, 320 (specimina serawschanica); Iljin in Tr. Turk. Nauchn. Obshch.
2, 191.—J. lasiopoda f. integrifolia I\jin in herb.—Exs.: H.F.A.M. No.
487.

Perennial. Plants 20-60 cm high. Root woody, usually branched,
often many-headed above. Stems solitary or few, erect, simple or with
few branches, sulcate, rather short and somewhat sparsely hairy, at some
places flocculose-arachnoid-hairy, especially strongly below capitulum,
woolly-tomentose near base, leafy up to middle. Leaves morphologically
resembling those of willow, oblong-lanceolate or lanceolate, undivided
and entire, acute or acuminate, green, very finely rugose, almost glabrous
and rather scabrous above, grayish arachnoid-hairy beneath, especially
young leaves, and with weaker arachnoid-hairy pubescence scattered
glandular hairs visible; basal leaves narrowed into somewhat winged
petiole; cauline leaves distinctly decurrent, sessile, more or less narrowed
below, uppermost leaves reduced to linear. Capitula solitary. Involucre
narrow cupuliform, 15-25 mm long and 7-12 mm wide, somewhat
narrowed toward base, arachnoid-hairy; outer bracts ovate-oblong to
linear, strongly reduced, sometimes appressed, green in upper part; middle
bracts strongly arachnoid-hairy, narrowly lanceolate, purple along margin;
inner bracts exserted, finely acuminate and membranous, purple, lax,
without serration along margin, like other bracts. Receptacle densely
covered with unequal smooth bristles, reaching up to half of involucral
height. Corolla pink, pendulous, 23-30 mm long, narrowed over 8-12
mm, with 5.5—7 mm long limb, and scattered glandular hairs, transversely
rugose in middle; basal appendages of anthers setaceous. Pappus
persistent, serrate, with dense barbules shorter than thickness of bristles,
pappus bristles rather numerous, snow-white; achenes 5—6 mm long,
yellow, indistinctly four-angled, glabrous, smooth, spiny-toothed above
along edges with cupuliform pappogenus at apex, with nectary in center.
Flowering V—VI; fruiting VI-VII.

Gravelly slopes of foothills and varicolored rocks.—Soviet Central .

610

604

Asia: Pamiro-Alai (western part).-Endemic. Described from the Aksai
Mountains in the Zeravshan Valley. Type in Leningrad.

61. J. gracilis Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada, V, 4
(1924) 60; Iljin in Tr. Turk. Nauchn. Obshch. 2, 19, Plate 1.

Perennial. Plants 25-60 cm high, many-stemmed, light green. Roots
relatively thick, usually with short many-headed caudex. Stem virgate,
branched from base to middle or up to two-thirds length, and leafy in
this part, striate-sulcate, glabrous or subglabrous, woolly-tomentose at
base. Leaves oblong to lanceolate, undivided and entire, or with sparse
teeth in lower half, acute, glabrous on both sides, green with scattered
appressed glandular hairs beneath, smooth or slightly scabrous above,
sessile, long and rather broadly decurrent; basal leaves narrowed gradually
into petiole with slightly broadened base, withering early; uppermost
leaves small, linear to filiform, not decurrent. Capitula solitary. Involucre
12-20 mm long, cupuliform, not arachnoid-hairy, five- or six-rowed;
outer involucral bracts more lax, sometimes recurved, green, acute or
subobtuse, reduced, other bracts lanceolate, appressed, purple,
membranous along margin and somewhat serrate, acuminate. Receptacle
densely squamose, scales smooth, reaching almost up to half to involucral
height. Corolla pink, pendulous, 22-28 mm long, narrowed over 8-12
mm, with 5-7 mm long limb, densely covered with sessile glandular
hairs; basal appendages of anthers setaceous; filaments glabrous. Pappus
persistent, snow-white, with unequal dense bristles, bristles short-barbate,
with barbules shorter than thickness of bristles, or almost as long; achenes
four-angled, obpyramidal, about 6 mm long, yellow, smooth, spinose-
toothed above along edges, usually with three-fid teeth, often with row
of similar spinules slightly below margin, with broad cupuliform,
thickened, finely toothed pappogenus at apex, with large five-toothed
nectary in center, on short stalk. Flowering V—VI; fruiting VI-VII(?).

Low-hill steppes, on varicolored rocks, stony slopes.—Soviet Central
Asia: Pamiro-Alai Region (extreme west). Endemic. Described from
Baisun. Type in Leningrad.

Note. In the herbarium of the Botanical Institute, Academy of
Sciences of the USSR, this species has been identified by V.I. Lipsky
as J. trautvetteri var. glabrata Lipsky.

62. J. popovii Iljin in Izv. Glavn. Bot. Sada SSSR, XXVII, 1 (1928)
80.

Perennial. Plants usually with many-headed caudex. Stems numerous,
usually branched in lower part, less often simple, sulcate, glabrous, leafy
up to middle. Leaves densely arranged, oblong, oblong-oval or broadly
lanceolate, undivided and entire, acute, narrowly revolute and strongly

61

—

605

scabrous, green and glabrous on both sides, or slightly scabrous above,
with prominent veins beneath, sessile, broadly decurrent. Capitula solitary,
on long peduncles, medium. Involucre narrowly cupuliform, 15-20 mm
long and 10-15 mm wide, glabrous, with linear or narrowly lanceolate
bracts; outer bracts short, weakly recurved, other bracts appressed;
innermost bracts prominent and acuminate. Receptacle densely covered
with smooth bristles. Corolla pink, exserted from involucre, outer
pendulous, 22-25 mm long, narrowed over 8-10 mm, with about 5 mm
long limb; basal appendages of anthers setaceous, almost not divided,
2-3 mm long; filaments smooth. Pappus persistent, white, densely
barbate; achenes (immature) glabrous, four-angled, toothed above along
edges; pappogerius cupuliform enclosing five-lobed nectary. Flowering
V; fruiting VI(?).

Herb slopes.—Soviet Central Asia: Pamiro-Alai Region. Endemic.
Described from Kugitang near the village of Kizyl-Alma. Type in
Leningrad.

Note. In the herbarium of the Botanical Institute, Academy of
Sciences of the USSR, this species has been identified by M.G. Popov
as J. trautvetteri var. nuda M. Pop.

63. J. cephalopoda Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, VII, 10 (1937) 244.—Ie.: Ibid. Fig. 1.

Perennial. Semishrub up to 75 cm high, with many-headed caudex.
Stems erect, flocculose-arachnoid-hairy, longitudinally sulcate, branched;
branches short, somewhat arcuate, leafless, one-headed. Leaves linear,
acute, green, glabrous above, one-veined, gray arachnoid-hairy beneath,
entire, involute; basal and lower cauline leaves numerous, 10-20 cm
long and 2-5 mm wide, gradually narrowed toward base, somewhat
broadened at base; cauline leaves shorter, sessile, not decurrent. Capitula
on 2.5—6.5 cm long peduncles, terminal, arcuately ascending. Involucre
15-17 mm long, pubescent, greenish, slightly dirty purple above;
involucral bracts appressed, with prominent veins, along margin narrowly,
almost transparently frilled; outermost bracts few, strongly reduced,
oblong-ovate, middle bracts lanceolate, innermost longest, linear-
lanceolate, acuminate. Receptacle densely covered with short, about 5—
6 mm long hairs. Florets purple, ray florets pendulous; corolla 20—22
mm long, narrowed over 9-10 mm, with 4.5—5 mm long lobes of limb.
Pappus persistent, white, 7-8 mm long, with barbate hairs; barbules 2—
3 times as long as hair diameter; achenes 6—7 mm long, 4-angled,
obpyramidal, with dentate corona, teeth at edges decurrent into long
parallel folds, with large, cupuliform, thickened, minutely dentate
pappogenus at apex along edges, enclosing poculiform, 5-dentate nectary.
Flowering IV(?)—V; fruiting V—VI.

612

606

Gravelly and sandy mountain ridges in desert zone.—Soviet Central
Asia: Karatau. Endemic. Described from foothills of Karatau, from the
banks of Kyzyl-Kum. Type in Leningrad.

Series 2. Pinnatae [ljin—Leaves lyrate, pinnatisect or toothed.

64. J. bucharica Winkl. in Tr. Peterb. Bot. Sada, IX, 2 (1886) 525;
O. and B. Fedtsch. Perech. Rast. Turk. IV, 300; Ijin in Tr. Turk. Nauchn.
Obshch. 2, 19.

Perennial. Plants 15-60 cm high. Roots somewhat thick, woody-
fibrous, bearing caudex. Stems solitary or usually many, simple or often
more or less long-branched; branches usually in lower half or two-thirds,
striate and finely sulcate, at base usually arachnoid hairy-woolly-
tomentose, less often weakly tomentose, remaining part flocculose-
arachnoid-hairy, leafy. Leaves whitish-gray beneath due to more or less
densely arachnoid-hairy pubescence, greenish above with only arachnoid-
hairy coating, often glabrescent, or even entirely glabrous, green, with
very fine appressed glandular hairs; basal leaves long-petiolate, petioles
somewhat broadened at base; lower cauline leaves on shorter petioles,
also somewhat broadened at base; middle cauline leaves sessile, usually
with small auricles or somewhat short-decurrent; upper leaves distinctly
reduced, sessile, usually without auricles and not decurrent, lanceolate
or linear; basal, lower, and middle cauline leaves pinnatisect, and then
usually with toothed rachis, or pinnatipartite, pinnatilobate or lyrate,
occasionally all undivided (f. integrifolia Winkl.), with flat margins or
narrow-sinuate, especially into pinnatisect laminas, sometimes in all
leaves or only in uppermost leaves. Capitula solitary. Involucre narrow
cupuliform, (15)20—25 mm long (7)10-15 mm wide, not or weakly
arachnoid-hairy only in upper half, four-or five-rowed; lowermost bracts
oblong-ovate, green, practically without frill or with quite narrow
membranous border; middle bracts lanceolate, acuminate, more broadly
membranous-frilled, pink-purple in upper part; innermost bracts remote,
linear-lanceolate, pink-purple, attenuate in thin subulate cusp, on outer
side, like middle bracts, covered with rather fine crisped hairs, often
with isolated serration; all bracts with prominent veins, especially midrib.
Receptacle covered with more or less densely smooth bristles, up to
one-third as long as_involucre, less often with shorter bristles. Corolla
pink, 22-25 mm long, narrowed over 9-12 mm, with 5—6 mm long
limb, covered with scattered glandular hairs, usually transversely rugose
in middle, outer ones pendulous; basal appendages of anthers setaceous.
Pappus persistent, snow-white, barbate or less often barbate-serrate;
achenes obpyramidal, four-angled, 5-6 mm long, shorter than pappus,
with prominently raised longitudinal ridges, especially thickened edges,

Plate XLV.

1—Jurinea arachnoidea Bge.; 2—J. abolinii Ijin.

613

614

608

in upper one-third modified in teeth, often bifid, with toothed corona
above, glabrous, sometimes with sparse fine glandular hairs between
teeth; with cupuliform pappogenus at apex, usually 10-lobed, enclosing
large tive-fid nectary. Flowering IV—V; fruiting V—VI.

Foothill zone: adyrs’, conglomerates, stony gravelly slopes, rocks.—
Soviet Central Asia: Pamiro-Alai Region. Endemic. Described from
Kodzhakazian. Type in Leningrad.

65. J. psammophila Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada, V,
4 (1924) 58.

Perennial. Plants 20-50 cm high, grayish due to arachnoid-hairy
pubescence, with thick, woody root, bearing densely crowded caudices,
woolly-tomentose. Stems branched. Leaves usually several; basal leaves
numerous, like lower cauline leaves, narrowed into relatively long petiole,
pinnatilobate, pinnatipartite or pinnatisect; weakly sinuate, other cauline
leaves sessile and decurrent; uppermost leaves linear, acuminate,
undivided, with thin, glabrescent, flocculose-arachnoid-hairy tomentum
on both sides. Capitula on very long leafless peduncles, solitary. Involucre
about 25 mm long and 10-15 mm wide, grayish-tomentose; involucral
bracts appressed, linear-lanceolate, acuminate; outer bracts significantly
reduced, innermost finely acuminate, narrowly membranous and purple
along margin. Receptacle covered with short hairs, one-sixth to one-
fifth as long as involucre. Corolla bright pink, glandular-hairy, 25—30
mm long, gradually narrowed over 12-15 mm, lobes of limb 5—6 mm
long, peripheral florets sharply pendulous. Pappus persistent, snow-white,
10-15 mm long, barbate-plumose, barbules four times as long as
thickness of bristles. Achenes 5-7 mm long and about 3 mm thick,
grayish, obpyramidal, with longitudinal ridges, alternating with more or
less deep furrows, continuous below, discontinuous above due to
tubercles, terminating into spines appressed above, with large corona
above, with broad, irregularly sinuate marginally fleshy pappogenus at
extreme tip, enclosing thick, large, five-angular nectary. Flowering V;
fruiting at the end of V and VI. (Plate XL, Fig. 8.)

Stony slopes of foothills, outcrops of conglomerates, sandy steppes.—
Soviet Central Asia: Kyzyl-Kum (southwest). Endemic. Described from
the foothills of the Aktau Mountains in Kyzyl-Kum. Type in Leningrad.

66. J. lasiopoda Trautv. in Tr. Peterb. Bot. Sada, VIII, 2 (1883)
510; O. and B. Fedtsch. Perech. Rast. Turk. IV, 303; Ijin in Tr. Turk.
Nauchn. Obshch. 2, 19.—Exs.: H.F.F.M. No. 492.

Perennial. Plants 30-60 cm high; roots bearing one-headed or few-

“Low foothills bordering Fergana depression—Translator.

615

609

headed caudex, covered with broadened bases of leaves. Stems solitary
or few, somewhat thick, erect, striate-sulcate, arachnoid-hairy, woolly-
tomentose near collar, branched only in upper half or one-third. Leaves
green and glabrous above with scattered, fine, appressed glandular hairs,
grayish beneath due to more or less densely arachnoid-hairy pubescence,
oblong or lanceolate, usually sparsely pinnatipartite or pinnatilobate, less
often pinnatisect, occasionally entire, acute or acuminate, subobtuse,
usually weakly sinuate, narrowly convolute; basal leaves, and sometimes
lowermost cauline leaves, narrowed into more or less long petiole,
abruptly broadened at base; cauline leaves sessile, often somewhat
decurrent, uppermost lanceolate to linear; apical leaves almost entire.
Capitula solitary, on relatively short peduncles, mostly incurved.
Involucre almost cylindrical, 15—20 mm long and 7-10 mm wide, weakly
flocculose-arachnoid-hairy, with appressed bracts, outermost bracts
triangular-ovate, acute; middle bracts oblong to lanceolate, narrowly
purple-bordered, acuminate; innermost bracts lax, narrow-lanceolate,
attenuate into purple membranous cusp; all bracts with distinct veins,
midrib prominent. Receptacle densely covered with smooth bristles, up
to one-third as long as involucre. Corolla pink or pink-purple, pendulous
(in peripheral florets), 14-23 mm long, narrowed over 8-11 mm, with
4—6 mm long limb, weakly transversely rugose in broader part, with
scattered glandular hairs throughout; basal appendages of anthers
setaceous. Pappus persistent, snow-white, 7-9 mm long, with two longer,
short-plumose bristles; achenes obpyramidal, four-angled, 5—6 mm long,
with deeply incised longitudinal ribs, smooth in lower half, spinulose in
upper half along edges, with toothed-spinose upper edge, pappogenus
connate with pappus, fleshy, cupuliform, enclosing five-lobed nectary.
Flowering V; fruiting V—VI.

Rocks, varicolored substrates, gypsum-clayey slopes of foothills —
Soviet Central Asia: mountainous Turkmenia, Tien Shan, Pamiro-Alai.
Endemic. Described from Akhalteke in Turkmenia. Type in Leningrad.

Note. Trautvetter, while describing this species, compares it with J.
adenocarpa Schrenk; however, these are not related species and belong
to different sections.

67. J. karatavica Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada, V, 4
(1924) 60.

Perennial. Plants about 40 cm high, green, weakly arachnoid-hairy.
Stem erect, weakly branched in inflorescence. Cauline leaves narrow,
almost linear, with less often distant and long teeth, convolute, green
on both sides, sessile and short-decurrent; lower cauline leaves not
decurrent, petiolate, uppermost filiform, bracteal. Capitula on short
peduncles. Involucre broadly turbinate, but with roundish-narrow base,

616

610

20-23 mm long and about 10 mm wide, green, with purple tinge in
upper part, not arachnoid-hairy, with sessile glandular hairs; involucral
bracts linear-lanceolate, acute, inner bracts acuminate, with scarcely
visible appressed bristles. Receptacle covered with bristles, almost one-
third as long as involucre: Corolla bright purple, covered with sessile
glandular hairs, 21-25 mm long, narrowed over 10-12 mm, with 5-6
mm long lobes of limb; peripheral florets sharply pendulous. Pappus
persistent, snow-white, 9-10 mm long, barbate-plumose, with plumes
three to four times as long as thickness of bristles; achenes covered
with sharp tubercles along edge and parallel ribs, with patelliform
pappogenus at tip, enclosing nectary. Flowering VI; fruiting VII(?).

Boyalych [Salsola arbuscula] deserts —Soviet Central Asia: Balkhash
Region (not far from the Karatau Mountains). Endemic. Described from
the deserts between Lake Ashchikol and the Karatau Mountains. Type
in Leningrad.

Note. Only one authentic specimen is present in the collections.

68. J. botschantzevii Iljin sp. n. in Addenda XXVI, 722.

Perennial. Plants (15)25—40(50) cm high. Stems usually few, finely
sulcate-ribbed, floccose-arachnoid-hairy, covered with numerous sessile
glandular hairs, branched and leafy almost from base to middle or up to
two-thirds; branches more or less erect, elongate and leafy only on lower
side. Leaves pinnatipartite or bipinnatipartite, pinnate, or pinnatilobate,
with oblong, less often linear, ovate or even triangular lobes, obtuse

and coalesced with more or less broadly winged rachis, always distinctly

finely sinuate, with sessile glandular hairs on both sides, sparsely
arachnoid-hairy, more weakly pubescent above, basal and lowermost
cauline leaves gradually narrowed into long petiole, distinctly broadened
at base; middle cauline leaves sessile, semiamplexicaul, coalesced with
lower part of lamina to the corresponding branch, uppermost leaves
linear. Capitula solitary terminal, with many florets. Involucre cupuliform
to cyathiform, usually 20-25 mm long, entirely covered with soft fluff,
and scattered glandular hairs; outer involucral bracts oblong-ovate; middle
bracts oblong-lanceolate, acuminate, inner long-aristate, usually purple,
glandular-hairy and with short.hairs on outer side and above, besides
soft bristles, with prominent midrib, and two weak lateral veins.
Receptacle squamose; scales acuminate, up to half of involucre. Corolla
pink or pink-purple, with scattered sessile glandular hairs, 22-30 mm
long, gradually narrowed over 10-13 mm, with 5—7 mm long lobes of
limb. Pappus persistent, of very fine bristles, almost as thick as human
hair, with distant, more or less short, barbules on all sides, strongly
lustrous at base; achenes slightly yellowish, distinctly four-angled,
obpyramidal, about 6 mm long, with longitudinal smooth ridges and

617

611

deep furrows, terminating into dense teeth above, almost connate with
corona, with scattered sessile glandular hairs in furrows and between
teeth; pappogenus somewhat thick, five-lobed, cupuliform, with roundish
lobes, very finely toothed along edges, nectary large, five-dentate, teeth
in turn with fine teeth, on rather. short manubrium. Flowering IV—V;
fruiting V—VI. .

Limestone outcrops, stony thin-soiled slopes—Soviet Central Asia:
Pamiro-Alai. General distribution: Possible in Afghanistan. Described
from Souther Tadzhikistan from the Koikitau Mountains northwest of
the village of Aivadzhi. Type in Leningrad.

69.J. winkleri Iljin sp. n. in Addenda XXVI, 722.—J. bucharica
var. integrifolia Winkl. ex O. and B. Fedtsch. Perech. Rast. Turk. IV
(1911) 300.

Perennial. Plants (15)20-50 cm high, forming somewhat dense turf
and many-headed caudex, with woolly buds in axils of upper scales,
and remains of lower leaves. Stems few, erect, flocculose-arachnoid-
hairy, glabrescent and with scattered sessile glandular hairs, finely ribbed-
sulcate, branched up to middle or even three-fourths; branches sharply
spreading upward. Basal and lower cauline leaves, sometimes also middle
leaves, petiolate, petiole shorter than lamina, less often equal; sometimes
upper leaves sessile, semiamplexicaul, with lower part coalesced with
corresponding branch and very short-decurrent, sometimes indistinctly
decurrent; all leaves green or grayish-green above due to fine arachnoid-
hairy pubescence, finely rugose, with reticulate wrinkles, usually
continued all over, and not very strongly depressed veins, white-
tomentose or grayish white-tomentose beneath due to dense arachnoid-
hairy tomentum, also with sessile glandular hairs; lower and middle
leaves oblong-oval, lyrate, less often pinnatilobate, occasionally almost
undivided, toothed or cristate in lower part, lobes and lobules usually
finely toothed along margin; uppermost leaves strongly reduced, oblong
or almost linear. Capitula solitary, with many florets. Involucre 20—25
mm long; all involucral bracts appressed, with arachnoid-hairy
pubescence, membranous border, and prominent veins, especially midrib,
and two pairs of lateral veins on each side; outer bracts oblong-ovate,
acute, like middle bracts, dark green below tip; middle bracts oblong-
lanceolate, acuminate, inner acuminate, less often aristate, purple above,
outer surface pubescent with sessile glandular hairs. Receptacle squamose;
scales acuminate, smooth, one-fourth as long as involucre. Corolla pink
or pink-purple, with scattered sessile glandular hairs, 23-30 mm long,
gradually narrowed over 9-13 mm, with 5-7 mm long lobes of limb.
Pappus persistent, half to two-thirds as long or less often almost as long
as achene, pappus bristles barbate-plumose, with dense, two-rowed .

618

612

barbules, innermost bristles four or five, broad and short-barbate, shorter
than thickness of bristles; pappogenus broadly lobate, fleshy, cupuliform, ~
red on inner side, enclosing cyathiform nectary on short manubrium;
achenes about 5 mm long, thick, sharply four-angled, obpyramidal, light
gray, on edges with numerous, large, compact, parallel ridges, with
narrowed and deep canals, terminating into teeth above, connate with
teeth of corona, latter as well as bottom of canals covered with sessile
glandular hairs. Flowering V—VI; fruiting VI.

Clayey, pebbled adyrs, and conglomerates, also sands and clayey
plains of the Fergana Valley.—Central Asia: Syr-Darya. Endemic.
Described from the Margelan District, eastern slope of conglomerate
ridges near the village of Assake. Type in Leningrad.

70. J. tadshikistanica [jin sp. n. in Addenda XXVI, 723.

Perennial. Plants 20-50 cm high, sometimes developing dense many-
headed caudices, with woolly fluff in the axils of scaly leaves. Stem
entirely covered with arachnoid-hairy pubescence, with scattered sessile
glandular hairs especially along furrows, branched in lower half and
with long, leafless erect branches in upper half, arising at an acute angle.
Leaves usually crowded in lower half of stem, basal and lower cauline
leaves long-petiolate, all leaves lyrate or lyrate-lobate, less often mixed
with pinnatipartite and pinnatilobate leaves, occasionally with almost
undivided leaves, but nevertheless laminas toothed in lower half, lobes
ovate or oblong, obtuse, grayish beneath due to arachnoid-hairy
tomentum, with numerous glandular hairs under tomentum, with
depressed midrib and lateral veins above, strongly finely rugose, with
sessile glandular hairs deep in pits between wrinkles, weakly arachnoid-
hairy, smooth, not crimped along margin; middle cauline leaves sessile,
usually fused with corresponding branch in lower part and often very
short and narrowly decurrent; uppermost leaves strongly reduced, oblong
or oblong-spatulate, undivided and entire or somewhat finely crenate,
usually narrowed toward base. Capitula solitary, with many florets.
Involucre cupuliform, 20-25 mm long, entirely arachnoid-hairy pubescent
with fine crisped hairs, outer involucral bracts oblong-ovate, short-
acuminate; middle bracts oblong-lanceolate, gradually acuminate, with
membranous border, truncate in lower part especially outer bracts,
ciliolate in upper part; all bracts three-veined, innermost narrow-
lanceolate, gradually attenuate in long awn, with membranous purple
frill, with scattered glandular hairs on outer side, besides pubescence,
and with scarcely visible bristles above. Corolla pink or pink-purple,
usually 22-24 mm long, gradually narrowed over 8-10 mm, with 5—7
mm long lobes of limb, and with numerous sessile glandular hairs. Pappus
persistent, pappus bristles less often short-barbate, barbules either shorter

619

613

or one and one-half to two times as long as thickness of bristles,
semiappressed to axis; pappogenus broadly cupuliform, whitish on outer
side, reddish on inner, enclosing five-lobed nectary. Flowering V—VI(?).

Stony and gravelly slopes.—Soviet Central Asia: Pamiro-Alai.
Endemic. Described from the Vakhsh Valley, from the Buryutau ~
Mountains. Type in Leningrad.

Series 3. Apodae [ljin—Plants without stem of almost stemless.

71. J. impressinervis Iljin in Izv. Tadzh. Bazy Akad. Nauk SSSR,
1, 1 (1933) 65.

Perennial. Plants low, about 10 cm high, caespitose. Stem simple,
arachnoid-hairy. Basal leaves numerous, oblong-oval or almost oval,
subacute, somewhat convolute, entire, subglabrous in lower part, dull
green above, finely arachnoid-hairy, glabrescent, with depressed veins,
grayish tomentose beneath, gradually narrowed into petiole; cauline
leaves four to six, smaller, uppermost leaves linear, sessile. Capitula
with about 17 mm long and 10 mm wide involucre. Involucre flocculose-
arachnoid-hairy; outer and middle bracts oblong and lanceolate, acute,
dark green in middle and above; inner bracts linear-lanceolate, acuminate,
purple. Corolla pink, almost pendulous, gradually narrowed over 7—8
mm, with 4-5 mm long lobes of limb. Pappus persistent(?), snow-white,
with unequal bristles, short-barbate, with sparse barbules, slightly
exceeding thickness of bristles. Flowering V.

Montane habitats —Soviet Cental Asia: Pamiro-Alai. Endemic.
Described from Kulyab, from the Takhta-Kuvat Mountains near the
Tashiktash Pass. Type in Dushanbe; cotype in Leningrad..

72. J. apoda Iljin in Izv. Tadzh. Bazy Akad. Nauk SSSR, 1, 1
(1933) 66.

Perennial. Plants stemless or with scarcely visible stem, caespitose,
semishrub with numerous woody caudices, underground over large part,
densely covered with remains of old leaves as narrow scales. Leaves
densely rosulate at tip of caudices, usually 2-4 cm across, oblong or
oblanceolate, obtuse, gradually narrowed into short petiole, entire or
less often weakly toothed, with depressed midrib above, grayish-green,
almost concolorous, finely pubescent or subglabrous, usually 1-2 cm
long. Capitula subsessile, solitary terminal on caudices. Involucre 20—
25 mm long and about 10 mm wide; outer and middle bracts grayish-
green, lanceolate or oblong, acuminate, distinctly attenuate above, inner
bracts very prominent, acuminate, purple, later pale ferruginous. Corolla
pink, 25-30 mm long, pendulous, gradually narrowed over long distance
with about 5 mm long lobes of limb. Pappus deciduous, barbate-plumose,
about 20 mm long, pappus bristles connate into ring, having common

620

614

base with rather large nectary enclosed by fleshy pappogenus; achene
four-angled, about 4 mm long and 2 mm thick, glabrous, smooth, with
toothed border above, spinulose, and with hemispherical pappophorus at
tip. Flowering VI(?)-VII; fruiting VII. (Plate XLVIII, Fig. 2.)

On limestones.—Soviet Central Asia: Pamiro-Alai Region (only in
the Babatag Range). Endemic. Described from Bish-Archa Peak. Type
in Leningrad.

Note. J. caespitosa Winkl. was not formally described by Winkler,
but was annotated in the herbarium with a question about its affinity to
the genus Jurinea. M. Ijin (1925) described and illustrated (Fig.10) this
species in a review of Turkistanian members of the genus Jurinea Cass.
However, the herbarium specimen and the description based on the
incomplete specimen do not satisfactorily demonstrate its specific status.
Therefore, having not referred J. caespitosa Winkl. to the synonyms of
our species, which is distinguished by a caespitose habit, we assign the
name J. apoda Iljin to the Babatag, Koikitau and Kodzha-Kazian
specimens.

Section 8. Olgaea Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XXI (1961) 390.—Capitula narrowly cyathiform, numerous; florets
bisexual; corolla of peripheral florets distinctly pendulous or gradually
narrowed toward base; anther yellowish; filaments scarcely tuberculate.
Achenes oblong, with toothed margin. Pappus deciduous, pappus bristles
connate at base into ring, enclosing five-toothed nectary; only in J.
xeranthemoides I\jin, ring is connate with membranous base, where it
forms patelliform pappogenus, with a nectary in the center.

1. Pappus comprising serrate bristles, inner bristles longest, broad,
flat, exserted from involucre and resembling inner series of

- involucral bracts of Xeranthemum... 76. J. xeranthemoides Iljin
+ All pappus bristles barbate, longest four hairs do not differ in their
MPACUIS IONE QUITCTS cece se ere aeee thee eckson aoe Z:

2. OUECE INVOlUCKal DIaCts, FECUIN CG cos. cee se eee ee a ee 3.
All involucral bracts appressed ....... 73. J. olgae Rgl. and Schm.
3. Plants whitish-grayish due to arachnoid-hairy pubescence; stem
eglandular; outer involucral bracts recurved, whitish arachnoid-
hairy, gradually narrowed upward, without constriction, others
appressed, purple along border and in upper half; corolla pink or
PINK PUNDG re noc ete re esetac ee eset ee 74. J. ferganica [ljin

+ Plants bright green, glabrous or with scarcely visible arachnoid-
hairy pubescence below in early stages. Stem with more or less
densely scattered sessile glandular hairs. Outer involucral bracts
small, usually green, with constriction above, involucre, dark purple

in remaining part, like corolla ............. 75. J. kuraminensis [jin

+

62

—

615

Series 1. Olgaeoides [ljin—All pappus bristles similar; pappogenus
not developed.

73. J. olgae Rgl. and Schm. in A.P. Fedtsch. Putesh. v Turk. II,
No. 18 (1881) 51; Tr. Peterb. Bot. Sada, VIII, 2, 512; O. and B. Fedtsch.
Perech. Rast. Turk. IV, 295; M.M. Iljin in Tr. Turk. Nauchn. Obshch.
2, 15.—Exs.: H.F.A.M. No. 494.

Perennial. Herbaceous plant, often semishrub, 15—40(45) cm high,
with strong fibrous tap root, many-headed. Stem woody below, strongly
branched up to middle, with tomentose fluff in leaf axils in woody part,
simple in upper half, together with leaves grayish, grayish-green (f.
arachnoidea I\jin) due to weak arachnoid-hairy pubescence or glabrous
and green (f. glabra I\jin), without glands or less often with scattered
sessile glandular hairs, angular-sulcate. Leaves sessile, linear; lowermost
leaves subsequently often withering, almost flat, narrowly involute. only
at extreme margin, up to 10 cm long, others involute up to midrib,
green or arachnoid-hairy-grayish on both sides, or strongly arachnoid-
hairy beneath; with depressed midrib above, punctate-alveolate due to
deep-seated glandular hairs, midrib prominent beneath, with scattered
sessile glandular hairs, tip rather’ short cartilaginous above, gradually
reduced upward, usually not reaching up to capitula. Capitula numerous,
solitary terminal on stem. Involucre poculiform, about 2—2.5 mm long
or slightly shorter, finely arachnoid-hairy, usually on lower side, or
without arachnoid-hairy pubescence; involucral bracts imbricate, in many
rows; outer bracts short, some obtuse, other bracts lanceolate, with
prominent midrib and two lateral veins, reddish in upper half, gradually
acuminate, membranous-cuspidate along margin, with or almost without
sessile glandular hairs, but then with scarcely visible, short, stiff hairs,
especially along margin, or with both types together; inner bracts usually
attenuate into aristate cusp, covered with short hairs. Receptacle short
squamose; florets few, peripheral florets pendulous, pink or pinkish-
purple, 20-25 mm long, narrowed over 7—10(12) mm, with 5—6(7) mm
long lobes of limb, slightly unequal, with scattered sessile glandular
hairs; basal appendages of anthers about 3 mm long, setaceous, divided
below; filaments yellowish, tuberculate. Pappus easily shedding entirely,
longer than achene, barbate, with many bristles, snow-white; bristles
connate into ring at base, recurved on maturation, barbules in-middle
part of bristles one and one-half to two times as long as thickness of
bristles; achenes 6—8 mm long and 2—3 mm thick, oblong, obpyramidal,
four-angled, brownish, glabrous, with toothed slightly deflexed corona
above, with a row of sharp two- or three-toothed tubercles below, with
dome-shaped pappophorus at apex, terminating into nectary exserted: from
pappus ring. Flowering IV—VII; fruiting VII(end)-IX. (Plate XL, Figs.
ha (05)

622

616

Adyrs, varicolored rocks, gypsums, red sand-stones, conglomerates,
rocks in foothill zone, stony, gravelly mountain slopes in the zone of
steppes and mountain xerophytes up to 3,000 m.—Soviet Central Asia:
Tien Shan, Pamiro-Alai Region. Endemic. Described from the Zeravshan
Valley not far from Bakhud. Type in Leningrad.

Note. An extremely polymorphic species. The form with arachnoid
tomentum (f. arachnoidea Iljin) is predominant; however, distinctly green,
glabrous or subglabrous forms (f. glabra Iljin) are also .not rare.
Sometimes the latter predominate in some localities, for example, in
the vicinity of the village of Urgut. Almost all the plant organs vary,
especially the vegetative parts.

74. J. ferganica (Ijin) Iljin comb. n.—J. olgae var. ferganica
Ijin in Tr. Turk. Nauchn. Obshch. 2 (1925) 16.—Exs.: GRF No.
2778.

Perennial. Plants 25-40 cm high, with long fibrous tap root. Stem
as also leaves grayish-green due to fine arachnoid-hairy pubescence,
angular-ribbed, somewhat spreadingly branched in lower half, more or
less geniculately at nodes, eglandular, whitish-brown or dirty purple at
base and leafless, with dense tomentose fluff in axils of scales. Leaves
sessile, linear, strongly involute up to midrib, entire, with strongly
appressed sessile glandular hairs above, sparsely arachnoid-hairy, grayish
beneath due to dense arachnoid-hairy cover, subobtuse with small
cartilaginous tubercle at apex, more or less recurved; lower leaves about
5 cm long or slightly longer, gradually reduced upward and completely
absent in uppermost part of stem. Capitula numerous. Involucre narrowly
cupuliform, usually about 20 mm long, less often slightly longer or
smaller; outer involucral bracts short, slightly thick, greenish, not carinate,
arachnoid-hairy, very short-cuspidate, decurved; other bracts significantly
larger than outer, entirely appressed, acuminate, with prominent midrib
and also two lateral veins, usually not arachnoid, with numerous, fine,
sessile glandular hairs and scarcely visible appressed hairs on outer side;
innermost bracts aristate, exceeding all other bracts. Receptacle scales
short, yellowish. Corolla pink, about 22—25 mm long, gradually narrowed
over 7-8 mm length, with 5S—7 mm long lobes of limb, scattered glandular
hairs below, corolla of peripheral florets recurved and pendulous; apical
appendages of anthers acuminate, almost linear, membranous, about 3
mm long, undivided, fimbriate only below; filaments very finely
tuberculate. Pappus deciduous, pappus bristles barbate, barbules in middle
part of bristles usually three to four times as long as thickness of bristles,
reduced above and below, bristles connate into ring; achenes four-angled,
about 7 mm long and 3-4 mm in thick, brown, with smooth dome-
shaped pappophorus at apex, terminating with small five-angled

623

617

membranous nectary. Flowering IV—VIII; fruiting VI-IX. (Plate XLVII,
Fig. 23)

Rocks, screes, gravelly and shaly mountain slopes——Soviet Central
Asia: Pamiro-Alai. Endemic. Described from northern slope of the Alai
Range, Shakhimardan Basin, near the village of Pulgan. Type in
Leningrad.

75. J. kuraminensis Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XXI (1961) 291.

Perennial. Plants 20-35 cm high, with strongly fibrous tap root,
many-stemmed, bright green, glabrous, subglabrous, or weakly arachnoid-
hairy. Stems especially in lower part angular-sulcate, divaricately
branched, somewhat geniculate at nodes, with scattered sessile glandular
hairs. Leaves sessile, linear-filiform, almost cylindrical, revolute up to
midrib, with subacute cartilaginous apex, undivided and entire, both
sides green, punctate-alveolate above, with sessile glandular hairs beneath;
lower leaves 5-8 cm long, lowermost almost flat, gradually reduced
and almost disappearing upward. Capitula numerous, solitary terminal
on stems. Involucre poculiform, not arachnoid-hairy, or arachnoid-hairy
only at base, generally dark purple; outer involucral bracts short, greenish-
yellow, purple along margin, with dark green, slightly extended and
recurved apex; middle and inner bracts entirely dark purple, lanceolate,
acuminate, with brownish-cusp, prominent midrib, and two less prominent
lateral veins, densely covered with sessile glandular hairs, short ciliate
along border and at apex, innermost bracts most prominent, with thin,
brownish, stiff-hairy awn, resembling transition to bristle. Receptacle
squamose; scales white, pointed, smooth, one-fourth to one-third as long
as involucre. Florets less numerous, peripheral florets semipendulous;
corolla purple, 17—19(20) mm long, covered with sessile glandular hairs,
gradually narrowed over 5-8 mm length, with 3—4(5) mm long lobes of
limb; basal appendages of anthers membranous-setaceous, divided below.
Pappus deciduous, barbate, about 10 mm long (excluding four long
bristles); all bristles connate into ring, enclosing nectary. Achenes (6)7—
8 mm long, oblong-linear, glabrous, brownish-olive, with toothed corona
and row of teeth below it, with gradually conical pappophorus at apex
terminating with nectary. Flowering VI-—VII; fruiting VIII-IX. (Plate
XLVII, Fig. 3.)

Gravelly and stony slopes in grass-wormwood steppe of subalpine
zone.—Soviet Central Asia: Tien Shan (Kuramin Range). Endemic.
Described from the Kuramin Range. Type in Leningrad.

Series 2. Biformes Iljin.—Pappus bristles of two types: outer as in
preceding series, and few inner bristles broad, flat, and prominent as in
Xeranthemum.

624

618

76. J. xeranthemoides Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada,
V, 11-12 (1924) 11; Tr. Turk. Nauchn. Obshch. 2, 9.

Perennial. Plants 15—40 cm high, many-stemmed, whitish to grayish-
green, with strong fibrous tap root. Stem angular-sulcate, divaricately
branched in lower half. Leaves sessile, linear, almost subulate, convolute
up to midrib undivided and entire, weakly arachnoid-hairy above, with
sessile glandular hairs in alveolae, more densely arachnoid-hairy beneath,
with short weakly cartilaginous cusp; lower leaves 5—8 cm long, withering
at anthesis, gradually reduced upward and absent below capitula. Capitula
numerous, solitary terminal on stems, narrowly cylindrical during
flowering, later very narrowly poculiform. Involucre 20-25 mm long,
with appressed bracts; outer bracts very short, greenish, arachnoid-hairy;
middle bracts narrowly lanceolate, acuminate, green in middle, broadly
membranous along margin; inner bracts prominent, narrower, gradually
narrowed into awn, purple, like middle bracts with prominent midrib,
and two lateral veins at base, almost entire in upper part, scarcely toothed
in lower part, without arachnoid-hairy pubescence outside, with numerous
sessile glandular hairs. Receptacle squamose; scales yellowish, smooth,
acuminate, one-fifth to one-fourth as long as involucre, less numerous.
Corolla pink-purple, about 20 mm long, narrowed over 5—6 mm length,
with 4-5 mm long lobes of limb, somewhat unequal, with lower lobe
slightly longer in pendulous peripheral florets; basal appendages of
anthers 4-5 mm long, membranous, undivided above, setaceous below;
filaments very finely tuberculate. Pappus not shedding easily, pappus
bristles toothed, of which 4 bristles prominent, thicker, flat, somewhat
carinate, also toothed, resembling inner row of involucral bracts of genus
Xeranthemum; all bristles connate ‘into ring with continuous base, on
which thickened patelliform pappogenus is developed, continued in
stalked nectary at tip. Achenes 5-6 mm long, flattened, four-angled,
narrowly obpyramidal, brownish or olive colored, glabrous, smooth,
corona toothed with dome-shaped pappophorus inside. Flowering V—IX;
fruiting IX—X. ‘(Plate XLVII, Fig. 1.)

On adyrs.—Soviet Central Asia: Pamiro-Alai (Fergana Valley).
Endemic. Described from the Fergana Valley, between the village of
Uch-Kurgan and the Arpalyk Mountains. Type in Leningrad.

Section 9. Insculptae Iljin sect. nova in Addenda XXVI, 723.—
Leaves in all species always differently divided, decurrent usually as
teeth, rows of appendages or toothed wing; pappus, with rare exception,
entirely deciduous, with pappogenus and nectary (exceptions J. asperifolia
Iljin, J. serratuloides Ijin, J. hamulosa Rubtz., and J. lithophila Rubtz.);
achenes variable in shape, usually with dome-shaped or mitriform nectary,
smooth below edges or with band of teeth below corona.

619

APL WHIZ
XR |

oN aN
AN NNT

625 Plate XLVI.

1—Jurinea multiceps \\jin, 2—J. fedtschenkoana Iljin; 3—J. suffruticosa Rgl.

626

620

+

. Pappus entirely deciduous with pappogenus and nectary; achenes

With; pappophorusts33 oil ee) 202i. --- ag hat MR. <<-- RR o Z
Pappus persistent; pappophorus abeent cy Ae a le 3 ET.

. Stem strongly flocculose-arachnoid-hairy; leaves not decurrent,

pinnatisect, with obtuse segments, green above, gray arachnoid-
hairy beneath; involucral bracts flocculose-arachnoid hairy and
strongly uncately recurved (southwestern Tadzhikistan)................
Bo ceeds eine iii 3-0 ok | eae Se. aaa 91. J. baissunensis [lyin
All other species of section with decurrent leaves (however, see,
US atwopurpiirea Wik.) ..... PRPS Saccc. SeatesnaectcoMehss ee aac BES J

. Plants stronger, with 40-60 mm wide capitula (including florets);

outer and middle involucral bracts strongly uncately or arcuately
HEL) TCL Wipe cer hae | naa alae, ea Om ima MOREY. fo | MBP 4.
@apitula amuch ‘smaller .. 1530 etter... 5. Ben. Oe. «sch ence tae Pescesks Se

. Basal and lower cauline leaves usually bipinnatisect, usually green

on both sides; outer involucral bracts arcuately recurved and finely
spinescent, covered with short bristles from base of recurved part;
pappus about 10 mm long, with barbate bristles, barbs exceeding
thickness of bristles; achenes smooth, without teeth along ribs
(northern ranges of Pamiro-Alai) .............. 77. J. maxima Winkl.
Basal and lower cauline leaves pinnatisect, usually bipinnatisect,
but with entire terminal lobe; outer involucral bracts with thin
arachnoid-hairy pubescence in recurved part and very short bristles
only at tip; pappus 12-15 mm long, with serrate bristles, always
shorter than their thickness; achenes with few excurrent teeth below
corona along ribs (southern foothills of Pamiro-Alai) ...................
eicgoAl be Cae nares em Mamet PES A Sd 78. J. bipinnatifida Winkl.

-’ @orolla’ blackish-purple .. 25. Goes 4.4...:.. ee ee 6.

Corolla; pinkspurple '\\12 ee eee eee cee 2... ee oe en Te

. Leaves pinnatipartite, with remote lobes, somewhat hard, almost

not decurrent; outer and middle involucral bracts sharply recurved
and also somewhat hard; pappus bristles short-barbate (Babatag
and adjoining adyIs............c::cesccees 89. J. atripurpurea Wink.
Leaves lyrate, thin, distinctly decurrent with row of oblong
appendages; outer involucral bracts thin, scarcely recurved; pappus
bristles plumose-barbate (Western Pamir) .............:..cccccceseecesseeeenees
Bh carb SR i 4 Af A RR. aeeei 82. J. propinqua I\jin

. Cauline leaves decurrent with row of isolated oblong appendages

Coe eosasesescersessse essa eseEeee EEE EEE EHD Ee EEE E EEE E EEE DEO EET EESSSOH SEE SEECEE EE EEEE SESE EEE EEEESEL

Cauline leaves entirely decurrent or with sparsely toothed wing =
SOMENPlACES MeN EIN Loree reces aaecere eenrne ean Tene en an eee ae oh

. Stem erect; outer and middle involucral bracts long, erect,

contorted, as in J. tortisquamea I\jin; pappus bristles short-barbate;

+ Stem bent and leafless above; cauline leaves finely pinnatisect;
outer involucral bracts somewhat divergent, semiappressed, all other
bracts appressed; achenes glabrous and smooth (Gissar, Zevar) ..
Sat A ee AMS LO a 80. J. serpenticaulis [ljin

9. Leaves weakly sinuate, smooth; outer and middle involucral bracts
shortly recurved; pappus bristles plumose; -achenes with sessile
glandular hairs (Dzhungarian Alatau).... 85. J. semenovii Winkl.

+ Leaves not sinuate, rough; involucral bracts appressed or outer
bracts lax; pappus bristles barbate, achenes eglandular........... 10.

10. All leaves linear, undivided and entire; all involucral bracts
appressed; achenes glabrous, smooth, with longitudinal striations
(Gissar Range, Kshtut)........00.000.... 83. J. helichrysifolia M. Pop.

+ All leaves, except uppermost, oblong, pinnatilobate or pinnately

— toothed; outermost involucral bracts slightly divergent; achenes
glabrous and smooth, with longitudinal striations (Nuratau Range,
Lyangan) Sie ee. Ae Ee eee 2 84. J. zakirovii Ijin

11. Cauline leaves sessile and decurrent as row of longitudinal
appendages; pappus bristles short-barbate; achenes glabrous and
SHITOOUHID, 202 SENG, Sots eee amen a erecta eee cen tT Pe na Oe koe es

HP appust DrIstles’ PIUMOSe Sie s.c!.- ta eece st eee eee gis

12. Leaves lyrate with large, attenuate terminal lobes (Shakhrisyabs)
U2, SMES TRUE AE. ARERR, AULA cote, oan ere 81. J. asperifolia [jin

+ Leaves pinnately divided (Denau) ........ 90. J. sangardensis [jin

13. Plants bright and light green; leaves somewhat hard and rough,
pinnatisect to pinnately toothed; all involucral bracts appressed
(SauraRange wer 208 Se aR ee 88. J. serratuloides [lin

+ Plants with linear undivided and entire leaves, less often with isolated
teeth; outer and middle involucral bracts uncate (Dzhungarian
Auber) Uae se ee, SU, SSO MRE SEY, SEE ahaa ee tees 14

14. Plants arachnoid hairy-tomentose............ 86. J. hamulosa Rubtz.

Hiab lants: slabrous Wien OMe ee! 87. J. lithophila Rubtz.

Series 1. Appendiculatae Iljin——Leaves decurrent as row of isolated
appendages.

621

achenes glabrous, smooth, with a band of spines only below corona
(Gissar Range, Tupalang) ............... 79. J. pjataieviae Ijin

77. J. maxima Winkl. in Tr. Peterb. Bot. Sada, XI, 2 (1892) 336;

O. and B. Fedtsch. Perech. Rast. Turk. IV, 302 ex p.; Ijin in Tr. Turk.
Nauchn. Obshch. 2, 24.

Perennial. Plants 25-80 cm high; root strong, short, developing dark _

thread-like adventitious roots. Stems few, erect, somewhat thick, simple,
ribbed-sulcate, covered with sessile glandular hairs and crisped many-

628

622

celled hairs, flocculose-arachnoid hairy above, especially below
capitulum. Basal and lower cauline leaves petiolate, with lobes decurrent
along petiole, up to 25 cm long, others sessile, very short-excurrent
with row of fimbriate appendages; all leaves usually bipinnatisect right
up to apex, with oblong-linear pinnae or only spatulate or toothed, green
on both sides and covered with crisped many-celled hairs, less often
grayish beneath due to fine arachnoid hairy pubescence, with scattered
sessile glandular hairs, somewhat rugose above, with strongly depressed
veins (raised beneath); uppermost leaves small, simple, absent in upper
half of stem. Capitula solitary, multiflorate, up to 60 cm wide. Involucre
hemispherical, 20-25 mm long, lower part floccose-arachnoid-hairy, often
glabrescent; outer rows of involucral bracts distinctly arcuately recurved,
dark green from base of recurved part, terminating into long and thin
spiny cusp, somewhat thick, usually punctate-alveolate, always covered
with short bristles throughout, yellowish in basal decumbent part, without
bristles, with almost inconspicuous or poorly developed midrib; other
bracts appressed, acuminate, membranous along margin, with prominent
veins, somewhat ciliate along margin, always distinctly or less visibly
covered on outer side with short semiappressed bristles; receptacle
densely squamose with long, thin acuminate scales. Corolla purple, with
sparsely scattered sessile glandular hairs, 22-25 mm long, narrowed over
10-14 mm length and with 5-8 mm long lobes of limb; apical
appendages of anthers 3 mm long, finely attenuate; basal appendages
about 2.5 mm long, setaceous-caudate, bifid; filaments glabrous but not
entirely smooth; styles glabrous; stigma about 2 mm long, divided up to
base into divergent branches, papillate, with tuft of basal bristles. Pappus
shedding only when extremely mature, snow-white, usually about 10
mm long, two bristles exceeding others, one and one-half to two times
longer, bristles connate into ring forming membranous base, with
cupuliform pappogenus, with cyathiform, five-angled and five-toothed
nectary in center, with furrows equal or one and one-half to two times
exceeding thickness of bristles; achenes obpyramidal, sharply four-angled,
6—7 mm long, whitish, glabrous, smooth, uniformly toothed above along
margin. Flowering IV—VI; fruiting VIIJ—VIII.

Alpine steppes——Soviet Central Asia: Pamiro-Alai (Zeravshan and
Gissar ranges). Endemic. Described from the Zeravshan Valley, near
Kshtut. Type in Leningrad.

78. J. bipinnatifida Winkl. in Tr. Peterb. Bot. Sada, XI, 2 (1892)
322; O. and B. Fedtsch. Perech. Rast. Turk. IV, 302, ex p.; Iljin in Tr.
Turk. Nauchn. Obshch. II, 25.

Perennial. Plants 25-80 cm high; root short, non-fibrous, producing
strong, filiform, adventitious roots. Stems few, erect, simple or occasionally

629

623

with solitary branches, cylindrical, distinctly ribbed-sulcate, densely
covered with sessile glandular hairs, lower part covered with flexuous,
many-celled, granose appressed hairs, finely arachnoid-hairy above,
especially below capitulum, or only weakly flocculose-arachnoid-hairy,
leafless above or with solitary filiform leaves. Basal leaves up to 25—40
cm long, petiolate like lowermost cauline leaves, with lowermost
segments of lamina distinctly divergent, oblong to almost linear, entire
or toothed, lamina pinnatisect in middle part, almost undivided,
pinnatipartite, pinnatilobate, or even deeply toothed in upper part, with
large, almost undivided terminal lobe; cauline leaves less numerous,
sessile, with excurrent row of parallel, long, linear appendages, strongly
reduced upward, finely divided; all leaves green and rugose above, with
scattered, granose, somewhat thick hairs, sessile glandular hairs, and
somewhat depressed veins, very narrowly revolute, usually with thin
arachnoid-hairy coating beneath and with similar granose hairs, along
prominent veins, having dense, large, lustrous, sessile glandular hairs.
Capitula hemispherical, up to 40—60 cm wide, solitary, with many florets.
Involucre about 25 mm long; outer bracts strongly recurved, partially
sinuate; recurved part herbaceous, green, somewhat thick, slightly
corrugated, finely arachnoid-hairy from its base, only modified in
lowermost part into short prickle, covered with very short bristles; middle
and strongly exserted inner bracts narrowly lanceolate; all bracts
appressed, attenuate into long cusp, with distinctly prominent veins,
sessile glandular hairs and short, scarcely visible hairs. Receptacle densely
squamose; scales long, smooth, acuminate. Corolla purple, densely
covered with sessile glandular hairs, 25-28 mm long, narrowed over
13-16 mm, with 5-6 mm long lobes of limb; apical appendages of
anthers about 3 mm long, abruptly attenuate into thin cusp; basal
appendages caudate-setaceous, 2-3 mm long; style glabrous; stigma
divided up to base into two divergent branches with short tuft of basal
bristles. Pappus deciduous, 12-15 mm long, snow-white, many-rowed
and with unequal bristles, bristles always serrate, with serrations shorter
than thickness of bristle, connate into ring, surrounding nectary; achenes
oblong-obpyramidal, four-angled, whitish, 6—8 mm long, glabrous, with
longitudinal, parallel, not strongly prominent, ribs, with few teeth along
ribs below toothed corona, pappophorus obtusely conical, inside corona.
Flowering IV—V; fruiting V—VII.

Varicolored rocks, red sandstones, pistachio plantations on adyrs up
to 1,500 m in foothills—Soviet Central Asia: Pamiro-Alai. Endemic:
Described on the basis of a single specimen from a place near the source
of the Chiglin in Tadzhikistan. Type in Leningrad.

79. J. pjataieviae Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XX (1961) 348.—Ie.: Ibid, Fig. 3.

630

624

Perennial. Plants 40-80 cm high, many-stemmed. Stems usually
branched, with few long upward spreading branches, finely sulcate, green,
weakly arachnoid-hairy, especially below capitulum. Basal leaves
petiolate, pinnatipartite or less often pinnatisect, with larger oblong-
oval terminal lobe and many oblong lateral lobes, reduced toward petiole,
less often almost lyrate; cauline leaves sessile, short-decurrent as wing,
consisting of two parallel rows of appendages-segments of leaves, with
upper lobe longer and only slightly broader; all leaves green and glabrous
above or with poorly visible papillae, grayish arachnoid-hairy beneath
due to fine tomentum, both sides with fine, sessile glandular hairs, lobes
of leaves usually entire, not rough, upper half or one-third of stem
leafless, or equipped with sparse, small, filiform, deciduous apical leaves.
Capitula solitary, with many florets. Involucre cupuliform, during fruiting
broadly cupuliform, about 15-20 mm long; outer bracts herbaceous or
with herbaceous tip, lax, contorted, but all bracts erect (similar to J.
tortisquamea), not recurved; middle and inner bracts broadly lanceolate,
membranous-scarious, glabrous, usually with five to seven parallel veins,
acuminate and here very finely toothed, pubescent at apex and weakly
arachnoid-hairy; all bracts appressed. Receptacle densely squamose; scales
numerous, setaceous, acuminate, almost up to half as long as involucre.
Corolla pink at apices of lobes and purple along margin, 18-20 mm
long, narrowed over 8-10 mm, with 5—6 mm long lobes of limb; basal
appendages of anthers caudate-setaceous, about 2—2.5 mm long, strongly
divided, crisped. Pappus snow-white to smoky, 8-10 mm long, of
numerous, barbate, unequal bristles, with barbules as long as or one and
one-half times as long as thickness of bristles; achenes brown, four-
angled, with unequal teeth of corona at apex, glabrous, with few rows
of spinules only below corona, immature achenes 5—6 mm long.
Flowering VI; fruiting VII—VIII.

Montane steppes.—Soviet Central Asia: Pamiro-Alai. Endemic.
Described from the valley of the Tupalang River in the Gissar Range.
Type in Tashkent.

80. J. serpenticaulis Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XXI (1961) 384.—Ie.: Ibid., Fig. 3.

Perennial. Plants 40-60 cm high, many-stemmed taproot fibrous,
with many-headed caudex bearing woolly stems at base, branched only
in lower part, simple and leafless above, straight or slightly flexuous;
branches upward spreading, greenish, slender, about 1 mm thick, about
2 mm thick at base, finely ribbed-sulcate, very sparsely covered with
thin arachnoid-hairy pubescence, more densely arachnoid-hairy below
capitulum, with sparse sessile glandular hairs. Basal and lowermost
cauline leaves 10-15 cm long, gradually narrowed into petiole; cauline

631

632

625

leaves sessile, short-decurrent as wing consisting of few parallel rows
of appendages, imparipinnate-pinnatisect, with attenuate, narrow-
lanceolate or often linear, entire, terminal lobe, only with isolated, linear,
solitary teeth on lower part, revolute, reduced toward base; all leaves
with depressed veins, rugose, scabrous above due to acute cartilaginous
papillae, punctate-alveolate, with sessile glandular hairs, weakly
arachnoid-hairy, glabrescent, sparsely arachnoid-hairy beneath, with
sessile glandular hairs and prominent veins; uppermost leaves almost
filiform. Capitula solitary terminal on stem and branches, with many
florets. Involucre cylindrical at anthesis, 8-10 mm long, on roundish
base, broadened to 15—18(20) mm at fruiting, almost cupuliform,
arachnoid-hairy below; outer bracts attenuate into herbaceous-green apex,
more or less contorted and erect, other bracts entirely appressed,
lanceolate, acuminate, purple along margin and tip, with membranous
margin and usually seven prominent veins, very short-ciliate (serrate),
on outer side densely covered with scarcely visible decumbent and
isolated sessile glandular hairs; innermost bracts acuminate, broadly
membranous along margin, recurved, otherwise like middle bracts.
Receptacle squamose; scales smooth, acuminate from middle. Corolla
pink-purple, bordered with darker purple margin, with sessile glandular
hairs, about 20 mm long, narrowed over 9-10 mm, and 5—6 mm long
lobes of limb; apical appendages of anthers 2—2.25 mm long, abruptly
attenuate upward into thin incurved tip; basal appendages setaceous-
caudate, 2.5—3 mm long; filaments thick, more or less smooth; style
glabrous; stigma 2.5-3 mm long, divided almost up to base, densely
covered with divergent small acute papillae, with short, erect setae at
base. Pappus deciduous, with fused base, enclosing obtusely five-toothed
nectary, smoky, 8-10 mm long (excluding two long bristles), bristles
barbate, as long as or one and one-half times as long as thickness of
bristles, two long bristles quite prominent, thicker, serrate, all bristles in
immature condition not connate, deciduous, later connate into ring;
achenes immature, brownish, obpyramidal, four-angled, glabrous, smooth,
with toothed upper edge, about 5—5.5 mm long, with smooth dome-
shaped pappophorus at tip, with column in center penetrating pappus
base bearing five-toothed obtuse nectary.

On limestone slopes.—Soviet Central Asia: Pamiro-Alai. Endemic.
Described from the southern slopes of the Gissar Range, from the vicinity
of the village of Zevar. Type in Tashkent.

81. J. asperifolia Iljin in Tr. Turk. Nauchn. Obshch. 2 (1925) 25.—
Ic.: Ibid., Plate VI.

Perennial. Plants 30-80 cm high. Stems few, erect, simple or less
often with solitary branches, cylindrical, finely ribbed, sulcate, with

633

626

scattered, fine, sessile glandular. hairs and granose, long, somewhat thick
hairs, finely arachnoid-hairy above, especially densely arachnoid-hairy
below capitula. Basal leaves short-petiolate, like lower and middle cauline
leaves; cauline leaves sessile, lyrate, short-decurrent with row of parallel
fimbriate segments, terminal lobe rather large, oblong-lanceolate,
gradually acuminate and decurrent up to base lateral segments, that are
sparse, oblong-oval or oblong-linear, absent in upper one-third of stem;
uppermost leaves bracteal, almost undivided, linear; all leaves weakly
rugose and scabrous above due to scattered, somewhat stiff, granose,
long hairs, with fine sessile glandular hairs, arachnoid-hairy beneath,
with similar granose hairs along veins, and numerous sessile glandular
hairs. Capitula solitary, with many florets. Involucre cupuliform, 22—28
mm long; outer involucral bracts usually flocculose-arachnoid-hairy, dark
green in upper part, arcuately recurved, finely spinescent, with very
short bristles in outer part under arachnoid-hairy coating; other bracts
appressed, with prominent veins having purple tinge, covered with very
short appressed bristles; innermost bracts aristate, membranous along
margin, scarcely ciliate. Receptacle squamose; scales relatively long,
acuminate. Corolla pinkish-purple, often with sessile glandular hairs 20—
25 mm long, gradually narrowed over 10-15 mm, and with 4-6 mm
long lobes of limb; apical appendages of anthers about 3 mm long,
abruptly attenuate into thin apex; basal appendages setaceous, about 3
mm long, filaments smooth, distinctly thickened; style glabrous; stigma
about 3 mm long, divided up to base, finely papillate with tuft of short
bristles. Pappus always dirty-smoky, short-barbate, with barbules slightly
exceeding or two times as long as thickness of bristles, on an average,
about 10 mm long (excluding two long hairs); achenes (not completely
mature) 5-6 mm long, oblong-obpyramidal, four-angled, pale, smooth,
with toothed corona at apex, and row of additional teeth below it.
Flowering VI-—VII; fruiting VII.

Limestone rocks and screes; shale screes.—Soviet Central Asia:
Pamiro-Alai (northwest). Endemic. Described from the Shakhak Pass in
the Shakhrisyabs Region (southern Uzbekistan). Type in Leningrad.

82. J. propinqua [ljin in Bot. Mat. Gerb. Glavn. Bot. Sada, V
(1924) 10; [jin in Tr. Turk. Nauchn. Obshch. 2, 26.

Perennial. Plants about 50 cm high; rhizome short, producing strong
filiform adventitious roots, less often with tap root. Stems few, simple,
finely sulcate, weakly flocculose-arachnoid-hairy, covered with gronose,
short, crisped hairs, leafy only below. Leaves lyrate, with large oblong
subobtuse or slightly subacute terminal lobe, and many lateral lobes or
small lobules, gradually reduced toward base, usually subobtuse. Basal
leaves short-petiolate; cauline leaves sessile, somewhat decurrent as lobes

634

627

or teeth; all leaves scabrous above due to scattered, short, mitriform,
somewhat stiff hairs, dark green, grayish beneath due to similar, but
soft and dense, thicker hairs, dense and longer along veins. Capitula
solitary, with many: florets, 2.5—3 cm wide. Involucre cupuliform, 1.5-
2 cm long, usually four-rowed, more or less dark purple; outer bracts
irregularly recurved, usually weakly arachnoid-hairy, often dark green, |
others appressed, narrowly lanceolate, finely acuminate, innermost bracts
long-aristate, acuminate, with 3—5 prominent veins, covered with short
bristles, ciliate. Receptacle densely squamose; scales smooth, acuminate,
almost one-third to half as long as involucre. Corolla dark purple,
glabrous, less often with isolated sessile glandular hairs, about 18-20
mm long, narrowed over 7-8 mm, with 4-5 mm long lobes of limb;
apical appendages of anthers about 3 mm long, abruptly finely acuminate
at apex, basal appendages caudate-setaceous, about 3 mm long; filaments
finely tuberculate; stigma about 3 mm long, with finely papillate
divergent branches, basally with short tuft of erect bristles; style glabrous.
Pappus deciduous(?), snow-white, 8-10 mm long (excluding bristles),
many-rowed, bristles unequal, plumose-barbate, with barbules several
times exceeding thickness of bristles; pappus bristles weakly connate
into ring; achenes immature, with toothed corona, glabrous, toothed in
upper half, teeth in few rows, glabrous elsewhere. Flowering V—VI. (Plate
XLIX, Fig. 2.)

High-mountain steppes of western Pamir.—Soviet Central Asia:
Pamiro-Alai. Endemic. Described from Shungan in the Garmchasma
River Valley. Type in Leningrad.

Series 2. Integroalatae Iljin—Leaves decurrent on undivided wing;
pappus barbate.

83. J. helichrysifolia M. Pop. in Tr. Uzb. Gos. Univ., Nov. Ser.
27, Biol., 14 (1941) 105, Fig. 4; Addenda XXVI, 724.

Perennial. Plants 20-40 cm high, with strongly fibrous root, bearing
many-headed vertical rhizomes, aerial parts covered with fine tomentum.
Stems few, erect, simple, yellowish-greenish; cylindrical, finely sulcate-
barbate, covered with scattered, short, crisped hairs and fine sessile
glandular hairs, not arachnoid-hairy or only below capitulum. Basal leaves
and leaves on vegetative shoots gradually narrowed into long, basally
broadened petiole, up to 10-12 cm long and up to 5 mm wide, somewhat
revolute; cauline leaves sessile, narrowly decurrent, up to middle of
stem; all leaves linear, undivided and entire, semiappressed, cuspidate
with brown cartilaginous cusp, grayish-green, with depressed midrib
above, strongly rugose, punctate-alveolate, scabrous due to short
cartilaginous papillae, weakly flocculose-pubescent, revolute, usually up

635

628

to: midrib, with sessile glandular hairs in pits, more strongly arachnoid-
hairy beneath and also with glandular hairs; uppermost leaves strongly -
reduced, filiform. Capitula solitary with many florets. Involucre
cupuliform, about 20 mm long, imbricate, usually four-rowed, appressed;
outer bracts oblong, yellowish, dark green above, short-cuspidate, with
few prominent veins; middle bracts lanceolate, with prominent veins,
especially midrib, with longer cusp, dirty purple in upper half; innermost
bracts usually involute, long-aristate; all bracts covered on outer side
with sessile glandular hairs and isolated arachnoid hairs, small-toothed,
sometimes outer bracts with stalked glandular hairs in lower part, corolla
similar (without pendulous peripheral florets), pink-purple, with scattered,
fine, sessile glandular hairs, 17-18 mm long, narrowed over 5—6 mm,
with 4—5 mm long lobes of limb; apical appendages of anthers about 3
mm long, abruptly attenuate into cusp; basal appendages setiform, 2.5—
3 mm long, divided below; style glabrous; stigma divided almost up to
middle, covered with subacute papillae, with erect short bristles below
branching. Pappus snow-white, barbate (like true Jurinea); achenes
immature, glabrous, obpyramidal, four-angled, with toothed corona above,
and with distinct longitudinal folds.

Soviet Central Asia: Pamiro-Alai. Endemic. Described from the spurs
of the Gissar Range in the Kshtut Region, on the left bank of the Negnot
River. Type in Tashkent.

Note. M.G. Popov gives a short description of this species only in
the Russian language. Besides, the accompanying figure gives only a
general impression. I have provided a more detailed description and
consider it proper to retain M.G. Popov as the authority for this good
species. The author placed it near J. olgae Rgl. and Schm., with which
I do not agree.

84. J. zakirovii Iljin sp. n. in Addenda XXVI, 723.

Perennial. Plants up to 50 cm high; rhizomes few, vertical, densely
covered with scales of remains of petioles, with woolly buds in axes.
Stems few, erect, sometimes ascending at base, with few elongate
branches, cylindrical, sulcate-ribbed, covered with fine sessile glandular
hairs and scattered, short, crisped, white hairs, arachnoid-hairy below
capitula, leafless in upper half or one-third. Basal leaves and leaves on
young branches 10—15 cm long, narrowed into basally broadened petiole,
oblong, pinnatilobate or pinnatidentate, with oblique, opposite, remote,
obliquely rising lobes and teeth and acuminate, gradually acuminate
terminal lobe with cartilaginous cusp, very narrowly convolute; cauline
leaves sessile, distinctly decurrent, entire or rarely with one to two pairs
of triangular teeth; all leaves dull green on both sides, with depressed
midrib above, scabrous due to stiff, short, crisped hairs, strongly rugose,

629

with fine sessile glandular hairs, lower surface with prominent midrib
and distinctly craspedodromous venation, longer, crisped, scattered hairs
and dense sessile glandular hairs, otherwise similar to basal leaves.
Capitula solitary, with many florets. Involucre cupuliform, 15-20 mm
long, basally appressed, yellowish-green, four-rowed; outer and middle
row bracts oblong-lanceolate, with green recurved apices, cuspidate; inner
bracts narrowly lanceolate, gradually acuminate, innermost gradually
attenuate into long, aristate purple cusps; all bracts ciliate with numerous,
fine, sessile glandular hairs on outer surface interspersed with papillae.
Receptacle squamose; scales acuminate, smooth, one-fourth as long as .
involucre. Corolla pink, 16-18 mm long, narrowed over 5—6 mm, with
4—S mm long lobes of limb, densely covered with fine sessile glandular
hairs; apical appendages of anthers white, about 3.5 mm long, abruptly
attenuate into fine cusp, basal appendages setose, 2.5—3 mm long;
filaments finely tuberculate; style glabrous; stigma at base with short,
erect bristles, its branches almost as long as fused part, covered with
fine papillae. Pappus deciduous, with bristles connate into ring, slightly
yellowish due to unequal barbate bristles, two inner bristles considerably
longer than others; achenes obpyramidal, four-angled, 5 mm long,
glabrous and smooth, with longitudinal, poorly developed striations and
uniformly toothed corona above, along margin enclosing conical
pappophorus, with nectary in center on elevation passing through ring.
Flowering V; fruiting V—VII(?).

Soviet Central Asia: Pamiro-Alai. Endemic. Described from
Yangokla near Lyangara in Nuratau. Type in Tashkent.

Series 3. Montanae Iljin.—Leaves short-decurrent with entire wing;
pappus plumose.

85. J. semenovii Winkl. in Tr. Peterb. Bot. Sada, XI (1889) 170;
O. and B. Fedtsch. Perech. Rast. Turk. IV, 297; Iljin in Tr. Turk. Nauchn.
Obshch. 2, 15.—J. tianschanica auct. non Rgl. and Schm.: O. and B.
Fedtsch. op. cit—Saussurea semenovii Herd. in Bull. Soc. Nat. Mosc.
XL, 2 (1867) 142.

Perennial. Plants 40-50 cm high. Stems solitary or few, covered
with remains of old leaves at base, with large woolly buds in their
axils, branched in lower half, with long, arcuately divergent branches;
branches cylindrical, ribbed-sulcate, weakly and sparsely arachnoid, more
densely above, particularly below capitulum, with scattered sessile
glandular hairs. Leaves pinnatipartite, with three or four pairs of oblong,
distant lobes, usually obliquely rising above, cuspidate; uppermost and
lower lobes reduced to large teeth, with upper lobe more attenuate,
rugose above, with sessile glandular hairs in pits and depressed arachnoid-

637

630

hairy midrib, more strongly arachnoid-hairy beneath, with scattered sessile
glandular hairs, green, convolute; basal and lowermost cauline leaves
gradually narrowed into petiole, others sessile and slightly decurrent,
becoming smaller but undivided upward. Capitula solitary. Involucre
broadly poculiform, 20-25 mm long, somewhat arachnoid-hairy, usually
green; outer bracts with recurved, dark green tip, acute or subobtuse,
others densely covered with sessile glandular hairs, appressed; middle
bracts oblong, with short brownish apical spine, prominent midrib, and
1—2 lateral veins; innermost bracts attenuate into cuspidate tip, brownish
purple, very short-ciliate. Receptacle densely squamose; scales broad,
acuminate. Florets numerous; corolla pink-purple, with numerous
glandular hairs, 20-23 mm long, gradually narrowed over 7-8 mm, with
4-5 mm long lobes of limb; apical appendages of anthers 3.5-4.5 mm
long, gradually and finely acuminate, with slightly incurved tips, basal
appendages membranous, divided up to middle, 2.5—-3.5 mm long; anthers
glabrous; filaments strongly tuberculate; style glabrous; stigma surrounded
at base with short, obliquely erect hairs, stigma branches about 3 mm
long, covered with dense papillae on outer side, arising slightly above
its base. Pappus deciduous, many-rowed, about 15 mm long (including
long bristles), plumose, with two very conspicuous bristles, connate at
base into ring, shedding entirely on separation from achéne, ring
contiguous with membranous bottom, with pentagonal, cyathiform
membranous-fimbriate nectary. Achenes oblong, four-angled, smooth,
pale, with scattered glandular hairs, regularly toothed, 6 mm long corona
above, enclosing narrowly conical whitish pappophorus, from which
pappus detaches easily. Flowering VIII.

Sands, stony places.——Soviet Central Asia: Dzhungaria-Tarbagatai
(Dzhungarian Alatau), Balkhash Region. General distribution: Possibly
growing in Sinkiang. Described from the lower reaches of the Ili and
Chingilda rivers. Type in Leningrad.

86. J. hamulosa Rubtz. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, VIII, 5 (1940) 64.

Perennial. Plants 40-60 cm high, with few stems. Stems cylindrical,
sulcate, whitish due to glabrescent more or less dense te .néntum, covering
sessile glandular hairs, branched above base; branches upward spreading,
up to middle or above, long. Basal and lowermost cauline leaves
petiolate, withering before anthesis, with woolly buds in their axils,
others sessile; lower leaves usually somewhat decurrent, middle and upper
not decurrent; all leaves linear, strongly revolute, often up to midnb,
undivided and entire, less often lower leaves with isolated teeth,
cuspidate, glabrous or subglabrous, lustrous, rugose and punctate-alveolate
above, with arachnoid-hairy coating only, strongly along midrib, grayish-

638

631

white beneath due to sparse arachnoid-hairy pubescence, with numerous
sessile glandular hairs, obliquely spreading upward, reduced upward to
short setaceous. Capitula solitary with many florets. Involucre poculiform,
20-23 mm long, arachnoid-hairy; lower and often middle involucral
bracts dark green, narrow-brownish along margin, somewhat thick,
recurved; middle bracts broadly oblong, greenish, with prominent midrib
and two less raised lateral veins, short-acuminate and cuspidate at apex,
membranous-fimbriate along margin, yellowish, with scattered glandular
hairs, ciliate; inner bracts gradually narrowed into tuberculate spine,
broadly membranous-fimbriate, ciliate along border, with sessile glandular
hairs and appressed short hairs on inner side, more dense at tip. Corolla
pink-purple, 20-22 mm long, narrowed over 6—7 mm, with 3-4 mm
long lobes of limb, with scattered sessile glandular hairs, less often
with mixture of stalked hairs; apical appendages of anthers about 3.5
mm long, gradually attenuate into thin, slightly incurved awn at tip;
basal appendages setaceous, divided, 3—3.5 mm long; filaments
tuberculate; style glabrous; stigma divided up to two-thirds, branches
papillate. Pappus snow-white, persistent, with plumose, many-rowed
bristles, two of them longer than others, inner bristles weakly connate
into ring, outer bristles shorter, pérsistent; achenes oblong, 6—7 mm long,
four-angled, almost smooth, glabrous, glandular-hairy, with toothed
corona at tip, enclosing membranous obtuse-angular nectary. Flowering
VINI-IX; fruiting IX.

Rocks.—Soviet Central Asia: Dzhungaria-Tarbagatai. Endemic.
Described from the Dyuvantau Mountains along the southern slopes of
the Dzhungarian Alatau. Type in Alma-Ata; cotype in Leningrad.

87. J. lithophila Rubtz. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, VIII (1940) 64.

Perennial. Plants 40-60 cm high. Stems branched with long, upward
spreading branches slightly above base in lower half, dark dirty purple
below, green above, glabrous or with isolated arachnoid hairs and sessile
glandular hairs, ribbed-sulcate, cylindrical. Basal and lowermost cauline
leaves gradually narrowed in petiole, broadened near base, bearing woolly
axillary buds, oblong, with sparse lobules or teeth in lower part of:lamina,
withering early; other leaves linear, convolute, usually up to midrib,
cuspidate, sometimes with solitary teeth in lower part, sessile, distinctly
short-decurrent, green, glabrous or usually with isolated arachnoid hairs,
rugose and punctate-alveolate above, somewhat arachnoid-hairy: beneath,
with numerous sessile glandular hairs, obliquely rising, slightly convolute,
reduced upward, absent below capitula. Capitula solitary. Involucre 20—
25 mm long, stalked; outer and partly middle involucral bracts oblong-
ovate or oblong, green, with dark green, somewhat thick and slightly

632

arachnoid-hairy tip, recurved, with, scarcely visible brownish cusp, other
bracts appressed; middle bracts ovate, attenuate with short brown cusp,
pale green, with prominent midrib and two less prominent lateral veins,
covered with scattered sessile glandular hairs, margin membranous,
ciliate; inner bracts oblong-linear, prominent, acuminate, broadly
membranous, ciliolate along margin in upper part, with short crisped
whitish hairs on outer side and hairy tip. Receptacle alveolate, alveoli
surrounded by basally connate scales. Corolla pink-purple, about 20 mm
long, gradually narrowed over 7-8 mm, with 3.5—4 mm long lobes of
limb; apical appendages of anthers 3—3.5 mm long, oblong, gradually
attenuate above, incurved, basal appendages about 3 mm long, setaceous,
divided; style glabrous; stigma 2.5-3 mm long, divided up to half into
two recurved, papillate branches, pubescent at base; filaments verrucose.
Pappus immature, persistent, plumose, of unequal bristles, of which two
very conspicuous; achenes quite immature, densely spinulose in upper
part, coronate, with pappophorus at apex, enclosing nectary. Flowering
VITI-IxX.

Rocks.—Soviet Central Asia: Dzhungaria-Tarbagatai (Dzhungarian
Alatau). Endemic. Described from the Dyuvantau Mountains on the
southern slope of the Dzhungarian Alatau. Type in Alma-Ata; cotype in
Leningrad.

88. J. serratuloides [Ijin in Tr. Turk. Nauchn. Obshch. 2 (1925)

2h
Perennial. Plants 25-40 cm high, with thick root and short caudices
covered by scales of old leaves and woolly pubescence. Stems few,
simple or less often with one or two branches, glabrous, green, many-
leaved, leafless only below very tip. Leaves oblong or oblong-lanceolate,
pinnatisect or pinnatipartite, with oblong-linear, entire or often weakly
640 toothed, revolute, green and glabrous on both sides, slightly scabrous
above, with sessile glandular hairs beneath; basal and lowermost cauline
leaves petiolate, somewhat broadened at base; cauline leaves sessile and
short-decurrent, uppermost reduced, auriculate. Capitula solitary.
Involucre narrowly cupuliform, 15-20 mm long and 10-15 mm wide,
glabrous, yellowish-green, often with purple tinge above; involucral bracts
lanceolate, gradually narrowed, especially in middle, with finely scabrous-
serrate and slightly deflexed, 3-5 mm long cusp; innermost bracts
gradually attenuate and with very short bristles. Receptacle covered with
short, yellowish, scales, one-fifth to one-sixth as long as involucre and
connate into membranes. Corolla pink-purple, about 20 mm long,
narrowed over 8-9 mm, with 4-5 mm long lobes of limb, glandular.
Pappus persistent, 7-10 mm long, pappus bristles plumose; achenes
oblong, usually 6-7 mm long, smooth, somewhat flat to four-angled,

633

Plate XLVII.
Jurinea xeranthemoides Ijin; 2—J. ferganica Ijin;

639

1

3—J. kuraminensis l\jin.

64

—

634

light brown, toothed above along edges, with two-tiered pappophorus at
tip; lower tier patellate, occupying almost entire area, with cupuliform
and five-toothed nectary in the center of upper tier. Flowering VII—VIII.

On montane steppe slopes—Soviet Central Asia: Balkhash Region
(Zaisan). Endemic. Described from the Saur Mountains near the
confluence of the Oi-Karachai River with the Aba River. Type in
Leningrad.

Note. Known only from one location.

89. J. atripurpurea Winkl. ex Iljin in Tr. Turk. Nauchn. Obshch.
2 (1925) 26.—J. arachnoidea var. atropurpurea Winkl. ex Iljin ibid
(1925).

Perennial. Plants up to 80 cm high; tap root strong, woody fibrous,
with cord-like strong adventitous roots. Stems erect, cylindrical, finely
ribbed-sulcate and arachnoid-hairy, flocculose-arachnoid-hairy in upper
half, with golden, scattered sessile glandular hairs, simple or with isolated
long branches, in upper half always unbranched and leafless. Leaves
pinnatisect or pinnatipartite, less often almost entire or irregularly toothed,
usually with equally broad and subobtuse lobes, terminal lobe often
strongly attenuate, lateral lobes remote; basal (up to 25 cm long) and
lower cauline leaves gradually narrowed into narrow-winged petiole,
with woolly axillary buds; other cauline leaves sessile, often somewhat
decurrent, uppermost linear or filiform; all leaves greenish or dull green
and weakly rugose above, glabrous, somewhat hispidulous, or finely
arachnoid-hairy, grayish beneath due to more or less arachnoid-hairy

pubescence, with light-colored sessile glandular hairs and prominent

yellowish veins. Capitula with many florets, solitary. Involucre
cupuliform, 15—20 mm long; outer involucral bracts somewhat thick
in upper half, herbaceous, short-acuminate, finely arachnoid-hairy or
floccose-arachnoid-hairy, sharply uncately decurved; middle bracts
appressed, purple with prominent veins, and attenuate, arachnoid-hairy,
recurved tip; inner bracts appressed, acuminate, straight, dark purple,
not arachnoid-hairy, ciliolate, densely covered with scarcely visible small
bristles above, sometimes dotlike in size. Receptacle squamose; scales
smooth. Corolla almost blackish-purple, subglabrous or with sessile
scattered glandular hairs, 17-20 mm long, narrowed over 7-9 mm with
3.5-5 mm long lobes of limb; apical appendages of anthers purple,
subacute, 2.5-3 mm long; basal appendages 2-3 mm long; filaments
glabrous; stigma 3.5—4 mm long, with short papillae, and short spreading
basal bristles; style glabrous or with bristles only at extreme tip. Pappus
7-8 mm long, snow-white, later turning yellowish, of unequal many-
rowed bristles, of which two much longer than others, short-barbate,
with barbules less often equal, and usually two times as long as thickness

642

635

of bristles, all bristles somewhat connate into, broad ring with narrow
opening, easily detaching from achene at maturity; achenes 5—5.5 mm
long, yellowish brown, four-angled, oblong-obpyramidal, often slightly
deformed, with apical corona of brownish hairs, with girdle of colorless
hairs in upper half, later turning brown, with whitish, smooth, conical
pappophorus at tip, with short stalk, terminating in cyathiform, dark,
somewhat five-toothed nectary with membranous border, enclosed by
ring. Flowering III—V; fruiting 1V—second half of VI. (Plate XLIX,
Fig. 1.)

Varicolored rocks in lower hills of southern Tadzhikistan, up to
1,000 m; zone of pistachio and bean caper [Zygophyllum] woodlands.—
Soviet Central Asia: Pamiro-Alai (Babatag, Khodzha-Kazian,
Gardaniushtli Range, Gazi-Mailik and others). Endemic. Confirmed from
the vicinity of Iomid in the Gazi-Mailik Mountains between the Vakhsh
and the Kafirnigan rivers. In view of the poor preservation of the original
specimens and lack of a description, | would propose to consider the
plants.of Varivtseva and Nepli from the Tardani-Ushtli Range, No. 501,
collected in 1947, as the type specimens.

90. J. sangardensis [Ijin in Tr. Turk. Nauchn. Obshch. 2 (1925)
25.—Ie.: Ibid. VII.

Perennial. Plants about 50 cm high. Stem simple, finely’ sulcate-
striate, flocculose-arachnoid-hairy, with sessile glandular hairs. Basal and
lower cauline leaves petiolate, oblong-oval, pinnate, with linearly
attenuate, entire, subacute segments; middle cauline leaves sessile,
decurrent with row of fimbriate appendages; uppermost leaves reduced,
all leaves green and slightly scabrous above, finely arachnoid-hairy and
grayish beneath. Capitula large, solitary. Involucre up to 20 mm long,
with appressed bracts having prominent veins, glabrous or subglabrous;
middle and inner bracts gradually acuminate and with scarcely visible
appressed bristles on outer side. Pappus dirty-yellowish, persistent, pappus
bristles densely barbate; achenes glabrous, smooth. Yellowish, with few
longitudinal folds, without teeth, with somewhat hard, toothed corona
above along edges, about 6 mm long, corona enclosing fleshy, later
hardening, thick, broad cupuliform pappogenus, outer surface ivory-
colored, inner beet red, with nectary at base in center. Fruiting V—VI.

Foothills.—Soviet Central Asia: Pamiro-Alai (Denau). Endemic.
Described from Denau, Sangardak District. Type in Leningrad.

Series 4. Baissunenses Iljin——Leaves not decurrent.

91. J. baissunensis [ljin in Tr. Turk. Nauchn. Obshch. 2 (1925)
26.—Ie.: Ibid. IX.

643

636

Perennial. Plants 30-75 cm high; tap root, densely fibrous. Stem
solitary or few, somewhat thick, fistular or in upper part filled with
spongy tissue, finely barbate, cylindrical, usually simple, less often with
solitary long branches, densely covered with flocculose-arachnoid-hairy
erasable pubescence and fine glandular hairs throughout, leafy only in
lower half, with remains of petioles of old leaves at base and here
flocculose-woolly-tomentose. Basal leaves numerous, narrowed into
relatively long, somewhat flat petiole, with broad oblong lamina,
pinnatisect in linear or oblong subobtuse lobes, in lower part with few
subobtuse teeth or less often with lobes, gradually reduced toward petiole;
leaves narrowed above in oblong attenuate obtuse tip; finely rugose,
subglabrous, often with few depressed: veins, and greenish above, grayish
beneath due to arachnoid-hairy coating and with scattered sessile
glandular hairs; cauline leaves reduced upward, initially short-petiolate,
later sessile, isolated leaves very short above, narrow-linear. Capitula
solitary, large. Involucre cupuliform, 20-30 mm wide and about 20-30
mm long, comprising numerous bracts; outer and middle bracts densely
flocculose arachnoid-hairy, somewhat thick and uncately recurved in
upper half; inner bracts erect, subglabrous or with small appressed
bristles, with five to seven prominent veins (especially midrib), finely
acute, finely ciliate-toothed. Receptacle densely covered with smooth
membranous, acuminate scales, almost up to half as long as involucre
or slightly shorter. Corolla purple, with scattered glandular hairs, 24—28
mm long, narrowed over 12-15 mm, with 6—7 mm long middle broader
part and 5—7 mm long limb, lobes of limb shortly and uncately incurved;
apical appendages of anthers about 3 mm long, basal 2 mm long,
contorted at tip. Pappus about 10 mm long, colorless or somewhat
yellowish, comprising numerous unequal, serrate-barbate bristles connate
into ring, inner bristles much longer than others, with cupuliform dome
in center, punctured at base through which pappophorus is exserted;
mature achenes oblong-obpyramidal or obovoid, more or less four-angled,
toothed along margin, with circular row of teeth below, whitish, smooth,
glabrous, 5—6(7) mm long, with conical pappophorus at apex, terminating
in nectary. Flowering V; fruiting VI—VIII.

On red grounds of foothills or montane steppes.—Soviet Central
Asia: Pamiro-Alai (Kugitang, between Kugitang, Baisun and Babatag
ranges). Endemic. Described from Baisun. Type in Leningrad.

Section 10. Derderia (Jaub. and Spach) Boiss. Fl. or. III (1875)
578.—Derderia Jaub. and Spach Ill. Pl. or I (1842-1843) 129.—
Cordifoliae DC. Prodr. VI (1837) 674.—Platycephalae Benth. in Benth.
and Hook. Gen. Pl. II, 1 (1873) 474 p. p.; Pflanzenfam. IV, 5 (1873)
321 p. p.—Corolla long, gradually narrowed towards base. Pappus

644

637

20-35 mm long, innermost pappus bristles with dense brush or barbules,
all bristles plumose, pappus persistent or not entirely persistent (bristles
somewhat connate into ring, detaching on extraction). Pappogenus always
developed, pappophorus present only in some Northwest Asian species
(with deciduous pappus), pappogenus and pappophorus developing
simultaneously. Achenes always oblong, glabrous, with- more or less
sessile glandular hairs, smooth, with light longitudinal striation, folds,
or streaked. Cauline leaves sessile, semiamplexicaul and auriculate, or
only auriculate (exception J. auscheriana DC. and partly J. karabugasica
Iljin). Tall, branched plants, leafy up to capitula.

This section mainly grows in Southwest Asia (J. multicaulis DC.,
J. cataonica Boiss. and Hausskn., J. macrocephala DC. etc.); five species
found in the USSR.

1. All leaves undivided and entire, oblong-oval and oval; basal and
lower cauline leaves petiolate, other leaves sessile, but not
auriculate and not decurrent (southern Caucasia) .............:::e
Pe ee cee ee Le mee tre 92. J. auscheriana DC.

tte entral, Asian. plantsjas. carted. wchaseta: olioane. dupetbeas. dane, tt ps!

2. Leaves without auricles; cauline leaves, especially upper, pendant,
pinnatisect or sinuate-toothed (KarabugaZ) ...............csceseeseeeeseeees
ee ee ee ee ee ee 90. J. karabugasica Iljin

+. Leaves alWays iAUnICUlAte, ancl gpetvoseel . sasemmnsees .. aceepene dette . tins gi
3. Involucre glabrous (Kara-Kum)......... 94. J. transhyrcanica Iljin
+ Involucre always arachnoid-hairy ...............:::cssccceesceeeseeeeeseeeeeeees 4.
4. Tall, strongly branched plants, grayish-green due to crisped

arachnoid-hairy pubescence, sometimes almost glabrous; involucre
turbinate, with appressed bracts, outer bracts always, middle bracts
sometimes, somewhat recurved (Kara-Kums) ...........::c:ecceseeeeeeeee
ot emotes mate tebe A. arty 93. J. derderioides Wink.
+ Plants white-tomentose, not more than 25—30 cm high. Involucre
large, cupuliform; bracts spinose, recurved, flocculose, except
innermost row, which is less recurved and uncate (Karatau) .......
ee rn een! Meee. 95. J. eximia Tek.

Series 1. Oblongifoliae [ljin—Leaves undivided without auricles.

92. J. auscheriana DC. Prodr. VI (1837) 674; Boiss. Fl. or. II],
578; Grossh. Fl. Kavk. IV, 171; Sosn. Obz. Kavk. Predst. (1926) 198.—
J. oblongifolia Boiss. and Hausskn. in Boiss. 1.c—Derderia cheirifolia
Jaub. and Spach. Ill. pl. or. (1844-1846) tab. 180.—Ic.: Jaub. and Spach
Ibid.

Perennial. Plants 25-50 cm high; roots bearing short caudex. Stems
solitary or few, strong, erect, ribbed-sulcate, densely covered with crisped

638

hairs and weakly arachnoid-hairy, with sessile scattered glandular hairs,
poorly branched; branches upward-spreading. All leaves undivided and
entire; basal leaves gradually narrowed into petiole; petiole withering
early; lowermost cauline leaves short-petiolate, others sessile, narrowed
toward base and upward, acute or short-cuspidate, somewhat thick and
hard, revolute, upper surface very scabrous due to cartilaginous papillae
and finely rugose above, weakly arachnoid-hairy-glabrescent, with
scattered glandular hairs, grayish beneath due to arachnoid-hairy
pubescence and densely glandular due to sessile glandular hairs, with
prominent midrib. Capitula large, solitary, with many florets. Involucre
cupuliform, 25—30 mm long, with somewhat divergent outer and middle
bracts; all bracts linear-subulate, with three prominent veins, prominent
midrib and less prominent lateral veins, ciliate, densely covered with
crisped hairs and scattered sessile glandular hairs; outer bracts spinescent;
inner bracts usually purple, narrowly lanceolate, finely aristate.
Receptacles densely squamose; scales yellowish, smooth, acuminate,
usually one-third as long as involucre. Corolla pink-purple, 28-35 mm
long, gradually narrowed over 10-15 mm, with 6-8 mm long lobes of
limb and scattered sessile glandular hairs; apical appendages of anthers
4.5-5 mm long, gradually attenuate in deltoid tip; basal appendages
3.5—4 mm long, membranous, flexuous, setaceous toward end, somewhat
divided below; stigma 2.5—3 mm long. Pappus deciduous, dirty yellow,

_ 23-24 mm long, pappus bristles plumose, unequal, but without two to

645

four prominent bristles, with plumes six to seven times as long as
thickness of bristles, all bristles weakly connate into ring, with scaly
base, on short conical structure, terminating in nectary, enclosed in
scaly circle; achenes glabrous, oblong, with unequal-toothed corona at
apex, pappophorus represented by scarcely visible smooth elevation, and
alveolus in center. Flowering VI-—VII; fruiting VII—VIII.

On desert slopes.—Caucasus: Southern Transcaucasia. General
distribution: Asia Minor, Iran. Described from Iran on the basis of

- Aucher’s collections, No. 3204. Cotype in Leningrad.

Series 2. Auriculatae Iljin—Leaves sinuate-toothed, auriculate.

93. J. derderioides Winkl. in Tr. Peterb. Bot. Sada, IX, 2 (1886)
524; O. and B. Fedtsch. Perech. Rast. Turk. IV, 303; Iljin in Tr. Turk.
Nauchn. Obshch. 2, 17; Fl. Turkm. VII, 234.

Perennial. Plants 30-60 cm high, creamish, entirely covered with
crisped arachnoid hairs. Stem branched from base, modified into long and
slender rhizomes. Leaves numerous, right up to capitula, weakly rugose-
foveate due to sunken glandular hairs, sessile and semiamplexicaul,
auriculate, ovate, oblong-ovate or almost round, sparsely sinuate-toothed,

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639

often with long teeth, usually withering before anthesis. Capitula almost
cupuliform, solitary. Involucre grayish due to fine tomentum, 20-25 mm
long and 13-18 mm wide; outer involucral bracts oblong-ovate, acute,
slightly recurved, others oblong-lanceolate to lanceolate, erect, appressed.
Receptacle pubescent, bristles very much shorter than involucre. Corolla
light pink, often yellowish on drying, glandular-hairy, 27-30 mm long,
gradually narrowed over 12-16 mm, with 3.54.5 mm long lobes of limb.
Pappus persistent, 13—-15(20) mm long, dirty yellowish, pappus bristles
barbate, plumose, with more or less distant barbules; achenes oblong,
yellowish, 6.5—8 mm long and 2-3 mm thick, smooth, longitudinally striate
with sessile glandular hairs and frequent long teeth at apex, with smooth
and conical pappophorus and nectary. Flowering V—VI; fruiting VI-VIII.

Saline sands, salt pans among sands, foot of sand hills and
hummocky sands.—Soviet Central Asia: Aralo-Caspian Region, Kyzyl-
Kum (western); Kara-Kum. Endemic. Described from the Mary Region.
Type in Leningrad.

94. J. transhyrcanica I[ljin in Tr. Turk. Nauchn. Obshch. 2 (1925)
We

Plants up to 40-50 cm high. Stems strongly branched, leafy, finely
sulcate, glabrous, with numerous sessile glandular hairs. Lower leaves
oblong-lanceolate, 7-10 cm long, gradually narrowed into petiole, erect,
acuminate, with remote, triangular, cuspidate teeth, later withering and
dropping off; middle and upper leaves reduced upward, oblong (about 5
cm long) to oval (uppermost leaves 1.0—1.5 cm long); all leaves short-
acuminate, also sparsely short-toothed, especially in lower half, sessile,
with short auricles, somewhat hard, with numerous sessile glandular hairs
in depressions on both sides, leaves usually somewhat pendant or almost
horizontal, almost reaching capitula. Capitula solitary, numerous.
Involucre cupuliform, comprising numerous imbricate’ bracts, more or
less appressed, only very tips of outer bracts slightly recurved and green,

like other bracts in remaining part, yellowish, almost coriaceous,

acuminate, glabrous, only along margin very short and frequently
scabrous-ciliate, with prominent midrib on dorsal surface. Receptacle
squamose; scales frequent, short, several times shorter than involucre,
flat, smooth. Corolla longer than pappus, narrowed over 10 mm, covered
with scattered glandular hairs. Pappus persistent, bristles numerous,
unequal, not connate into ring, plumose, innermost three or four bristles
longer than others and modified into tuft at tip with still longer barbules,
more densely and compactly arranged upward; achenes about 7 mm
long, obovoid, yellowish, glabrous and smooth, ribbed, with large,
broadly cupuliform, thick, apical pappogenus, and nectary in center.
Flowering V—VI; fruiting VII-IX.

640

Known only in the Turkmenian SSR, in Kara-Kums.—Soviet Central
Asia: Kara-Kums. Only two specimens are known: 1) near Daz-Kuyu
well, 1912, No. 281, collected by D.P. Gedevanov and D.A. Dranitsin;
and 2) 4 km due southeast of Kara-Tegelek, 1953, No. 20, collected by
I. Rustamov.

95. J. eximia Tek. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
VII, 6 (1938) 123.—Ie.: Ibid. 125. .

Perennial. Tap root woody, with compact fibrous pith and detaching
layer of outer bark, bearing short caudices near soil surface, densely
covered with compact rosettes of basal leaves, with dense, white, soft
tomentum in their axils. Stems herbaceous, solid, branched, 15-45 cm
high, with longitudinal furrows, densely leafy up to capitula, covered
with dense, erasing grayish-whitish arachnoid-hairy tomentum. Basal
leaves oval-lanceolate, long-petiolate, 5—13 cm long; cauline leaves also
oval-lanceolate, sessile, auriculate, semiamplexicaul, more or less
decurrent, reduced toward apex, with erasing. grayish white pubescence
on both sides. Capitula fairly numerous. Involucre 2.5-3 cm long and
2 cm wide, globose-ovoid, imbricate-scaly, grayish-pubescent, often
with pale purple tinge; bracts linearly lanceolate, arcuately recurved,
subulately-pointed. Corolla dark pink, with sessile glandular hairs, 30—
35 mm long, narrowed over 18-20 mm, with 5—6 mm long lobes of
limb. Pappus 18-20 mm long, with plumose-barbate bristles, connate
into ring at base, barbules remote, longer than thickness of bristles;
three to five inner bristles with dense barbules above; immature achenes
about 3 mm long, glabrous, dark brown, longitudinally barbate. Flowering
VII; fruiting VIII.

Stony habitats in mountains.—Soviet Central Asia: Tien Shan
(Karatau). Endemic. Described from Kara-Dzhatyk area not far from
the village of Uch-Uzen. Type in Leningrad.

Series 3. Pendulifoliae Iljin——Leaves not auriculate; cauline leaves
pendent.

96. J. karabugasica Iljin in Bot. Mat. Gerb. Bot. Inst..Akad. Nauk
SSSR, VII, 1 (1937) 54; Fl. Turk. VII, 232.—Ie.: op. cit. (1937) Fig. 2.

Perennial. Plants 40-50 cm high, grayish due to flocculose-arachnoid-
hairy pubescence. Stems numerous, flocculose-woolly at base, branched
almost up to middle, branches elongate, leafy up to capitula. Basal leaves
petiolate, gradually broadening toward base, withering before anthesis,
only their petiolar remains persisting near stem base, pinnatisect, with
distant short segments; all cauline leaves sessile, concolorous, strongly
broadened at base, pinnatisect, with short lanceolate or linear segments,

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641

acuminate, remote; leaves often runcinate, sinuate-toothed, with few
usually recurved lobes. Capitula solitary, initially with cylindrical, later
campanulate, involucre; involucre 20-30 mm long and 10-13 mm wide,
gray due to flocculose-arachnoid-hairy pubescence, with appressed bracts;
outermost bracts slightly recurved; inner bracts gradually pointed,
yellowish; outer and middle bracts slightly carinate, short- and dark-
cuspidate, oblong-ovate to linear. Receptacle squamose; scales very much
shorter than involucre. Corolla pale pink, 28-33 mm long, gradually
narrowed towards base, with about 4 mm long lobes of limb. Pappus
persistent, about 25 mm long, pappus hairs plumose; achenes 5-6 mm
long, glabrous with equidentate corona enclosing thick, yellowish, deeply
lobate pappophorus, with a large, high, poculiform, dark yellow nectary
in center.

Stony habitats ——Soviet Central Asia: Kara-Kum (western). Endemic.
Described from Karabugazgol. Type in Leningrad.

Section 11. Stechmannia Boiss. Fl. or. III (1875) 569 (s. str.) —
Stechmannia DC. Prodr. VI (1837) 553.—Stenocephala Benth. in Benth.
and Hook. Gen. pl. II, 1 (1873) 473 p. p.; Pflanzenfam. IV, 5 (1897)
321 p. p—Plants with woody branches, caespitose, with unilaterally
decurrent entire leaves, involucral bracts always spinose; pappus
deciduous, pappus bristles always plumose, connate into ring with narrow
opening, broad scaly base, with pappogenus enciosing nectary; achenes
oblong, glabrous, smooth, with dome shaped, five-angled or lobate apical
pappophorus. Plants mainly from Southwest Asia; so far four species
have been recorded in the USSR, all in Kopetdag.

Note. The section is described on the basis of Kopetdag species and
some Southwest Asian species, which belong in this section in its narrow
sense; these are J/. stachelinae DC., J. dumulosa Boiss., J. ramosissima
DC., J. eriobasis DC.

Type of section: J. pungens Boiss.

—

. Entire plant uniformly grayish arachnoid-haity ..................:006 Ze
+ Plants glabrous, only sometimes with densely tomentose, decurrent,
EWEN lL Tibet teint el deci ace ell al Ra ee 3.
2. Stem strongly bent at nodes; involucral spines strongly divaricate
andswith’ thickened (DASE ...2.6.-..-f...-.08 +. 97. J. antonovii Winkl.
+ Stem erect, slightly bent at nodes, spines of involucral bracts thin
-aciioneatt lc bret aectc a as ea ere nila eee i te 98. J. kultiassovii: [jin
3. Dense woolly globules developed in leaf axils, decurrent on stem
as wings; spines of involucral bracts strongly divergent (Malye
HIFCSSCM | ESAIKMANG) i c.cheo tet ct rre cece sa cetenscccartt a: 99. J. lydiae Ijin
+ Woolly globules not present in leaf axils; involucral bracts modifed
as divergent spines (central Kopetdag) ..... 100. J. ludmilae [jin

649

642

97. J. antonovii Winkl. in Tr. Peterb. Bot. Sada, XI, 1 (1890) 145;
O. and B. Fedtsch. Perech. Rast. Turk. IV, 295; [jin in Tr. Turk. Nauchn.
Obshch. 2, 14; Fl. Turkm. VII, 228.—Ic.: Winkl. op. cit. t. II, f. 4.—
Exs.: P. Sintensis. Iter transcasp.-persic. 1900-1901, No. 1947.

Perennial. Semishrub, 8-25 cm high, divaricately branched from
base. Root strong, often branched, strongly fibrous. Stems densely
covered with remains of hard, relatively long, sharply divergent petioles
of fallen leaves, broader at base, and woolly in axils, arachnoid-hairy
pubescent, more or less branched, nodose, usually appearing three-angled
due to decurrent petioles. Leaves oblong-linear, undivided, with distant,
sparse, subobtuse, small teeth or margin almost entire, involute, with
very short cartilaginous tip; lower and middle leaves 4-10 cm long and
2-8 mm wide, gradually narrowed in more or less long petiole, carinate
below, with broadened base; upper leaves shorter, narrower, sessile.
Capitula solitary, often aggregated, with 15-25 florets, five to seven
florets only in less developed forms. Involucre cylindrical, 20-30 mm
long and 5-7 mm wide (including 15—25 mm long divaricate bracts),
grayish-green due to fine pubescence; outer and middle involucral bracts
attenuate from relatively broad base into divaricate, long, narrow, stiff
structures, terminating into brownish, almost glabrous spine; inner bracts
narrow oblong-linear, flat, gradually acuminate, membranous, with
scattered sessile glandular hairs and fine pubescence on outer side, dirty
purple in upper part. Receptacle squamose; scales short, glabrous and
smooth, acuminate, usually 3—5 mm long. Corolla prominent, pink, very
densely covered with sessile glandular hairs, especially in upper part,

25-35 mm long, gradually narrowed at 10-12 mm, with 4-7 mm long

lobes of limb; apical appendages of anthers 6-8 mm long, whitish-
pinkish, gradually acuminate, very deeply divided above, basal
appendages setaceous, colorless, 4-6 mm long; slightly divided only at
tip. Pappus deciduous, 15—22 mm long, ferruginous either entirely or
only in lower part, comprising long-barbate unequal bristles, with barbules
3—4 times as long as thickness of bristles, almost as long, or even shorter,
connate into ring with narrowed base; achenes oblong, 5—6 mm long,
glabrous, smooth, with small and many-toothed corona enclosing
hemispherical pappophorus with.membranous, five-lobed, stalked, nectary
in center, protruding through opening in pappus. Flowering V—VII;
fruiting VII—X.

Clayey and gravelly mountain slopes, in steppe and semisteppe
zone.—Soviet Central Asia: mountainous Turkmenia. Endemic. Described
from the vicinity of Germab in Kopetdag. Type in Leningrad.

Note. This species varies in plant size, length of corolla, pappus
and especially its bristles, which are mostly barbate, with the barbules
three to four times as long as the thickness of the bristle, straight and

650

643

obliquely directed upward relative to the bristles, which are sometimes
more scattered, of almost the same length, but arcuately recurved, and
less often the serrate-barbate bristles are as long as or shorter than the
thickness of the bristles. The specimens collected by M.V. Androsov
and L.M. Bubyr from the Kurtsu Region near Ashkhabad on June 27,
1912, No.- 2431, are particularly noteworthy. They have entire leaves
that are more appressed to the stem and serrate pappus bristles (f.
serratisetosa Ijin f. n.).

98. J. kultiassovii Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XX (1960) 352; Fl. Turkm. VII, 226, Plate 36.

Perennial. Plants 25-30 cm high, with many-headed caudex, and
strongly fibrous roots; many-stemmed, grayish. Stems more or less erect,
simple or only with isolated short branches, angular, gradually and
weakly bent at nodes arachnoid hairy-flocculose-pubescent, dirty purple
in lower part, leafy up to tip, densely squamose with stiff scales, the
remains of old petioles near base. Cauline leaves numerous, obliquely
directed upward, narrowly lanceolate or narrowly oblong, undivided and
entire, acuminate, one-veined, narrowly involute, narrowed toward base,
grayish on both sides due to sparse arachnoid-hairy pubescence and with
sessile glandular hairs, often poorly visible due to pubescence, 4-8 cm
long, 3-6 mm wide; lower leaves distinctly petiolate; middle and upper
leaves sessile, not enclosing capitulum. Capitula solitary, about 4—5 cm
long (including florets). Involucre narrowly cylindrical, about 2.75—3
cm long and 1 cm wide, grayish due to arachnoid-hairy pubescence,
with numerous imbricate bracts, terminating into glabrous, rusty-brown
awns, shorter in outer bracts, similar to large number of middle bracts,
gradually attenuate outward, with straight awns, from outer to inner
bracts, elongated up to 4—5 mm. Receptacle squamose; scales somewhat
stiff, very wide, 2.5-3 mm long, irregularly divided, sometimes up to
middle or even below. Florets significantly longer than involucre, 12—
20 in capitulum. Corolla pink, 30-35 mm long, gradually narrowed in
middle, with 5—6 mm long lobes of limb, exserted, whitish; apical
appendages of anthers scatteredly pubescent, similar to basal appendages,
5—6 mm long. Pappus entirely deciduous, 12-15 mm long, pinkish-rusty,
bristles barbate-plumose, with barbules four to five times as long
as thickness of bristles, connate into ring, forming continuous
membranous base, bearing small nectary; achene immature, four-angled,
glabrous, smooth, with short-columnar, five-angled pappophorus.
Flowering VI.

On mountain screes.——Soviet Central Asia: mountainous Turkmenia.
Endemic. Described from the vicinity of the village of Sharlak. Type in
Ashkhabad.

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644

99. J. lydiae [ljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XIX (1959) 390, Fig. 1; Fl. Turkm. VII, 229.

Perennial. Light green semishrub, (6)8—-15 cm high, floccose with
long fibrous woody root, forming branched caudex. Stems numerous,
simple, branched above, angular, glabrous or often with dense and soft
flocculose angles, especially in lower part. Leaves numerous, oblong,
subobtuse, glabrous on both sides, with prominent midrib beneath,
decurrent and gradually narrowed into three-angled petiole; lower leaves
falling before anthesis, middle (6)8—10 cm long, upper reduced and
subtending capitulum. Capitula numerous, 4—4.5 cm long. Involucre
cylindrical at anthesis, 22-30 mm long and 18—23 mm wide, 7-10 mm
wide without divaricate involucral spines; involucral bracts numerous,
imbricate, outer and middle bracts gradually narrowed, attenuate trom
broad yellow base, somewhat thick, greenish, glabrous on outer side,
puberulent on inner side, terminating in smooth rusty spine, distinctly
arcuately recurved; innermost bracts erect, narrow, linear, gradually
acuminate, purple above, glabrous on both sides, often ciliolate only
along border. Receptacle squamose; scales smooth, acuminate, one-eighth
to one-seventh as long as involucre. Corolla pink, densely glandular,
usually 32-35 mm long, with 13-16 mm long tube and 6—7 mm long
lobes of limb. Pappus deciduous, all pappus bristles plumose, connate
into ring, extended in scaly base with five-partite globose pappogenus
and whitish scaly nectary falling entirely with them; achenes oblong,
glabrous, smooth, immature achenes about 5 mm long, with toothed
corona above along edges, and with smooth dome-shaped pappophorus
inside. Flowering VI-—VII; fruiting VII.

Wormwood and saltwort waste lands.—Soviet Central Asia:
mountainous Turkmenia. Endemic. Described from Lesser Balkhans.
Type in Leningrad.

100. J. ludmilae Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XIX (1959) 392, Fig. 2; Fl. Turkm. VII, 226.

Semishrub with long, fibrous, tap root, connecting rhizome covered
with old petioles. Stems of current year numerous, 5—8 cm long,
hemispherical, unbranched or less often with solitary branches in lower
part, green, ribbed, like entire plant almost glabrous, with slightly
arachnoid hairy-coating at base. Leaves linear, entire or with few teeth,
strongly convolute, subobtuse, with prominent midrib beneath, usually
30-50 mm long, slightly broader at base, arcuately recurved, without
distinct tuft of hairs in axil. Capitula numerous. Involucre about 20 mm
long, infundibuliform due to strongly divergent imbricate, linear, ‘erect,
unequal bracts, somewhat broadened from base itself; attenuate into
somewhat stiff, thin, dirty purple spine. Florets in capitulum about 20;

645

Wii
WwW

Py G

\ (2
VIZENGNA

INS
SENN

SSAA

SS

653 Plate XLVIII.
1—Jurinea tapetodes \\jin; 2—J. apoda lljin.

652

646

corolla pink, with scattered sessile glandular hairs, usually 27-30 mm
long, gradually narrowed toward base, with 10-12 mm long narrow basal
part, lobes of limb 5—6 mm long. Pappus ferruginous, 15—20 mm long;
bristles barbate-plumose, deciduous, -all bristles connate into ring with
extended scaly base bearing cyathiform, longitudinally five-fid pappogen,
with columnar whitish nectary at apex; achenes glabrous, immature,
longitudinally striate. Flowering VI-—VII; fruiting VII.

Gravelly mountain screes.—Soviet Central Asia: mountainous
Turkmenia. Endemic. Described from Chuli. Type in Leningrad.

Section 12. Nanae Iljin sect. n. in Addenda XXVI, 725.—
Semishrubs, usually short, stemless or with short stem, mountainous or
alpine plants, with undivided or finely divided leaves, predominantly
rosulate; pappus serrate and barbate, always shedding easily; pappogenus
like pappophorus always developed; achenes glabrous, often with smooth
ridges.

Type of section: J. abramovii Rgl. and Herd.

1. Plants stemless, forming dense turf; capitula always sessile..... 2.
+). Stemsi- somewhatideveloped:... 53 ee sca stves eesti eceeeteeseeas 3.
2. Leaves pinnatipartite, with short lobes, gray- -arachnoid-hairy on both
sides; involucral bracts appressed, only sometimes their tips slightly
recurved; pappus barbate; achenes with scarcely visible, smooth
longitudinal ridges (Kugitang) .................. 102. J. tapetodes Iljin
+ Leaves undivided and entire, short-petiolate, oval or oblong-oval,
greenish above, white tomentose beneath; involucral bracts
arcuately recurved; pappus bristles serrate; achenes glabrous, smooth
(Ropetdag) tse oP siocd cache ea ee 104. J. antoninae I\}jin
3. Plants with pinnatisect leaves, with thin, gray-arachnoid-hairy,
sometimes linear segments; stems with solitary leaves and
pendulous capitula; involucral bracts appressed or slightly recurved
at tip; pappus bristles barbate; achenes glabrous with longitudinal
smooth ridges (mountainous ZETAVSMAN))! MewidescsSusseleereh vain ee
oT Ce er. culien Seale 101. J. abramovii Rgl. and Herd.
+ Leaves undivided and entire, less often with solitary teeth...... 4.
4. Stems almost leafless, sometimes reaching 25 cm in height; leaves
lanceolate or oblong, or with solitary teeth, grayish-green above,
white-tomentose beneath; outer and middle involucral bracts
recurved, pappus bristles serrate; achenes small, with longitudinal
smooth striations (southwestern Kopetdag) ..............ccceeeeeseeseeees
cibbasted foeaditcayer: Maan eg onteaielnn Biome aneel 103. J. bobrovii [lyin
+ Leaves linear, hard, entire, green on both sides; involucral bracts
almost equal, somewhat lax, finely acuminate; pappus bristles

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647

barbate; achenes larger, about 4 mm long, with smooth longitudinal
ridges, and isolated teeth above ........ 105. J. knorringiana [ljin

101. J. abramovii Rg!. and Herd. in Izv. Obshch. Lyubit Estestv.
Antroip. 1 Etn. XXXIV, 2 (1882) 53; O. and B. Fedtsch. Perech. Rast.
Turk. IV, 303; [jin in Tr. Turk. Nauchn. Obshch. 2, 27.

Perennial. Plants short-stemmed, 3-20 cm high, with dark brown
root branched above, bearing numerous, more or less elongate caudices,
densely covered with scaly, yellowish-brown remains of old leaves,
sparsely arachnoid-hairy-tomentose, often almost glabrescent. Stems and
leaves grayish-green or whitish due to arachnoid-hairy pubescence, few
with dense sessile glandular hairs. Leaves basal and cauline; cauline
leaves sparse, usually absent in upper part; leaves oblong or even linear,
pinnatisect into narrow linear or oblong, obtuse segments, sometimes
almost pinnatilobate, with more or less short lobes, involute almost up
to midrib. Capitula solitary, somewhat pendulous. Involucre cupuliform,
about 1.5—2 cm long, with few rows of imbricate, lax bracts, usually
dirty-purple, arachnoid-hairy pubescent in lower part, flat, with indistinct
veins and sessile glandular hairs; outer bracts shorter, oblong-ovate,
usually with somewhat recurved apex, subacute, others oblong-lanceolate,
acuminate, in lower part with sparse and irregular small teeth, like inner
bracts aristate, somewhat hairy in upper part, usually along margin; upper
bracts scabrous. Receptacle with very short, about 1-2 mm long, smooth
scales. Corolla pink or pink-purple, with numerous sessile glandular hairs,
18-25 mm long, narrowed over 7-12 mm, with 6.5—9 mm long tube
and 4—5 mm long limb, bent in peripheral florets, as a result corolla
slightly pendulous, corolla of central florets erect; apical appendages of
anthers whitish, about 3 mm long, basal appendages colorless, 2-3 mm
long, divided up to base. Pappus deciduous, 12-20 mm long, snow-
white, bristles barbate, with barbules two to four times as long as
thickness of bristles; bristles connate into ring, with extended scaly base,
bearing cyathiform scaly pappogenus with white, membranous, sinuate
nectary; achenes up to 7 mm long, oblong, four-angled, smooth, grayish-
brown, with longitudinal ridges; with membranous and sinuate corona
along edges, enclosing whitish conical pappophorus. Flowering VII—VIII;
fruiting VIII-IX. (Plate L, Fig. 2.)

Stony slopes, screes, and lake banks.—Soviet Central Asia: Pamiro-
Alai (western). Endemic. Described from Lake Kuli-Kalan. Type in

‘Leningrad.

102. J. tapetodes Iljin in Izv. Bot. Inst. Akad. Nauk SSSR, XXX
(1932) 344; Fl. Turkm. VII, 219.

Perennial. Plants forming dense turf with many-headed caudex, short,
2—5 cm high, stemless; caudices very densely covered with remains of

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648

old leaves. Leaves in dense rosette, usually 10-15 mm (5—25 mm) long,
on sparse, slightly elongate branches, sometimes up to 40 mm long,
sessile, with two to four pairs of short, linear to slightly lobate obtuse
lobes, grayish due to very dense arachnoid-hairy tomentum on both sides,
with scattered sunken glandular hairs, somewhat thick due to strongly
revolute margins almost up to midrib, depressed above, subtending
capitulum. Capitula sessile or on short peduncles. Involucre almost
cylindrical, slightly broadened above, 1.5—2.25 cm long and 0.7—1.0 cm
wide, with somewhat lax bracts; outer and usually middle bracts ovate
or oblong-ovate, greenish-gray due to relatively dense, very short, white
hairs, with short cusp, usually recurved, other bracts lanceolately linear;
upper bracts acuminate, like inner bracts prominent, narrower, attenuate
into long recurved cusp, dirty-purple, densely covered on outer side
with very short appressed hairs and scattered sessile glandular hairs.
Receptacle densely covered with short 1-2 mm long scales. Florets
exceeding involucre; corolla pink-purple, with sessile glandular hairs,
sometimes mixed with very short-stalked glandular hairs, 20-23 mm
long, gradually narrowed over 9-10 mm, with 4-5 mm long limb; apical
appendages of anthers whitish, with scattered hairs about 4 mm long,
basal appendages colorless, setaceous, 3-4 mm long and slightly divided
below longitudinally. Pappus shedding easily, up to 15 mm long, snow-
white, bristles barbate, barbules one and one-half to two (three) times
as long as thickness of bristles, up to four times long only in four
longer bristles, all bristles connate into ring with extended base, bearing
a pink, fleshy, lobate structure surrounding cyathiform nectary with five-
lobate yellowish membranous margin; achenes mature, 6-8 mm long,
almost smooth or with scarcely visible longitudinal helices, rugose above,
often modified into spiny tubercles along thickened edges, yellowish,
especially below corona. Corona distinctly toothed along edges and
terminal on achenes with conical whitish pappophorus. Flowering VI—
VIII; fruiting VII-VIII. (Plate XL, Figs. 13-16; Plate XLVIII, Fig. 1.)

Gravelly and stony plateau and slopes, a rocks in subalpine zone.—
Soviet Central Asia: Pamiro-Alai (Kugitang). Endemic. Described from
the vicinity of the village of Kugitang from the collections of E.G.
Bobrov. Type in Leningrad.

103. J. bobrovii [ljin sp. nov. in Addenda XXVI, 725.

Perennial. Semishrub with prostrate woody branches, bearing
numerous dense rosettes of leaves at top. Stems 15—25 cm high, striate,
flocculose-arachnoid-hairy, erect, simple, leafy in lower part. Basal and
rosette leaves lanceolate or oblong, entire, or with isolated teeth, subacute,
narrowed toward base; cauline leaves narrower, sessile; all leaves grayish
green above due to fine arachnoid-hairy tomentum, rugose and with

656

649

sessile glandular hairs in alveoli, white-tomentose beneath with prominent
midrib. Capitula solitary. Involucre cupuliform, arachnoid-hairy in lower
part; outer involucral bracts acuminate with arcuate tips; middle bracts
less recurved; inner bracts erect, long, and scaly-tipped; all bracts with
prominent veins and small teeth. Corolla 14-15 mm long, abruptly
narrowed over 4—5 mm, with 4—5 mm long lobes of limb, covered with
scattered sessile glandular hairs. Pappus entirely deciduous, bristles
connate into ring, scabrous, pappogenus on extended base in ring as
lobed cup, whitish on outer side, red inside, with pappophorus in center;
achenes 2—2.25 mm long, with smooth longitudinal striations, and equal-
toothed membranous corona, enclosing broad cupuliform pappophorus
with smooth margins.

Mountain slopes.—Soviet Central Asia: mountainous Turkmenia.
Endemic. Described from the western slopes of Khosar-Dag in Kopetdag,
from the collections of E.G. Bobrov. Type in Leningrad.

104. J. antoninae [ljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, VII,.3 (1937) .56.. Ie.: Ibid: p) 37.

Perennial, short semishrub forming dense turf. Root thick, woody,
producing numerous densely appressed woody branches, terminating with
rosette of leaves 2-4 cm across. Leaves oval or oblong-oval, spatulate,
gradually narrowed into petiole, one-veined, undivided and entire, less
often with isolated teeth, white-tomentose beneath, arachnoid-hairy above,
grayish-green, lamina 4—15 mm long and 3—6 mm wide. Stems of current
year short if developed, 0.5—3 cm high, simple, with oblong leaves,
one-headed. Involucre 10-13 mm long, cupuliform at fruiting, arachnoid-
hairy, with lanceolate bracts; inner bracts almost linear, acuminate; all
bracts appressed; outer and middle bracts somewhat recurved only at
tip. Receptacle densely squamose; scales, half as long as involucre.
Pappus deciduous, snow-white or yellowish, pappus bristles scabrous-
barbate, with barbules slightly shorter than thickness of bristles, sparse;
all bristles connate into ring with extended base, enclosing small five-
toothed nectary; achenes mature, 3-3.5 mm long, up to 1 mm thick
above, glabrous, smooth, toothed along edges, with fine pappophorus at
tip as tubercle. Fruiting VI—VII.

Stony slopes of mountains.—Soviet Central Asia: mountainous
Turkmenia. Endemic. Described from the Bakharden Region of Kopetdag
from the Tyazi-Tailanly Mountains, a short distance from the village of
Tyazi-Tailan. Type in Leningrad.

105. J. knorringiana Ijin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XXI (1961) 384.—Ie.: Ibid. Fig. 2. .

Perennial. A short semishrub, about 10 cm high. Stems few,
originating from apices of previous year’s stems, densely covered with

65

~

650

scaly remains of old leaves, erect, simple, slightly angular-ribbed, sulcate,
with scattered, somewhat stiff, short hairs, more or less densely covered
with sessile glandular hairs, weakly arachnoid-hairy at some places.
Leaves few, usually about 5 cm long, undivided and entire, sometimes
revolute up to midrib, short-acuminate; basal and rosette leaves narrowed
toward base and again broadened at the attachment; cauline leaves sessile,
broadened at base and decurrent with short and narrow, entire wing; all
leaves bright green on both sides, slightly rugose above, with sparse,
scattered, sessile glandular hairs and dense, stiff, almost cartilaginous
few-celled short hairs, often capitate, at some places with weakly
arachnoid-hairy, glabrescent pubescence, glabrous beneath, but with
numerous sessile glandular hairs and prominent, distinctly whitish midrib.
Capitula solitary terminal on stem, with fewer florets. Involucre
cupuliform, up to 20 mm long and up to 8-12 mm wide, five-rowed;
outer bracts relatively lax, linearly-lanceolate, finely acuminate; inner
bracts finely acuminate, outer flocculose-arachnoid-hairy, others short
appressed, with small teeth at some places. Receptacle densely squamose;
scales smooth, pointed, one-fourth to one-third as long as involucre.
Pappus deciduous, snow-white, 8—9 mm long (excluding bristles), short-
barbate, with barbules as long as or slightly exceeding thickness of
bristles. Achenes about 4 mm long, four-angled, obpyramidal, with
toothed corona above, and longitudinal ridges, with rows of teeth along
ridges in upper half, brownish, with smooth dome-shaped pappophorus
at tip, enclosing five-toothed nectary on long stalk exserted from opening
at pappus base.

Soviet Central Asia: Tien Shan (Karatau). Endemic. Described
from Zhan-Kurgan-Sai north of the village of Atabaevka. Type in
Leningrad.

Section 13. Molles Korsh. in Bull. Acad. Sc. Pétersb. I, 2 (1894)
123, pro stirpe.—Capitula large or medium, with many florets; florets
bisexual; pappus readily and entirely shedding, pappus bristles connate
into ring at base, short-barbate or almost scabrous, base of pappus
extended into scale, pappogenus with nectary detaching together with
pappus; besides pappogenus, conical or dome-shaped pappophorus always
present; nectary on short manubrium or sessile; achenes distinctly
scabrous due to discontinuous alveolate, parallel ribs on edges, alveoli
alternating with fine erect acute teeth and scattered numerous sessile
glandular hairs, four peripheral thicker ribs distinctly cartilaginous. Plants
woolly-arachnoid-hairy at base. A fairly large number of species of this
section are found in Europe and Northwest Asia, which are difficult to
distinguish among themselves.

Type of section: J. arachnoidea Bge.

658

65i

. Plants yellowish-green, with long and'slender branches, densely

glandular; outer and middle involucral bracts imbricate, reduced,
‘green, roundish below, with dark green apex and short cusp, arcuate
at apex; achenes entirely covered with glandular hairs masking its
markings (Oly) 117. J. demetrii [jin
Plants with different set of characters; achenes with distinct
alveolate-tuberculate and vallate structure, also always glandular
hainy;;butsnot*masking ats markings: 08. Av ROA... De

. Involucre arachnoid hairy-tomentose;-all involucral bracts appressed,

only triangular and dark green tips of reduced outer and middle
involucral bracts exserted from tomentum..............ccccceseeecees Be
Involucre different, often also arachnoid hairy-tomentose; bracts
appressed or without tomentum; triangular tips of outer and middle
bracts never exserted from tomentum, but as linear tips .......... 4.

. Leaves almost always with smooth margins, not sinuate (south-

eastern European Part and Ciscaucasita ...........ccccccccsccsssccesssceeseeeens
BR ca) TR RS SEO at i 106. J. arachnoidea Bge.
Leaves always sinuate (chalky deposits of eastern Ukraine) ........
SESE PE RENE BRO) RE BETIS 107. J. talievii Klok.

. Plants of Crimea; leaves always sinuate (exceptions rare)............

NER E ER LOR OTT RRA EEA 114. J. sordida Stev.
Leaves of different type, may be flat and sinuate, but never Crimean
polenta ch ety sie bs Deen ARR Id EARNS ANTS OE 0g >A

. Plants strong, branched from base; branches spreading upward,

strongly winged; leaves large, erect (alpine plant of the ‘Glavnyi’

[Main] Caucasus Range)......... 116. J. exuberans (Trautv.) Sosn.
Plants*wathedifferentiset! of characters 228.2202 ee Ss 6.
. Plants of Transcaucasia................. 115. J. blanda (MB.) C.A.M.
Plants of the European Part of the USSR and extreme western
KA ZAalShiStaT ES SPORE OPE NOES, RN, SO GN eka rt Je
. Leaves not sinuate (exceptions quite rare) ...........ccceeeeeee eens 8.

Leaves always sinuate. Involucre always arachnoid-hairy tomentose;
lower and middle bracts distinctly finely recurved (chalk deposits)
b EOE, IE EO WM CHENIN ke, | NE aks 11.

. Middle involucral bracts from their ovate-oblong, somewhat

swollen, pale base sharply narrowed to relatively long and narrow
apex, loosely appressed or even gradually recurved, dark green or
purple, sometimes deciduous (eastern and middle parts of the

European: territory)... 9h. 1014) 10... 110. J. ledebourii Bge.
All involucral bracts gradually acuminate, not swollen in lower
PCTS PS PR, er SL TN Sie EG oe o

. Capitula large; all involucral bracts, except one or two innermost

rows, gradually arcuately recurved, with long, gradually acuminate

652

659 bent part, not arachnoid-hairy tomentose; inner bracts peri
involute in upper part and long-ciliate along margin...............:....
<n gaieapany savages Races «eGeRE- Saye. TERMI. Sees oth 113. J. michelsonii tijin

+ Outer involucral bracts very short-acuminate or erect............. 10.
10. Entire plant densely arachnoid-hairy, almost whitish tomentose,
grayish green and arachnoid-hairy, above, white-tomentose beneath;
involucre always arachnoid hairy-tomentose, with appressed bracts;
outer and middle bracts with short-recurved apices (coastal zone
of western Ukraine) .............c:cceeeeeeeees 111. J. mollissima Klok.
+ Plants weakly arachnoid-hairy, but not arachnoid hairy-tomentose;
leaves green, often bright green, glabrous, or somewhat arachnoid-
hairy, white-tomentose beneath; involucral bracts more or less
appressed only in beginning, later lax, somewhat divergent,
sometimes not arachnoid hairy-tomentose, subglabrous or weakly
arachnoid-hairy, short-acuminate, with cusp slightly recurved
(middle part of Ukraine) ........00.0 ee 112. J. calcarea Klok.
11. Achenes with four to six discontinuous ribs, approximate on edges;
corona of achenes initially membranous, erect, bases of teeth fused
in lower half, later cartilaginous (chalk deposits along Lower Don)
saned daslnp « SERGE LENSER GE: Sats. - daaeeeSeenee REE oe 109. J. kasakorum [lyin
+ Achenes usually with three longitudinal alveolate-discontinuous ribs
on edges, not approximate, corona distinctly toothed, cartilaginous,
with few recurved teeth (chalk deposits of Don and Lower Volga)
cep aide rie sbesaeene «weet sapaachet 0eaQane. « ENS eee 108. J. cretacea Bge.

Series 1. Arachnoideae Iljin—Achenes with sharply developed
markings and scattered sessile glandular hairs not masking markings.

106. J. arachnoidea Bge. in Flora, XXIV, 1 (1841) 157; Ldb. FI.
Ross. II, 766 p. p.; Boiss. Fl. or. II, 574 p. p.; O. and B. Fedtsch.
Perech. Rast. Turk. IV, 301 p. p.; Klok. in Vizn. Rosl. URSR, 565;
Schchian in Fl. Gruzi, VIII, 447.—J.
Akad. Nauk, 1, 2 (1894) 124; Grossh. Fl. Kavk. IV, 173; Fedtsch. in
Fler. Fl. Evrop. Ross. 1005; Sosn. in Zhurn. Russk. Bot. Obshch. 11, 1-
2, 196.—J. mollis Schmalh. Fl. I] (1897) 110 p. p. non Rchb.; Ldb.. FI.
Ross. II, 766 p. p.; DC. Prodr. 676 p. p—vJ. arachnoidea var. dubia
Korsh. ex Iljin in Tr. Turk. Nauchn. Obshch. II (1925) 21; Fl. Yugo-
Vost. VI, 394.—J. dubia Ijin Ibid 394 (nomen nudum.).
blanda MB. FI. taur.-cauc. III (1819) 549 p. p.

Perennial. Plants 25-100 cm high; tap root branched above. Stems’
solitary or few, cylindrical, erect, striate, flocculose-arachnoid-hairy, like
petiole arachnoid-hairy at base, simple, less often with solitary branches,

660 leafless, except in lowermost part. Almost all leaves in basal rosette;

653

basal leaves gradually narrowed into petiole, pinnatisect into oblong,
lanceolate or less often linear, acuminate or acute segments on each
side, less often undivided (f. integrifolia Ijin); lowermost leaves short-
petiolate or almost sessile, nondecurrent in thin-stemmed forms, reduced
above, undivided, linear, sparse or totally absent; all leaves green above,
more or less scabrous due to short papillae, rugose, covered with
appressed, thin, long hairs emerging from tip of papillae, narrowly
involute, grayish or grayish-whitish due to thin appressed hairs, not
glandular. Capitula solitary terminal on stem large, with many florets,
with florets about 4-5 cm wide. Involucre broadly cupuliform, with
bend at junction, about 20 cm long, with appressed bracts, imbricate
from reduced outer to distinctly exserted inner bracts; outer and middle
bracts densely arachnoid-hairy-tomentose, protruding from tomentum by
only dark green cuspidate tips; bracts of inner row gradually narrowed,
_acuminate, involute with scattered glandular hairs or scarcely visible
appressed bristles, somewhat toothed or ciliolate along border. Receptacle
squamose; scales smooth, acuminate, approximately up to half as long
as involucre. Corolla dark purple, covered with golden sessile glandular
hairs, 15-20 mm long, narrowed over 6—9 mm, with 4-5 mm long lobes
of limb; apical appendages of anthers 2.5—3 mm long, abruptly narrowed
at apex, basal appendages subobtuse, 1.5—2.5 mm long, filaments
glabrous; stigma 2—2.5 mm long, divided up to base, without tuft of
basal bristles, with branches slightly divergent (but not arcuately
recurved), and thereafter fused. Pappus snow-white, 8-12 mm long,
pappus bristles connate into ring, entirely deciduous, scabrous-barbate,
with barbules almost as long as thickness of bristles; achenes
obpyramidal, four-angled, usually 4 mm long, uniformly tuberculate,
cartilaginous along edges, with two rows of cartilaginous tubercles
between edges, and with sessile or very short glandular hairs between
them, with distinctly toothed corona, teeth usually three-toothed in turn;
pappophorus dome-shaped, whitish. Flowering V—VII(VIII); fruiting: VI—
VIII (Plate XL, Figs. 1-2; Plate XLV, Fig. 1.)

In feather grass-sheep’s fescue and herb—sheep’s fescue steppes,
chalk deposits, limestone deposits, pine forests and waste lands.—
European Part: Middle Dnieper; Volga-Don; coastal area; Lower Don;
Trans-Volga Region; Lower Volga; Caucasus: Ciscaucasia, western
Transcaucasia (rarely). Endemic. Described from Volga, below Saratov.
Type in Leningrad.

107. J. talijevii Klok. in Bot. Zhurn. URSR, VII, 4 (1951) 50.
Perennial. Plants 30-90 cm high. Stems solitary or few, erect, simple
or branched in lower half, with elongate branches, finely sulcate, more
661 or less flocculose-arachnoid-hairy. Leaves crowded in lower part; basal

654

leaves numerous, like lower cauline leaves, often all pinnatisect, less
often undivided, usually oblong-linear or linear, sinuate-toothed,
sometimes with small lobes, always sinuate, segments winged along
rachis; other cauline leaves sessile, amplexicaul, somewhat decurrent;
all leaves greenish, finely arachnoid-hairy, rugulose and with scattered
sessile glandular hairs above, white-tomentose beneath. Capitula solitary
terminal on stem and branches, large, with many florets. Involucre
densely arachnoid-hairy; all involucral bracts appressed; deltoid tips of
outer and middle bracts exserted from arachnoid-hairy pubescence and
also appressed as in J. arachnoidea Bge. Corolla dark purple, with sessile
glandular hairs, 12.5-17 mm long, narrowed over 4—7 mm, with 4.5—6
mm long lobes of limb. Pappus snow-white, comprising few, short (5—
6 mm long), scabrous-barbate bristles, with densely arranged barbules,
connate into ring terminating with small nectary and pappophorus, and
shedding easily on maturation. Achenes 3.5—4 mm long, dark olive-
green, with thicker, usually continuous, but somewhat sinuate,
cartilaginous structures along four ribs, with two or three tuberculate-
foveate ribs, scattered sessile glandular hairs along edges, and scabrous
corona, with somewhat recurved teeth, and dome-shaped pappophorus
in center of upper areole. Flowering V—VI; fruiting VII.

Chalk deposits——European Part: Lower Don. Endemic. Described
from the vicinity of Belovodsk. Type in Kiev.

Note. The species, close to J/. cretacea Bge., is distinguished by
having an involucre as in J. arachnoidea Bge., 1.e., the bracts do not
have recurved tips as in the first species, but appressed ones as in the
second species.

108. J. cretacea Bge. in Flora, XXIV, 1 (1841) 158; Ldb. FI. Ross.
II, 763, spec. wolg.; Ijin in Fl. Yugo-Vost. VI, 394.—J. mollis var.
cretacea (Bge.) Fedtsch. and Fler. Fl. Evrop. Ross. (1910) 1005.—(?) J.
litwinowii Zing. Sb. Sved. (1885) 263.

Perennial. Plants 25-50 cm high. Stems solitary or few, simple or
branched, erect, brownish-purple or greenish, ribbed, arachnoid-hairy,
with scattered, fine, sessile glandular hairs; almost all leaves crowded
in lower half of stem; basal leaves numerous, on short, winged, petiole,
pinnatisect into oblong, linear, or almost oval segments, fused with wing
of leaf rachis, mildly sinuate-toothed, often slightly narrowed at base,
always sinuate, sometimes incised, acute or acuminate; cauline leaves
sessile, usually broadened at base, distinctly auriculate and decurrent
with entire or sinuate-toothed wing; lower and middle leaves divided,
like basal leaves; upper leaves usually undivided and entire, linear, less
often almost all leaves undivided (J. litwinowii); all leaves greenish or
grayish-green above, with fine, scattered, sessile, glandular hairs, leaf

655

662 surface soft, not scabrous, white-tomentose beneath due to thin, long,
many-celled hairs. Capitula solitary terminal on stem and branches, 1.5—
3 cm wide. Involucre hemispherical, appressed at base, densely arachnoid
hairy-tomentose, with outer and middle bracts exserted from tomentum;
bracts acuminate, dark green or dark purple, with recurved tips; inner
bracts straight, usually three-veined, aristate, ciliolate, recurved at fruiting,
covered on outer surface with very short, dense, appressed bristles and
sessile glandular hairs. Receptacle squamose; scales long, smooth,
acuminate. Corolla pink-purple, with scattered sessile glandular hairs,
14-16 mm long, narrowed over 5—7 mm, with 4—5 mm long lobes of
limb. Pappus deciduous, snow-white, 5-6 mm long, with short-barbate
bristles. Achenes 3—3.5(4) mm long, dark brown or ferruginous, with
thick ribs, edges usually with three longitudinal-parallel discontinuous
foveate ribs bearing scattered sessile glandular hairs, teeth of corona
somewhat recurved, with pappophorus in center. Flowering V—V]; fruiting
VI.

Chalk beds——European Part: Lower Volga; Lower Don(?). Endemic.
Described from the Kamyshinskii Range, from the vicinity of the village
of Belye Glinki. Type in Leningrad.

Note. S.J. Korshinsky (/zv. Akad. Nauk, 1, 2 (1894) 128) believed
that J. cretacea Bge. is neither a species nor a distinct variety, but only
a mixture of forms intermediate between J. mollis var. dubia and var.
typica, and often simply a form grazed by cattle. J. cretacea Bge. is a
good chalk species, collected by Claus from the chalk beds of the
Kamyshinsk District and by me and Yu.S. Grigorjev in 1926 at the
same place. I agree with Korshinsky only in that the specimen collected
by D.I. Litvinov from the chalk beds of the Ilovli River in the same
region is a modified, mainly pasture, form with entire leaves (J.
litwinowii Zing.), noted by V. Zinger as the species growing together
with sheep’s fescue.

109. J. kasakorum Ijin sp. n. in Addenda XXVI, 725.

Perennial. Plants up to 50 cm high. Stems few or solitary, erect,
simple or with few short branches, longitudinally sulcate, greenish or
purple, flocculose-arachnoid-hairy, with scattered, fine, sessile glandular
hairs. Leaves crowded in lower quarter or one-third of stem; basal leaves
numerous, short-petiolate; cauline leaves sessile, semiamplexicaul and
decurrent with toothed margin; all leaves simply pinnatisect into linear,
less often oblong-linear, distant, sinuate-toothed, acuminate segments;
fused with narrow-winged rachis also toothed at some places, strongly
involute, always crisped along margin; upper leaves distinctly reduced,
grayish-arachnoid-hairy and rugulose above, with fine sessile glandular
hairs, not scabrous, sparsely white-tomentose beneath with sessile

663

656

glandular hairs under tomentum. Capitula solitary. Involucre broadly
cupuliform, depressed at base, about 20 mm long; outer and middle
involucral bracts appressed, triangularly lanceolate, strongly arachnoid-
hairy, covered with sessile glandular hairs, modified apically into short
callose cusp, projecting from tomentum; innermost bracts acuminate-
aristate, not arachnoid-hairy, densely covered with sessile glandular hairs
and very small appressed bristles, ciliate and involute. Receptacle densely
squamose; sczles smooth, acuminate, one-third to half as long as
involucre. Corolla pink, 12-13 mm long, narrowed over 4—5 mm, with
4.5 mm long lobes of limb; apical appendages of anthers 2.5—3 mm
long, abruptly narrowed at apex into short, incurved cusp. Pappus
deciduous, snow-white, 6-8 mm long, with short-barbate, dense bristles,
connate into short-cylindrical structure, flattened above, with short upper
margin, surrounding pappophorus as cap, with small, five-toothed, nectary
in center. Achenes usually 3—3.5 mm long, pale yellowish, later dark
brown, thick-cartilaginous along ribs, with transverse, parallel, dense
folds, sometimes discontinuously tuberculate-alveolate, edges with
three or four cartilaginous, parallel more or less approximate ribs,
discontinuously foveate or sinuate, discontinuously scattered, fine, sessile
glandular hairs, and well-developed, erect, almost membranous, scabrous
corona, connate at base, later becoming almost cartilaginous. Flowering
V-VI1(?); fruiting VI.

Chalk beds.—European Part: Lower Don, Lower Volga. Endemic.
Described from the vicinity of Kletskaya Station. Type in Leningrad.

110. J. ledebourii Bge. in Flora, XXIV, 1 (1841) 157—J. pseudo-
mollis Klok. nom. nud. in Bot. Zhurn. VII, 4 (1951) 46.—J. mollis B.
dubia Korsh. in Izv. Akad. Nauk, 1, 2 (1894) 124 p. p—J. mollis y.
arachnoidea Korsh. Ibid. 125, specim. e Tula, Orel, Woron. Ryasan,
Tamb. Simb. Pensa, Kazan, Ufa, etc——J. mollis DC. Prodr. VI (1837)
676 (specim. sibir. Ross. merid.); Ldb. Fl. Ross. II, 2, 766 (specim.
uralens.); V. Zinger, Sb. sved. (1885) 264; Schmalh. Fl. II, 110 p. p.;
Fedtsch. and Fler. Fl. Evrop. Ross 1005 p. p.—J. arachnoidea auct.:
Stank. and Tal. Opr. Vyssh. Rast. (1957) 410 (specim. tamb. tatar.
baschk.)

Perennial. Plants (15)25—70(90) cm high. Stems few or solitary,
usually simple, less often with solitary branches, basally. sulcate and
finely striate, somewhat thick, 2.5-6 mm at base, covered with fine,
dense arachnoid hairs and scattered fine glandular hairs, usually in
furrows; usually leafy in lower part. Basal leaves numerous, narrowed
into winged petiole, covered with divergent, soft and long, hairs forming
woolly coating; cauline leaves also narrowed toward base, but slightly
broadened at point of attachment, often semiamplexicaul and decurrent;

664

657

uppermost leaves reduced and simple, usually filiform, sessile, not
decurrent; basal, lower and middle cauline leaves usually pinnatisect
into linear, oblong or lanceolate, less often entire or cuspidate segments,
fused basally with entire wing of rachis, less often all leaves undivided
and entire (f. integrifolia Iljin), or only lower leaves undivided and
entire, sometimes only their tip three-lobed, all leaves dark and grayish-
green or bright green above, slightly scabrous due to fine tubercles,
with long and thin many-celled hairs at apex usually semiappressed,
leaf rugulose, with scattered fine sessile glandular hairs, grayish or whitish
beneath due to sparse tomentum, covering sessile glandular hairs. Capitula
with many florets, usually 40-50 mm wide. Involucre broadly cupuliform,
usually 20-30 mm long; outer rows of involucral bracts more or less
distinctly arachnoid-hairy, except outermost two to five oblong bracts,
others narrowed from ovate-oblong, somewhat raised, convex more pale
base into more or less long and narrow, usually loosely appressed tip,
less often gradually recurved, dark green or purple sometimes decurved,
almost as in J. mollis (L.) Rchb., later breaking off; all bracts with
sessile scattered glandular hairs, ciliate; innermost bracts acuminate,
densely covered on outer side with short bristles or sessile glandular
hairs, or both, with declinate margin, distinctly ciliate. Receptacle densely
squamose; scales relatively long, smooth, acuminate. Corolla pink or
purple, with dense sessile glandular hairs, (15)17—23 mm long, narrowed
over (6)7—9 mm, with 5-89 mm long lobes of limb. Pappus with 8-12
mm long bristles, snow-white, becoming somewhat yellowish toward
maturation; bristles short-barbate, dense, connate into cylindrical cover
with small depression at apex enclosing nectary; on fruit maturation
cover slides down from pappophorus together with pappus. Achenes
dark brown or almost black, less often lighter, 44.5 mm thick, with
three or four parallel, approximate rows of discontinuous tubercles on
edges, ribs with thicker and more sparse, fine, sessile glandular hairs,
with toothed corona, surrounding dome-shaped pappophorus. Flowering
V-VI; fruiting VII—VIII.

Forest-steppe and steppe zone in meadows, feather grass-herb and
stony steppes, rocks, limestone deposits and chalk beds on slopes.—
European Part: Volga-Kama, Volga-Don, Trans-Volga Region. Endemic.
Described from Orenburg Region, from the vicinity of the village of
Spasskoye. Type in Paris.

111. J. mollissima Klok. in Bot. Zhurn. Ukr.RSR, VII, 4 (1951)
49. —J. mollis DC. Prodr. VI (1837) 676 (specim. bessarab.) non Rchb.;
Ldb. Fl. Ross. II, 766 (p.p. spec. cherson.); Schmalh. Fl. II, 111 p. p.—
J. mollis y. arachnoidea Korsh. in Izv. Akad. Nauk, 1, 2 (1894) 125
(specim. odess.).

66

n

658

Perennial. Plants (15)20-—50(70) cm high. Stems erect, solitary or
few, sulcate-finely ribbed, usually whitish due to more or less dense
tomentum or flocculose-arachnoid-hairy with erasing pubescence, with
scattered, fine, sessile glandular hairs, usually simple, less often weakly
branched. Basal leaves more or less numerous, sessile, but sometimes
gradually narrowed toward base, broadened at base and somewhat
amplexicaul, pinnatisect into linear or oblong, less often oblong-oval,
acuminate or cuspidate, usually entire segments, often revolute, less often
toothed and sinuate (f. crispa Iljin), basally fused with broadly winged
rachis, almost equal in width; lower cauline leaves like basal, mainly
sessile, semiamplexicaul and often asymmetrical, decurrent; upper leaves
often undivided, uppermost narrow, reduced, sessile; all leaves grayish
above due to sparse arachnoid-hairy pubescence, not scabrous, rugulose,
white-tomentose beneath. Capitula with many florets, 30-50 mm wide.
Involucre 15-20 mm long, broadly cupulate, densely arachnoid-hairy;
outer involucral bracts oblong-lanceolate, acuminate and involute above
middle, with somewhat colored tips; innermost bracts narrowly lanceolate,
acuminate-aristate, erect, densely glandular on outer side, ciliate, involute.
Corolla pink-purple, with sessile glandular hairs, 15—18 mm long,
narrowed over 5—7 mm, lobes of limb 4-6 mm long. Pappus 8-10 mm
long, snow-white, with short-barbate bristles, innermost bristles forming
short-cylindrical dome, with nectary in center; on maturation, cover slides
off pappophorus. Achenes 2.5—3.25 mm long, usually black or blackish-
brown, with rusty, somewhat thick nbs and 2-4 parallel longitudinal,
discontinuous ribs along margin and ferruginous corona; ribs at four
corners of achene cartilaginous, their broad and cuspidate margin
obliquely directed; fine sessile glandular hairs scattered between ribs.
Flowering V—VI; fruiting VI—VII.

In maritime zone; lagoons, herb-steppe and stony slopes.—European
Part: Black Sea Coast (from the mouth of the Danube River up to
Kherson).—General distribution: Rumania (Dobruja). Described from
Izmail Region, near the village of Budaki (town of Primorsk). Type in
Kiev; isotype in Leningrad.

112. J. calearea Klok. in Bot. Zhrun. Ukr.RSR, VII, 4 (1951) 47.—
J. mollis Ldb. Fl. Ross. II (1845-1846) 767 p. p. non Rchb.; DC. Prodr.
VI, 676 p. p., Schmalh. Fl. II, 110 p. p—dJ. mollis var. dubia Korsh. in
Izv. Akad. Nauk, 1, 2 (1894) 124 p. p—J. mollis var. arachnoidea
Korsh. Ibid, 124 p. p.

Perennial. Plants 30-100 cm high. Stems solitary or few, usually
erect, simple, less often branched, with solitary long branches, usually
finely ribbed-sulcate below, sometimes above middle, flocculose-
arachnoid-hairy, arachnoid-woolly below, with scattered sessile glandular

659

hairs, yellowish-green. Leaves crowded in lower part, sometimes up to

666 one-third of stem height; basal leaves more or less numerous, like lower
cauline leaves, gradually narrowed toward base and woolly here,
semiamplexicaul, pinnatipartite or pinnatisect into distant, oblong, oblong-
oval, lanceolate or less often linear, entire, acuminate or cuspidate
segments, less often segments crispate (f. crispata Iljin), occasionally
leaves almost bipinnatisect (f. bipinnata I\jin), sometimes all leaves
undivided or only upper leaves undivided and entire; middle cauline
leaves sessile, almost always semiamplexicaul, often auriculate and
decurrent; uppermost leaves sessile, smaller; all leaves rugulose, green
or finely arachnoid-hairy above, lacerate, very small, somewhat hard
papillae with long, many-celled, decumbent or semidecumbent hairs at
tip, leaves soft or weakly scabrous, grayish beneath due to loose
tomentum. Capitula solitary terminal on stem and branches, with many
florets, usually 3-4 cm wide. Involucre broadly cupuliform, depressed
below, 15—20 mm long, arachnoid-hairy; involucral bracts initially usually
appressed, later outer bracts slightly divergent, oblong, acuminate, slightly
recurved; innermost bracts oblong-lanceolate, aristate, involute and ciliate
densely glandular on outer side, with very short hairs. Corolla pink-
purple, densely glandular, usually 15-18 mm long, narrowed over 6-8
mm, lobes of limb 4-6 mm long. Pappus together with projecting, 10—
12 mm long, dense and short-barbate bristles; innermost bristles connate,
forming cover inside ring, with nectary at tip easily sliding off
pappophorus on maturation. Achenes 3-4 mm long, olive green, later
dark brown, with cartilaginous, lighter ribs, consisting of quite
approximate structures pointed upward with small teeth; edges with
parallel cartilaginous ribs, bearing scattered, fine tubercles and sessile
glandular hairs in-between, with cartilaginously toothed corona above,
along edges, enclosing a narrow dome-shaped papvophorus. Flowering
V-VII; fruiting VI-VIII.

Steppe slopes, forest edges, and river banks.—European Part:
Bessarbia, Upper Dnieper, Middle Dnieper, Black Sea Coast (northern
part); Volga-Don (west). General distribution: Poland (?). Described from
Kherson Region. Type in Kiev.

113. J. michelsonii Iljin sp. n. in Addenda XXVI, 726.

Perennial. Plants 50-60 cm high, with short vertical rhizome covered
with brown scales. Stems usually few, erect, striate, finely sulcate,
uniformly covered with soft, slightly erect, many-celled arachnoid
helicoid hairs and scattered, sessile glandular hairs with long branches
in lower one-third part, upward spreading. Basal and lowermost cauline
leaves broadened downward, broadly amplexicaul and here more densely

667 covered with soft hairs; cauline leaves with shorter, but also flat, and

668

660

strongly winged petioles, auriculate and decurrent; all leaves pinnatisect
into linear, acuminate, somewhat involute, distant, almost opposite’
segments, decurrent along flat rachis, green, rugulose and weakly
scabrous above, with scattered sessile glandular hairs, grayish beneath
due to soft, sparse arachnoid-hairy pubescence. Capitula usually 4-5 cm
wide, with many florets. Involucre broadly cupuliform, 20-25 mm long;
outer and middle involucral bracts arcuately recurved, acuminate, dark
violet in upper half, mixed with long arachnoid hairs; inner bracts erect,
acuminate, aristate, ciliate, with short semiappressed bristles and scattered
sessile glandular hairs on outer side. Corolla violet-purple, densely
glandular, 15-16 cm long, narrowed over 5—6 mm length, with 5-6 mm
long lobes of limb. Pappus entirely deciduous, bristles short-barbate,
barbs as long as or slightly shorter than thickness of bristles. Immature
achenes with alveolate tuberculate structure. Flowering V; fruiting ?.
Forest edges.—European Part: Upper Dniester. Endemic (?).
Described from the Maksimovka-Bogdanovka District, Romanovka, from
the collections of A.I. Mikhelson, June 9, 1916. Type in Leningrad.

114. J. sordida Stev. Taur. (1857) 227.—J/. roegneri C. Koch in
Flora, XXIV (1951) 414.—J. mollis DC. Prodr. VI (1837) 676 p. p. non
Rchb.; Schmalh. Fl. II, 110 p. p.; Ldb. Fl. Ross. II, 767 p. p.—J. mollis
8. dubia Korsh. and var. arachnoidea Korsh. in Bull. Acad. Sc. Pétersb.
V. sér. 1, 2 (1894) 123 p. p.—J. arachnoidea Boiss. FI. or. III (1875)
574 p. p. non Bge.—J. ledebouri Stev. op. cit. 228, non Bge.—Carduus
mollis MB. Fl. taur.-cauc. II (1808) 272, non Serratula blanda MB.
Ibid. (1819) 549.

Perennial. Plants 25-90 cm high. Stems erect, cylindrical, finely
striate, arachnoid-hairy, usually flocculose-arachnoid-hairy, solitary or
few, simple or less often with few long branches in lower half, less
often leafless up to middle. Basal leaves petiolate, with winged petiole;
cauline leaves usually approximate in lower part of stem, with lamina
pinnatipartite or almost pinnatisect into linear or oblong-linear, acuminate
lobes, occasionally undivided (f. integrifolia); in lower leaves often
oblong, subobtuse, with short cusp, wings sinuate, very rarely nonsinuate
(f. platyphylla), fused with winged rachis; all leaves dark green or grayish:
green above, more or less arachnoid-hairy, with sessile glandular hairs,
except uppermost, almost filiform, small, sessile, and decurrent with
simple or sinuate-toothed wing. Capitula with many florets, 2.5—-4 cm
wide, solitary. Involucre broadly cupuliform, 20-25 mm long, with many
bracts, densely arachnoid-hairy in lower half, less often almost glabrous;
outer bracts exserted from tomentum as long, acuminate, almost glabrous
tips with scattered sessile glandular hairs, ciliolate along margin, initially
appressed, later divergent and even recurved; inner bracts strongly

Plate XLIX.

1—Jurinea atropurpurea Winkl., 2—J. propingua Iljin.

661

670

662

projecting, aristate, pale in upper half, somewhat purple, revolute,
ciliolate, on outer side covered with sessile glandular hairs, often with
mixture of scarcely visible appressed bristles or only with latter.
Receptacle densely squamose; scales smooth, pointed, up to one-third
to one-fourth as long as involucre. Corolla 14-20 mm long, narrowed
over 5—8.5 mm, with 5-7 mm long lobes of limb, and scattered glandular
hairs. Pappus 10-12 mm long, snow-white, bristles barbate, strongly
connate into ring, entirely deciduous; barbules shorter than thickness of
bristles, dense, pappus dropping off with nectary. Achenes brown to
olive green, less often pale, 3-5 mm long, four-angled, obpyramidal,
discontinuously cartilaginous along ribs, somewhat thickened, with thinner
three or four cartilaginous rows of tubercles along edges, and fine sessile
glandular hairs in-between; corona well developed, two- or three-toothed
along edges; pappophorus dome-shaped. Flowering V; fruiting VI.

Chalk beds, limestone deposits, and marls.—European Part: Crimea.
Endemic. Described from Crimea. Type in Helsinki.

115. J. blanda (MB.) C.A.M. Verz. Cauc. (1831) 67.—J.
arachnoidea Sosn. and var. caucasica Sosn. in Zhurn. Russk. Bot.
Obshch. 11 (1926) 196, 197; Grossh. Fl. Kavk. IV, 173; Schchian in FI.
Kavk. VIII, 447—J. mollis var. arachnoidea Korsh. in Izv. Akad. Nauk
V, Ser. 1, 2 (1894) 124, 125.—J. arachnoidea Ldb. Fl. Ross. II (1845—
1846) 766 p. p. non Bge.; Boiss. Fl. or. III, 575; Perech. Rast. Turk.
IV, 301 p. p.; Iljin in Tr. Turk. Nauchn. Obshch. II, 24.—J/. mollis Ldb.
l.c. non Rchb.—? J. alkpigena C. Koch. in Linnaea, XXIV (1845) 414;
Sosn. op. cit.—Serratula blanda MB. FI. taur.-cauc. III (1819) 548 p. p.

Perennial. Plants 15-75 cm high. Stems solitary or few,
longitudinally striate, arachnoid-hairy, often floccose, erect, simple, less
often branched in lowermost part, with long branches, leafy only in
lower part. Leaves rosulate, petiolate, like cauline leaves usually
pinnatipartite or divided; middle cauline leaves sessile or more or less
decurrent; uppermost leaves reduced, simple, undivided and entire, often
crisped (f. crispata Iljin), less often all leaves undivided and entire (f.
integrifolia Iljin), green, glabrous or weakly arachnoid-hairy, and smooth
or scabrous above, grayish or white-tomentose beneath. Capitula solitary,
with many florets, medium or large. Involucre usually purple, especially
in upper part, sometimes pale green; involucral bracts appressed, but
often outer bracts recurved; inner bracts longer, erect, gradually
acuminate, all bracts small-toothed-ciliate. Pappus entirely deciduous,
short-barbate, almost serrate-barbate, all pappus bristles connate into ring,
enclosing more or less cylindrical pappogenus with five-toothed,
membranous, sessile, nectary at apex. Achenes 3-4 mm, less often 5
mm long, four-angled, with parallel, discontinuous tuberculate-foveate

67

—

663

ridges, sometimes indistinct and always covered with more or less sessile
or short-stalked glandular hairs; corona toothed above along edges,
sometimes not developed. Flowering ([V)V—VII; fruiting VI—VIII.

Mountain and subalpine meadows, rocks, especially limestone de-
posits, steppes.—Caucasia: Western, eastern and southern Transcaucasia.
General distribution: Asia Minor and Iran. Described from Caucasus.
Type in Leningrad.

Note. M. Bieberstein, while describing his species under the name
Serratula blanda MB., reported that this species is distributed throughout
the Caucasus, as well as in Southern Russia and Ukraine; this species
appears in the second volume of his Flora as Carduus mollis and is
reported only from Crimea, but in Volume III the author includes this
name among the synonyms of Serratula blanda. \n the herbarium of
Bieberstein, plants from the Caucasus, Crimea, Southern Russia, and
Ukraine are present on the same sheet under both names. Since the
Crimean plants are all similar, it is best to adopt Steven’s name, J.
sordida Stev., for them, which agrees well with the original specimens;
the Caucasian plants should be called J. blanda (MB.) C.A.M., as
reported by Bieberstein for the entire Caucasian region. As regards the
Ukrainian and Central Russian plants, the former, evidently, would be
J. mollissima Klok., and the latter J. ledebourii Bge. J. blanda (M.B.)
C.A.M. is an aggregate of elementary species whose borders cannot be
defined at this time.

116. J. exuberans (Trautv.) Sosn. in Zhurn. Russk. Bot. Obshch.
11 (1926) 197; Grossh. Fl. Kavk. IV, 173; Schchian in Fl. Gruzii, VIII,
448.—J. arachnoidea var. exuberans Trautv. in Tr. Peterb. Bot. Sada,
V, 1 (1877) 449.

Perennial. Plants very robust, at least 60-70 cm high. Stems erect,
somewhat thick, with long branches in lower part, arachnoid-hairy and
weakly scabrous like upper leaf surface due to fine papillae; branches
spreading upward. Basal and lower cauline leaves gradually narrowed
into long winged petiole, erect, 15—25 cm long, less often undivided,
usually pinnatipartite, sometimes with linear or lanceolate lobes only in
lower part of lamina, lobes directed upward, acute and cuspidate, distant,
entire; middle cauline leaves sessile and long-decurrent, with lamina
like basal leaves; uppermost leaves linear, almost filiform, undivided
and entire, usually bent; all leaves green, somewhat rugose above, gray
arachnoid-hairy beneath. Capitula solitary. Outer and middle involucral
bracts recurved, arachnoid-hairy, like inner bracts veined; innermost
bracts erect, with scarcely visible appressed bristles on outer side.
Receptacle squamose, scales slightly less than half as long as involucral
bracts, smooth, acute. Corolla pink-purple, 12-15 mm long, narrowed

672

664

at 3-4 mm, with 5-6 mm long limb. Pappus entirely deciduous, 6-8
long, pappus bristles connate into ring forming cylindrical pappogenus
at base, with sessile membranous nectary at truncate apex. Achenes 4—
5 mm long, with longitudinal transversely alveolate ribs along edges,
with toothed corona above, enclosing dome-shaped pappophorus along
edges. Flowering VI(?); fruiting VII.

Alpine zone.—Caucasus: Eastern Transcaucasia. Endemic. Described
from Atsunty from the Khidotan foothills between Khevsuria and
Tushetia. Type in Leningrad.

Note. This species was collected only by Radde. Other specimens
in the Harbarium of the Botanical Institute of the Academy of Sciences
of the USSR and the Botanical Institute of Georgia under this name do
not belong to it. The specimen collected by Shelkovnik from the Abakura
Pass, mentioned by D.I Sosnowsky, could not be seen; apparently, it is
present in Armenia.

Series 2. Anatolicae Iljin—Markings on achenes less sharp and
strongly masked by dense glandular hairs, sessile as well as short-stalked.

117. J. demetrii Iljin sp. n. in Addenda XXVI, 727.

Perennial. Plants 25-60 cm high, yellowish-green, slender. Stems
covered with large, brownish, scaly remains of bases of old leaves,
producing long, slender branches from base, angular-striate, with
numerous sessile glandular hairs, finely and sparsely arachnoid-hairy.
Leaves crowded in lower half of stem; basal and lower cauline leaves
on relatively short, winged, petioles, semiamplexicaul like middle leaves,
pinnatisect, sessile at base, auriculate and slightly decurrent, segments
distant, linear, cuspidate, with brownish, cartilaginous, short, cusp, entire,
sometimes involute up to midrib, often arcuately directed upward; upper
leaves undivided and entire, almost filiform; all leaves green above,
often greenish due to fine, sparse, arachnoid-hairy pubescence, rugulose,
scabrous due to cartilaginous tubercles, with sessile glandular hairs in
alveolae, and depressed midrib, whitish-grayish beneath due to more
dense pubescence, with promiment midrib. Capitula solitary, usually
medium, with many florets. Involucre cupuliform, 12—18(20) mm long,
slightly arachnoid-hairy only at base, outer rows of bracts recurved,
dark green and dark purple-green on outer side, with short, brownish,
cartilaginous cusp, erect, and easily breaking off; inner bracts linearly
lanceolate, gradually acuminate, purple, brownish above, very distinctly
toothed, covered with numerous sessile glandular hairs. Corolla 13-18
mm long, purple, glandular and arachnoid-hairy at some places, gradually
narrowed over 5—9 mm, with 5-6 mm long lobes of limb. Pappus
deciduous, 10-12 mm long, snow-white, pappus bristles scabrous-barbate

665

somewhat flat, connate into ring, barbules distinctly shorter than thickness
of bristles, very compact in two rows, with pappogenus inside ring on
membranous base, with nectary above; achenes 2.5—3 mm long, with
toothed corona, longitudinally ribbed along edges, with alternating
alveolae and tubercles, sparsely covered with lustrous, sessile or short-
stalked, glandular hairs with dome-shaped pappophorus in center.
Flowering VI; fruiting VI—VII.

On rocks.—Caucasus: Southern Transcaucasia(?). General
distribution: Asia Minor. Described from the former Karsk Region from
a location between Bardu and Promezhutochnyi. Type in Leningrad.

Note. This species differs quite sharply from all other species of
this section growing in the USSR in the structure of the achenes as well
as in general morphology.

Section 14. Jurinea——Sect. Platycephalae Benth. and Hook. Gen.
pl. III (1873) 473; subsect. 2. Lyratae Sosn. in Zhurn. Russk. Bot.
Obshch. 11 (1926) 195.—§ 3. Pinnatae Boiss. (p. p.) Fl. or. III (1875)
572 p. p.—Subsect. Alatae Sosn. op. cit. 198.—§ 33. Decurrentes DC.
Prodr. VI (1937) 675.—Large thistle-like plants with lyrate basal leaves.
Pappus of serrate or very short-barbate bristles connate at base into
ring, shedding entirely together with pappogenus and nectary. Achenes
small, with conical or dome-shaped pappophorus.

Type species: Type of section.

—

. Stem strongly winged ............... LSE. 118. J. alata (Desf.) Cass.
HaStempnot WANSeM earls! sorted, cree. Cott. tedodd. «ete. MOEN. ORES. be Dee
2. Involucral bracts narrow, linear, lax, like in J. cyanoides, not prickly

and not recurved; achene four-angled, obpyramidal, light brown,

with whitish ribs, corona, and hilum... 121. J. ruprechtii Boiss.
+ Involucral bracts two to three times broader, prickly, outer bracts
sharply recurved, like in thistles (C. nutans); achene different

3. Achenes strongly compressed, two-angled, but four-nbbed, see
with dark transverse spots, hilum lateral «0.0.0.0... eee
ee AES Marl whee. eA Oded tt 120. J. grossheimii Sosn.

+ Achenes four-angled, obpyramidal, blackish-brown, with somewhat
undulate surface and white-cartilaginous markings of edges. ........
sca one ee. wepeeeeene. TALE oe. 2 119. J. spectabilis Fisch..and Mey.

673 Series 1. Pterocladae [ljin—Stem winged.

118. J. alata (Desf.) Cass. in Dict. sc. nat. XXIV (1826) 288; DC.
Prodr. VI, 675; Ldb. FI. Ross. II, 2, 762; Boiss. Fl. orient. III, 576;

674

666

Grossh. Fl. Kavk. IV, 171; Schchian in FI. Gruzii, VIII, 448.—Serratula
alata Desf. Catal. Paris, ed. II (1815) 108.

Perennial. Plants 25—75 cm high. Tap root non-fibrous. Stems solitary
from base, less often strongly branched above middle, with long strongly
divergent branches, ribbed-sulcate, somewhat arachnoid-hairy, but green,
with scattered sessile glandular hairs. Basal leaves petiolate, with more
or less winged petiole; lower cauline leaves also petiolate, with broad,
entire or often large-toothed wing, strongly decurrent; other leaves sessile,
semiamplexicaul and decurrent, stems and branches almost leafless in
upper part, with filiform apical leaves; all leaves, except uppermost,
gradually acuminate, undivided -and mostly entire, lyrate, with large,
oval or oblong-oval, large-toothed, obtuse or cuspidate terminal lobe,
and usually three to five roundish and obtuse lateral lobes; all leaves
green above, greenish-whitish beneath due to fine pubescence. Capitula
numerous, with many florets. Involucre hemispherical, arachnoid hairy-
pubescent in lower part, six- to eight-rowed, outer rows of bracts narrowly
oblong, with recurved herbaceous tips, subobtuse in outer row, others
acuminate; inner rows straight, narrowly lanceolate, acuminate; bracts
of inner row usually long-aristate, membranous along apical margin; all
bracts with three to five (seven) prominent veins (except outer row),
densely covered with glandular hairs, and often also with scarcely visible
bristles, usually along veins, margins ciliolate, sometimes only in lower
part. Receptacle more or less distinctly alveolate, densely covered with
long, smooth, acuminate scales, at least up to half as long as involure.
Corolla pink-purple, with sessile glandular hairs, 12-17 mm long, on
5.5—7 mm long stalk, with 3.5-5 mm long lobes; apical appendages of
anthers 2.5—-3 mm long, gradually acuminate, basal appendages setaceous,
2—3 mm long, usually undivided filaments glabrous. Pappus deciduous,
snow-white, 6-7 mm long (excluding two to four prominent bristles),
pappus bristles short-barbate, inner row bristles shorter-barbate, flattened,
barbules arranged on one side along edge, all bristles connate into
ring with extended membranous base, enclosing a glass-shaped nectary
with five longitudinal lobes, easily shedding entirely. Achenes usually
4 mm long, obpyramidal, regularly four-angled, blackish-brown, lustrous,
deeply rugose, with small-toothed, usually whitish corona at
apex and with white ribs, often with two or three-toothed appressed
scales only in upper part and along ribs, surrounding smooth dome-
shaped pappophorus above. Flowering VI-VIII; fruiting VII-IX.

Stony xerophytic mountain slopes, limestone deposits, reaching
alpine zone.—Caucasus: Ciscaucasia. Endemic. Described from Terskaya
Region. Type in Paris.

Series 2. Apterae Iljin—Stem wingless.

675

667

119. J. spectabilis Fisch. and Mey. in Bull. Soc. Nat. Mosc. VI
(1833) 252; Fisch. and Mey. Ind. sem. horti. Petropol. IV, 39; DC.
Prodr. VI, 678; Ldb. Fl. Ross. II, 764; Boiss. Fl. or. UI, 573; Grossh.
Fl. Kavk. IV, 172; Sosn. in Zhurn. Russk. Bot. Obshch. XI, 195.

Perennial. Plants 15-40 cm high, possibly taller, with tap roots.
Stems solitary or few, thick, fistular, ribbed-sulcate, green or dirty purple,
more or less arachnoid-hairy, densely covered with sessile glandular
hairs, branched from base or commonly in upper two-thirds with upward
spreading branches, and remains of old leaves at base. Basal leaves
numerous, gradually narrowed into flat petiole, basally broadened; cauline
leaves short-petiolate, both sinuate-imparipinnate, divided, almost lyrate
above; lateral lobes numerous, oblong or oblong-oval, broadly sessile
and somewhat narrowed at base, usually sparsely sinuate-toothed,
subobtuse, subacute, or cuspidate, opposite or obliquely opposite, fused
with winged rachis; terminal lobe often larger, broadly ovate or oblong-
ovate, less often. undivided, usually deeply divided, sinuate-toothed, fused
with lateral lobes; upper cauline leaves lanceolate, toothed, uppermost
leaves linear, almost undivided; all leaves strongly rugose above,
glabrous, with scattered sessile glandular hairs in alveoli, arachnoid hairy-
pubescent along depressed veins, densely white-tomentose beneath, and
with dense glandular hairs under tomentum, in shade-loving forms (f.
umbrosa \\jin) somewhat grayish and distinctly glandular beneath.
Capitula 3—6 cm long, solitary, axillary almost from stem base. Involucre
20-30 mm long; outer involucral bracts divaricate and distinctly recurved,
acuminate, prickly, arachnoid-hairy, outer bracts single-furrowed (on inner
side); innermost bracts also acuminate, erect, with prominent vein, usually
with appressed, short, usually purple bristles, sometimes with scattered
sessile glandular hairs. Receptacle indistinctly alveolate, alveole
surrounded by relatively short more or less basally connate, acuminate,
smooth, scales. Corolla pink-purple, covered with sessile glandular hairs,
18-21 mm long, narrowed at 10 mm; apical appendages of anthers 3—
3.5 mm long, acuminate, basal appendages usually setaceous, 2-3 mm
long; filaments finely tuberculate (magnification < 56); stigma 2—2.5 mm
long, divided up to base, with short tuft of bristles at base, modified
into papillae above. Pappus colorless, detaching with difficulty, 10-15
mm long, fragile, pappus bristles bent, unequal, and connate into ring.
Achenes oblong-obpyramidal, four-angled, blackish-brown or grayish-
brown, with whitish cartilaginous markings, especially along ribs, and
with whitish rostellum, with weakly undulate surface, glabrous, forming
toothed border along edges, with conical pappophorus, 3-4 mm long.
Flowering and fruiting V—VII.

High mountain zone, mostly on rocks.—Caucasus: Southern and

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eastern Transcaucasia. General distribution: Iran(?). Described from
vicinity of Shusha. Type in Leningrad.

Note. In describing J. coelestis Sosn., D.I. Sosnowsky suggested a
hybrid (/. depressa x J. spectabilis) origin for this species. The material
examined does not give such an impression, although in its morphology
(achenes, pappus, and corolla) it shows a transition from the genus
Jurinella Jaub. and Spach. to the genus Jurinea Cass. A true corona has
still not developed; only the upper edges form the toothed border.
Formally this species should be assigned to the genus Jurinea Cass.

120. J. grossheimii Sosn. in Grossh. Fl. Kavk. IV (1934) 172.—J.
spectabilis var. monocephala Trautv. in herb.

Perennial. Plants 25-50 cm high or more. Stems erect, cylindrical,
branched in upper half, less often simple, sulcate, arachnoid-hairy, with
scattered sessile glandular hairs. Basal leaves rosulate, usually lyrately
divided or incised, with roundish. or broadly ovate, undivided, large-
toothed, terminal lobe, with cuspidate teeth or usually divided into three
lobes, lateral lobes oblong or obovate, toothed, reduced toward petiole;
lamina gradually narrowed into short or longer petiole broadened at
base; middle leaves reduced or almost undivided and entire, oblong-
lobate, with apical cusp, only uppermost leaves linear, undivided; all
leaves green and rugulose above, with sessile glandular hairs in
depressions, white-tomentose beneath or densely glandular under
tomentum. Capitula few or solitary in single-stemmed plants (f.
monocephala I\jin), 2.5-—4 cm wide, with many florets. Involucre
arachnoid-hairy at base or entirely glabrous; outer bracts narrowly
lanceolate, green, gradually acuminate, decurved, prickly, innermost
bracts erect, long, finely acuminate, purple. Corolla pink-purple, with
scattered sessile glandular hairs, 17-20 mm long, narrowed in middle,
with 6-7 mm long lobes of limb. Pappus divaricate; pappus bristles
fragile, scabrous, connate into ring, exceeding achene several times, two-
tiered pappogenus with upper dark purple, discoid part and lower lighter
part developed at base of ring, lower part terminating in membranous
nectary. Achenes strongly compressed, four-angular, oblong-triangular,
4—5 mm long, glabrous, smooth, dark brown, with few dark transverse
striations, and light-colored, equal small-toothed corona at apex,
surrounding whitish, flat, dome-shaped lower part of pappophorus with
truncate apex, hilum lateral. Flowering and fruiting VI—VIII.

Stony slopes.—Caucasus: Eastern Transcaucasia. Endemic?
Described from Karabakh, in the vicinity of Shusha Region. Type in
Tbilisi.

Note. Grossheim expressed doubt in the Flora Kavkaza [Flora of
the Caucasus] about the separate status of this species. In his opinion,

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669

this is only a form of J. spectabilis Fisch. and Mey. However, these
species are similar only in their growth habit, but are quite different in
their involucral bracts and the shape of the achenes.

121. J. ruprechtii Boiss. Fl. or. HI (1875) 573; Grossh. Fl. Kavk.
IV, 172; Sosn. in Zhurn. Russk. Bot. Obshch. XI, 195.

Perennial. Plants 15—75 cm high. Stem more or less erect, cylindrical,
sulcate-finely ribbed, appressed finely arachnoid-hairy, covered with
numerous scattered sessile glandular hairs, branched from base; branches
elongate and abruptly rising upward. Leaves in basal rosette numerous, .
usually undivided until development of stem, oval, long-petiolate, later
withering, other leaves in rosette lyrate, short-petiolate with large, ovate
or roundish, usually entire, terminal lobe and oblong or oval, also usually
entire, distant lateral lobes reduced below; caulir2 leaves not decurrent,
lower short-petiolate, incised like basal, upper leaves sessile, narrowed
toward base, oblong-obspatulate or less often oblong-oval, usually entire,
all leaves green but sparsely arachnoid-hairy. above, covered with
scattered sessile glandular hairs, white-tomentose beneath, also covered
with sessile glandular hairs. Capitula numerous, solitary, less often two
or three, terminal on stem and branches, with many florets. Involucre
cupuliform, 20-25 mm long, usually six-rowed; involucral bracts linear-
narrowly lanceolate, gradually acuminate; outer bracts finely arachnoid-
hairy, herbaceous-green in recurved half, usually squarrose; all bracts
sharply recurved after fruiting; innermost row of bracts erect, broader,
with distinct veins, scarcely toothed along upper margin; all bracts
covered with scattered sessile glandular hairs. Receptacle squamose;
scales relatively short, smooth, connate almost up to one-third or half
of their length and surrounding each alveole. Corolla pink-purple, densely
glandular, 12-13 mm long, with 4-5 mm long stalk, and about 2 mm
long lobes of limb; apical appendages of anthers acuminate, 2.5 mm
long, somewhat divergent; basal appendages sinuate-caudate; filaments
glabrous; stigma with short tuft of bristles at base, finely papillate on
outer side, about 2 mm long. Pappus about 10(12) mm long, three or
four pappus bristles longer and thicker, somewhat fragile, all scabrous,
connate into ring, detaching with difficulty. Achene about 3 mm long,
obpyramidal, four-angled, lustrous, blackish-brown, whitish-cartilaginous
along ribs and corona, weakly cylindrical, often with distinct
small teeth along ribs below corona. Corona with all equal small teeth,
and with flat dome-shaped pappophorus at apex. Flowering V—VII;
fruiting VI-VIII.

In alpine and subalpine zone on limestone and chalk beds.—
Caucasus: Dagestan. Endemic. Described from the region of Avarskoye
Koisu. Type in Leningrad.

678

670

Section 15. Chaetocarpae (Korsch.) Iljin comb. n.—Chaetocarpae
Korsh. in Izv. Akad. Nauk, 1, 2 (1894) 126 pro stirpe.-—Sect. Pinnatae
Boiss. Fl. or. III] (1875) 572 p. p.—Achenes always with parallel
longitudinal ribs on edges, tuberculate, discontinuous and modified into
teeth in upper part up to base of toothed corona, usually covered with
sessile glandular hairs. Pappus usually barbate, less often scabrous or
plumose, deciduous in almost half the species, with pappogenus and
nectary; achenes in such cases with pappophorus; weakly connate into
ringlet in other half of species, deciduous, some bristles remain attached
in such cases, pappophorus not developed, and stem always poorly
developed. Leaves usually pinnatipartite, pinnate or lyrate, often
decurrent.

Type of section: J. chaetocarpa Ldb.

=

. Pappus entirely shedding in immature achenes ...........0....0..00.0.. Z:
+ Pappus not deciduous, bristles connate ................ccccceeeeeeeees 14.
2. Plants stemless or with short stem; lower leaves as long as or

exceeding capitula, sometimes slightly below them; leaves always

white-tomentose beneath, usually divided...........0...0c0cccceeeee 3:
+ Stem always distinctly exceeding basal leaves............0.0ccc08 a:
3. All involucral bracts appressed (Isfairam River Valley)................
Heusen de lomendi | sei iui) rahe Orcs Lael UREN ELT) 138. J. densisquamea [Ijin

4. Plants stemless or with scarcely developed stem; outer favs
bracts long and arcuately bent (alpine and subalpine regions of

Tien Shan and Pamiro-Alai Region) .............. 136. J. algida [lyin
+ Plants always with short developed stem; outer involucral bracts
short and arcuately bent (Darvaz) ........... 137. J. darwasica Ijin
Sz deavess: decumentene witht) 6h De LOI), NS tol Hae LLM 6.
tH? (Leavesenotydecumrenitwy.. eee Onan, CERES A Tal RRREM oo 8.
6. Leaves undivided and entire, graminaceous (Kyzyl-Dzhar Region
ofiieasternuiergana} dx. 2a a 141. J. poacea [lyin
+ ssleaves:notiundivided and! motventire 282.02... oe Tb

7. Leaves large, oblong-oval or oval, lyrate, long-decurrent; outer
involucral bracts recurved; achenes more or less smooth and not
plandulatiatiniy.mecewbale on . oe 142. J. baldshuanica Winkl.

+ Leaves narrow, oblong to linear, occasionally undivided, usually

sparsely narrow-lobate or toothed; lower cauline leaves short-
decurrent. Outer and middle involucral bracts short-uncately
recurved. Achenes densely glandular (vicinity of Issyk-Kul’) ......
eal. det, ee ameter. 132. J. tianschanica Rgl. and Schn.
d-Ally mivolucralebractsappressedn s4). AG! Se ROet se 9.
+_sOuteranvolucralibractsrrecunved’:.24 2). Meee Lae A ad 12?

oo

679

11.

12.

14.

15)

16.

Wie

67]

. Pappus bristles plumose; outer and middle involucral bracts

imbricate, gradually reduced downward........ 130. J. lipskyi [ljin
Pappusebristlesybanbate sc: 04 eee. ooo 25 ih etadal.. ioalds. UES 10.

- Semishrubs; leaves cord-like, pappus bristles short-barbate and very

LSTA Sept Ft, ps. nett ou Ba es rab advenits aipltial a alee denne, Laity:
Leaves flat, pinnatisect, strongly arachnoid-hairy, leaf segments
somewhat sinuate; pappus bristles sparsely barbate (from
Muyunkums to almost Ust-Urt) ................ 126. J. multiloba [jin
Morphologically similar to J. suffruticosa. Leaves simple, undivided
(castemmkanataw) esis tii. aeped 145. J. ezilikinoana [jin
Leaves with distant long segments (chalk beds of Aktyubinsk
Region )) eal a. A cee ed 146. J. fedtschenkoana [Ijin
Stem almost leafless, with capitula inclined on one side; leaves
filiform; involucral bracts lax, with fine, slightly recurved, cusp;
achenes very densely covered with sessile glandular hairs. ‘Bolshie’

Greater | MBalkhans, ..5.25..2ees acdsee Balk 144. J. spissa [lyin
Stem uniformly leafy, flat; achenes sparsely covered with fine
plan alana ness: casement - Leneaien yeaa ol) haul she 13:

. Basal and lower cauline leaves lyrately divided, others undivided

and entire, all leaves very hard; outer and middle involucral bracts
arcuately recurved; achenes covered with longitudinal rows of
tubercles (‘Malaya’ [Lesser] Almaatinka) ...............ccccccescccseeeseeees
S. peesperaven iene R MORE oa ce nciectsesyenssnese 134. J. almaatensis Iljin
All leaves pinnatisect, especially white arachnoid-hairy beneath,
soft; outer and middle involucral bracts with long, arcuate, cusps;
achenes entirely covered with erect, somewhat stiff, appressed, long,
bristlesyatisine; fromm tubercles) (Zaisan)ixi2iu.. anche’. t. tas
pein 4 0 eysemepeus. Shroee lds, yeni hes fil | 122. J. chaetocarpa Ldb.
Outer involcural bracts arcuately recurved ..............00.c:cccccce IS.
All involucral bracts appressed or somewhat lax..............0.00.. 18.
Stem flat, usually solitary, short; leaves obtusely pinnatilobate, not
sinuate; achenes toothed only above, farinose-glandular

(Dzhungarian Alatau)............... 127. J. dshungaria (Rubtz.) [jin
Stem more or less branched, at least in lower part; often flexuous
bein ene RMN BH liek. ean Wns, eae, Muted a be fe Gs

Leaves sparsely pinnatilobate, with broad, triangular, obtuse or
subobtuse lobes, incised, large toothed, with obtuse basal auricles,
not sinuate; achenes also farinose (Balkhash Region)...................
WE es Oa vlan et ed eee bl ars oo, 4 124. J. suidunensis (Winkl.) Korsh.

eaviesppinnatisectasinuater.d34.0s./ TON ad el caladinek LZ
Outer involucral bracts arcuately bent (Ust-Urt and northern
Turkineiia) seh eh. Wak ert WC betes 123. J. persimilis [jin

Outer involucral bracts filiform, initially appressed, at anthesis and

680

672

18.

Jyh

especially at fruiting recurved to sides, sometimes also
decurved (foothills of Dzhungarian Alatau)..............ceeeeeeeeeeeeee
staasadtosshbdecshd RBA edh ieee 125. J. margalensis [ljin
Pappus bristles plumose; capitula small, 10-12 mm long and 8-10
mm wide, achenes densely glandular (steppes of Balkhash Region)
OR MiP RU RON RTS, VARTO MA. 131. J. adenocarpa Schrenk

Pappus bristles barbate or scabrous-barbate .........00.. eee 19.
. Stem simple, significantly exceeding leaf rosette; leaves undivided,
pinnatisect, or less often lyrate.............00.... 135. J. lanipes Rupr.
Plants different in appearance, without scapelike stem ........... 20.
. Leaves longer than or almost as long as stem.................0 PAN
Stem always longer than basal leaves 00.0.0... cee eeeeeeeeeeeeeeeeeeeeee OR

. Plants forming pad of sessile capitula on soil, with oblong,

pinnatipartite or pinnatidentate leaves in between; rhizomes vertical,
short and approximate, arising from tap root (Tien Shan, near the
Mouth of KyZyl-su)\-heeicttsceccveseseceesesicess 140. J. grumosa Ijin
Stem 8-20 cm high, simple, as long as, slightly longer or slightly
shorter than leaves; leaves oblong-oval or oblong-lanceolate,
gradually narrowed into petiole (Pamir) ..... 139. J. nivea Winkl.
Strongly branched semishrub, with spatulate, undivided, or toothed
or oblong spatulate lobes; leaves petiolate, whitish due to
continuous floccose pubescence (chalk beds of western Kazakhstan)
ERR UIES oo bates. squscensveaaestsencneests 143. J. kirghisorum Janisch
Herbaceous plants, sometimes with developed caudex ............ ZS):

. Tall plants with undivided oblong-lanceolate leaves, acuminate and

gradually narrowed into petiole (mountain meadows of Tien Shan)
betes ste Susevesacenaseadihes saateen eI Ret eee toes 133. J. foliosa (Ijin) Ijin
IVEAWES? PUI atiSe Cli sac..ccssecexaes ceisccessazaevecsn dette congersaeesacenencatesatatao 24.

. Stem strongly branched, with long branches; capitula relatively

small; involucre 10-15 mm long; corolla 10-15 mm long (sands
of Muyunkuma) ..............cceeeeee 129. J. krascheninnikovii [ljin
Stem simple or with few short branches; capitula cylindrical or
cyathiform at anthesis, later almost cupuliform; involucre 18-20
mm long; corolla 15-18 mm long (Sultan-Uiz-Dag).....................
sultanaeavatdss Seamteg basocpengneavangsabasegestetevectesazease 128. J. longicorollaris [jin

122. J. chaetocarpa Ldb. FI. Ross. II, 2 (1845-1846) 756; Boiss.

Fl. or. HI, 577 excl. spec. alt.; Kryl. Fl. Zap. Sib. XI, 2933: Korsh. in
Izv. Akad. Nauk 1, 2, 127; O. and B. Fedtsch. Rast. Turk. IV, 302,
spec. semipal.; Iljin in Tr. Turk. Nauchn. Obshch. II, 21.—Serratula
chaetocarpa Ldb. FI. alt. IV (1833) 42—J. chaetocarpa var. typica
Herder in Bull. Soc. Nat. Mosc. XL (1867) 167.—Ie.: Ldb. Ic. pl. fl.
Ross. V, tab. 490.

673

Perennial. Plants 20-30 cm high, grayish due to arachnoid-hairy
pubescence. Stems usually few, simple or somewhat branched, with few
leaves. Leaves mainly crowded at base, pinnatisect into oblong-oval or
often oblong, subobtuse, and entire lobes or lobes with few short teeth,
usually involute and sinuate, grayish on both sides, especially beneath,
due to fine arachnoid-hairy pubescence, petiolate; cauline leaves few,
smaller, linear, usually undivided, sessile; only lowermost leaves like
basal. Capitula solitary, relatively large. Involucre 18-20 mm long and
13-20 mm wide, cupuliform, somewhat concave at base, green;
involucral bracts lanceolate; outer and middle bracts floccose-arachnoid-
hairy, aristate and deflexed from middle; inner bracts glabrous, erect,
acuminate. Receptacle densely squamose; scales one-fifth to one-third
as long as involucre. Corolla pink-purple, with about 16-17 mm long
lobes of limb. Pappus deciduous when mature, snow-white, 7.5-10 mm
long, bristles barbate, with barbules slightly longer than thickness of
bristles. Achenes 4.5—6 mm long and about 1.5—2 mm thick, densely
covered with appressed, fine, fiber-like hairs above, modified into
tubercles, with patellate, somewhat thick, pappophorus and nectary at
apex. Flowering V—VI; fruiting VII. (Plate LI, Fig. 3.)

Sandy-stony steppes, stony slopes, and saline desert steppes.—Soviet
Central Asia: Balkhash Region. General distribution: Western Mongolia,
Sinkiang. Described from the Zaisan Region. Type in Leningrad.

123. J. persimilis Iljin in Bot. Mat. Gerb. Bot. Sada, V (1924)
164; in Tr. Turk. Nauchn. Obshch. II, 22; Fl. Turkm. VII, 225.

Perennial. Plants 10—30(40) cm high, floccose-arachnoid-hairy. Stem
simple or with few branches, less often strongly branched with long
branches (f. ramosa Iljin), but usually leafy in lower part. Leaves grayish-
green or even whitish due to tomentum, usually concolorous or somewhat
dark green above, pinnatisect into linear or oblong-linear, subobtuse lobes,
distinctly narrowly sinuate, occasionally almost undivided (f. indivisa
Ijin); basal and lower cauline leaves petiolate, upper sessile, linear,
usually toothed or even undivided along margin. Capitula solitary.
Involucre cupuliform, sometimes, especially at anthesis, narrowly
cupuliform, 15-20 mm long and 10-18 mm wide. roundish at base or
with slight curve, floccose-arachnoid-hairy with lanceolate and linearly
lanceolate, acuminate bracts; outer and middle bracts with recurved
apices; inner bracts appressed. Receptacle densely squamose; scales short,
one-fifth to one-fourth as long as involucre. Corolla pinkish-purple, 15—
19 mm long, narrowed over 5—7 mm, with 3.5-5 mm long lobes of
limb. Pappus persistent, 7-10 mm long, snow-white, barbate, barbules
dense, almost as long as thickness of bristles. Achenes 4.5-6 mm long,
less often up to 7 mm (f. macrachenia I\jin), obpyramidal, with

682

674

longitudinal row of tubercles, connate below, on upper side modified
into acute tubercles, with petallate pappogenus at apex, and long nectary
in center. Flowering V—VI; fruiting VI. (Plate LI, Fig. 1.)

On chalks, limestones, sandstones, less often sandy steppes, loessy
and gravelly loams.—Soviet Central Asia: Aralo-Caspian Region (Ust-
Urt); Kara-Kum (far west). Described from the Karatau Mountains on
the Mangyshlak Peninsula, the city of Aktau and Akmysh. Type in
Leningrad.

Note. In the south the species reaches Krasnovodsk. At the northern
border of Ust-Urt and Lake Sam and Asmantai it produces forms
intermediate to J. multiloba \ljin. This species is also known from the
northern shores of the Aral Sea.

124. J. suidunensis (Winkl.) Korsh. in Bull. Acad. Sc. Pétersb. V,
1 (1894) 128; O. and B. Fedtsch. Perech. Rast. Turk. IV, 301; [lin in
Tr. Turk. Nauchn. Obshch. II, 23—J. pollichii var. suidunensis Wink.
in herb. and in Korsh. op. cit.

Perennial. Plants 25-70 cm high. Stem branched, mainly in lower
half, with long branches, floccose-arachnoid-hairy, leafy, predominantly
in lower half. Leaves pinnatilobate or pinnatipartite, with broadly
triangular or oblong, obtuse or subacute lobes, finely arachnoid hairy-
pubescent above, grayish beneath due to fine tomentum; basal and lower
cauline leaves gradually narrowed into petiole; upper leaves undivided,
sessile, usually with obtuse auricles. Capitula solitary. Involucre 17—25
mm long and 15-20 mm wide, mildly bent at base, floccose-arachnoid-
hairy in lower part at some places, glabrous in upper half, with light
purple tinge; involucral bracts lanceolate, acuminate, outer bracts
recurved, inner finely acuminate, straight. Receptacle densely squamose;
scales one-fourth to one-third as long as involucre. Capitula pinkish-
purple, covered with glandular hairs, 15—19 mm long, narrowed over
5.5—7 mm, with 4—5.5 mm long lobes of limb. Pappus persistent, snow-
white, 8-10 mm long, barbate, barbules as long as thickness of bristles.
Achenes 5—6 mm long, covered with discontinuous parallel ribs, modified
into acute tubercles above; with toothed edges at tip; pappophorus two-
tiered; lower tier of small membranous ring fused with base of pappus
bristles, and upper tier petallately thickened, enclosing nectary. Flowering
V; fruiting VI.

Hummocky sands, saxaul [Haloxylon ammodendron|] forests.—Soviet
Central Asia: Balkhash Region (eastern part). General distribution:
Sinkiang. Described from Suidun in Kuldzha Region. Type in Leningrad.

125. J. margalensis [jin in Tr. Turk. Nauchn. Obshch. 2 (1925)
22, Plate IV.

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675

Perennial. Plants up to 40 cm high, floccose-arachnoid-hairy. Stem
erect, usually branched, less often simple, leafless in upper -half. Basal
leaves numerous, like lower cauline leaves, petiolate, pinnatisect into
oblong or oblong-linear, subobtuse sinuate lobes, middle leaves sessile,
uppermost undivided, linear-filiform, all leaves thin, sparsely tomentose
beneath, green above, floccose-arachnoid-hairy at some places. Capitula
solitary. Involucre cupuliform, depressed at base, 12-15 mm long at’
fruiting and up to 20 mm wide, glabrous, green, only slightly floccose-
arachnoid-hairy at base; involucral bracts lanceolate or linearly lanceolate,
acuminate; outer and middle bracts with recurved apices; inner bracts
appressed. Receptacle densely squamose; scales relatively short, up to
one-third as long as involucre or slightly longer, but not reaching its
half. Corolla pink-purple, 14-15 mm long, narrowed over 5—6 mm, with
3-4 mm long lobes of limb. Pappus persistent, snow-white, 10-12 mm
long, barbate, with barbules two to three times as long as thickness of
bristles. Achenes 5—6 mm long, oblong, obpyramidal, with slightly
projecting discontinuous ridges, modified at apex into acute tubercles,
covered with fine sessile glandular hairs, and with border of toothed
corona above along edges, with two-tiered pappophorus inside; outer
tier lobate, small-toothed, compactly fused, and fleshy, olive-brown, inner
tier appressed to entirely membranous pappus base; nectary very large,
columnar, membranous, twisted at apex, surrounded by the fleshy part
of pappophorus. Flowering V—VI; fruiting VII—VIII.

Sandy steppes, stony semideserts—Soviet Central Asia: Balkhash
Region. Endemic. Described from the Margaly area near the Ili River.
Type in Leningrad.

Note. The species was named J. nargalensis I\jin mistakenly in place
of J. margalensis, as this species is named after the Margaly area.

126. J. multiloba [jin in Bot. Mat. Gerb. Glavn. Bot. Sada, V, 11—
12 (1924) 2; in Tr. Turk. Nauchn. Obshch. 2, 22.

Perennial. Plants 15-40 cm high, with branched, floccose-arachnoid-
hairy stems. Leaves oblong, pinnatisect, grayish arachnoid-hairy on both
sides, especially beneath; segments oblong, with uniformly broad rachis,
and like latter, slightly involute and somewhat sinuate; basal leaves
numerous, more or less petiolate; upper cauline leaves sessile. Capitula
on long peduncles, with involucre at anthesis up to. 15—20 mm long and
10-17 mm wide. Involucral bracts floccose-arachnoid-hairy, less often
glabrous, and, besides, very distinctly appressed-hairy; outer bracts ovate
or oblong-ovate, middle and inner bracts linear, all middle bracts
acuminate. Corolla pink-purple, 14-16 mm long, narrowed over 5 mm
and with 4 mm long lobes of limb. Pappus entirely persistent, snow-
white, barbate, with barbules almost as long as thickness of bristles.

684

676

Achenes 5—6 mm long and 1.5—2 mm thick, with small-toothed border,
edges tuberculate, modified in upper part into short appressed spinules, _
on lower side connate into discontinuous ridges, with fleshy, broadly
cupuliform pappophorus at apex enclosing nectary.

On chalk slopes, limestone deposits, sands, sandy and wormwood-
covered steppes.—Soviet Central Asia: Aralo-Caspian Region, Balkhash
Region; Tien Shan (western); Kyzyl-Kum (Sultan-Uiz-Dag). Endemic.
Described from Bil-Aran Mountain, a short distance from the northern
coast of the Aral Sea. Type in Leningrad.

Subsp. aulieatensis Iljin comb. n—J. multiloba var. aulieatensis
Iljin in Bot. Mat. Gerb. Bot. Sada, V, 11-12 (1927) 3; Iljin in Tr.
Turk. Nauchn. Obshch. II, 22.—Mainly grows in the Dzhambul Region,
preferring stony and gravelly places; differs by the achenes that
have distinctly white-cartilaginous peripheral ribs, with dome-
shaped pappophorus at the tip ending with a nectary, an entirely
deciduous pappus, and the pappus hairs that are connate into ring (an
sp. nov.?).

127. J. dshungarica (Rubtz.) Iljin comb. n. hoc loco.—J.
chaetocarpa Ldb. subsp. dshungarica Rubtz. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, VIII, 4-9 (1940) 65.

Perennial. Plants usually 10-25 cm high, with rather dense caudex
covered with fine scales of remains of petiole bases, with woolly globules
in axils. Stem solitary, erect, simple, floccose-arachnoid-hairy, striate
covered with scattered sessile glandular hairs. Basal leaves petiolate,
with larger terminal lobe; lateral lobes three to seven, ovate, roundish
or oblong, obtuse, usually entire, simple and sessile, absent in upper
one-third part; all leaves grayish-green above with fine arachnoid-hairy
coating, weakly rugose, with scattered, fine, sessile glandular hairs, white-
tomentose beneath. Capitula with many florets, 15-20 mm wide.
Involucre broadly cupuliform, about 15 mm long; outer and usually
middle involucral bracts green or pale green, usually floccose-arachnoid-
hairy with sparse sessile glandular hairs, one- to three-veined, slightly
bent at tip; inner bracts oblong-linear, gradually acuminate, bright purple,
usually with three veins, finely ciliate and involute, on outer side
arachnoid-hairy pubescent and with scattered sessile glandular hairs, 14—
16 mm long, narrowed over 5.5-7 mm, with 4-6 mm long lobes of
limb. Pappus about 10-12 mm long, persistent, pappus bristles weakly
connate into ringlet, readily shedding, barbate, with barbules slightly
longer than thickness of bristles, usually approximate, becoming longer
toward upper side, two to three times as long as thickness of bristles,
structure at base of pappus with five-toothed, small, membranous nectary
and membranous border separating from flat-dome-shaped pappophorus.

677

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sy
4

<< a =i =<
5

—————

Y>.,
\ZA@
=

WES

Ya Se
MA A

yh
~ C= Y, . S\N

Plate L.
1—Jurinea spissa Mjin; 2—J. abramovii Rgl. and Schm.

685

686

678

Achenes about 5 mm long, grayish or grayish-olive, matty, with corona
along upper edge having approximate teeth on pale ribs discontinuous
in upper part, and with short cusps, edges with scarcely projecting,
parallel, discontinuous ribs, entirely covered with fine sessile glandular
hairs, less often light-colored, usually matty, almost farinose. Flowering
V-VI; fruiting VI-VIII.

Loess loams on the slopes of the Dzhungarian atate —Soviet
Central Asia: Dzhungarian Alatau. Endemic. Described from the
Borokhudzhir River Valley from the Ku area. Holotype in Alma-Ata;
isotype in Leningrad.

128. J. longicoroliaris (Iljin) Iljin comb. n—J. multiloba Tin var.
longicorolla \ljin in Tr. Turk. Nauchn. Obshch. 2 (1925) 22.—J/.
chaetocarpa auct. fl. As. Med.

Perennial. Plants 10—25(30) cm high. Stems solitary or usually few,
simple or short-branched, longitudinally striate, distinctly floccose-
arachnoid-hairy. Leaves oblong; basal leaves numerous, with petioles
shorter than lamina, and woolly globules in axil, like lower and middle
cauline leaves, pinnatisect or pinnatipartite, less often lobate or almost
undivided, with oblong, oval or oblong-ovate segments and lobes, almost
obtuse, less often subacute, segments obtusely-toothed, usually slightly
sinuate, fused with narrow wing of rachis; uppermost leaves sessile,
toothed or almost entire; all leaves soft due to pubescence grayish-green
above due to arachnoid-hairy pubescence, whitish beneath due to more
dense arachnoid-hairy tomentum. Capitula terminal on main stem and
branches, solitary, with many florets, (10)12-15 mm wide. Involucre
entirely arachnoid-hairy, floccose-arachnoid-hairy in lower part,
poculiform at anthesis, 18-20 mm long; all bracts appressed, usually
three-veined, with short cusp, later slightly recurved; innermost bracts
oblong-lanceolate, prominent, gradually almost acuminately aristate, with
sessile glandular hairs on inner side. Corolla 15-18 mm long, light pink,
with sessile glandular hairs, gradually narrowed over 5—6 mm, with 3.5—
4 mm long lobes of limb; apical appendages of anthers 3-3.5 mm long.
Pappus persistent, snow-white, together with two prominent, 10-12 mm
long, bristles; bristles short-barbate, barbules sparse, readily shedding.
Achenes ferruginous, 5-6 mm long, with low, toothed-prickly, corona
above along edges, with poculiform thickened pappophorus inside,
enclosing membranous five-lobed nectary on short stalk; achenes with
tubercles along edge, modified into spinules above, gradually obliterated
below. Flowering V; fruiting V—VI. (Plate LI, Fig. 2.)

Stony mountain slopes, rocks, pebbly deserts——Soviet Central Asia:
Kyzyl-Kum (Karakalpakia). Endemic of the Sultan-Uiz-Dag Range.
Described from the Sultan-Uiz-Dag Range.

687

679

Note. It is impossible to retain as the type the now fragmentary
specimen from the collections of M.G. Popov from which this species
was described as var. longicorolla Iljin, and I am selecting
Muravlyanskii’s specimen as the holotype.

129. J. krascheninnikovii Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada,
V, 11-12 (1924) 165, in Tr. Turk. Nauchn. Obshch. I, 23.

Perennial. Plants 15—45 cm high, entirely grayish-green due to sparse
arachnoid-hairy tomentum, especially leaves beneath, forming dense turf
of short caudices, woolly-tomentose in axils of old petioles. Stems few,
branched. Leaves oblong, pinnatisect, with oblong or linear segments,
narrowly involute and usually finely sinuate, almost as wide as middle
undivided part; basal leaves numerous, like lower cauline leaves petiolate;
upper leaves sessile, linear, undivided, gradually disappearing above.
Capitula numerous, smaller, solitary, about 20 mm long including florets.
Involucre narrowly cupuliform, 10-15 mm long and 5-12 mm long
bracts, slightly floccose-arachnoid-hairy, roundish below. Receptacle
densely squamose; scales one-third to two-fifths as long as involucre.
Corolla small, pink-purple, covered with glandular hairs, 10-15 mm
long, narrowed over 3.5—6 mm, with 2.5—-4 mm long lobes of limb.
Pappus persistent, snow-white, 8-10 mm long, barbate, with dense
barbules almost as long as thickness of bristles. Achenes 6-7 mm long,
oblong, not glandular, with elongate discontinuous ridges, modified above
into spinulose tubercles, with broadly cupuliform pappophorus above
corona, with oblong, narrow, five-toothed nectary in center, compactly
surrounding fleshy clavate part. Flowering VI—VII; fruiting VI-IX.

Sandy steppes, alluvial sands, sandstone outcrops.—Soviet Central
Asia: Balkhash Region, Aralo-Caspian Region (Far East). Endemic.
Described from the lower reaches of the Sarysu River in Muyunkum.
Type in Leningrad.

Note. | described var. arenicola Ijin (= J. arenicola Wjin l.c.) from
the Muyunkum sands, growing on hummocky sands, as a sharply marked
variety. It could be recognized as an entirely separate, young species,
distinguished by having a more dense tomentum, especially on the
involucre, more elongate and loosely disposed caudices, and more distant
leaf segments, the leaves sometimes being almost undivided. It was
described from the vicinity of Dzheras-Kuduk. The pappus and
pappophorus are like those of the type specimen, but the achenes are
about 1.5 mm thick, with glandular hairs.

130. J. lipskyi Iljin in Tr. Turk. Nauchn. Obshch. 2 (1925) 23,
Plate V.

Perennial. Plants up to 50 cm high, with erect, long-branched or
almost unbranched, weakly floccose-arachnoid-hairy stems. Basal leaves

688

680

numerous, like lower cauline leaves petiolate, pinnatisect into oblong,
somewhat sinuate, somewhat narrowly involute, and usually subobtuse
lobes; middle leaves less divided, upper leaves linear or almost filiform,
undivided; all leaves somewhat hard, finely gray-tomentose beneath,
green, and glabrous or weakly arachnoid-hairy pubescent. Capitula
solitary. Involucre cupuliform, with florets 12-16 mm long and 12-17
mm wide, depressed at base, almost glabrous or slightly arachnoid-hairy
below, with appressed, lanceolate and linearly lanceolate. acuminate
bracts. Receptacle covered with long bristles up to half as long as
involucre or slightly longer. Corolla pink-purple 15-27 mm long,
narrowed over 4.5—5.5 mm, with 4-5 mm long lobes of limb, with
sessile glandular hairs. Pappus entirely deciduous, up to 10 mm long,
snow-white, plumose, with barbules three to four times as long as
thickness of bristles, bristles connate at base into ring. Achenes 6—7
mm long, gray, oblong-obpyramidal, with parallel, elongate,
discontinuous ridges, modified above into two or three acute spinulose
tips, with sessile glandular hairs in between ridges and at tips of spinules;
corona cuspidate, slightly recurved above along edges, with flat
pappophorus inside, modified into narrowly conical tip, with columnar
nectary, terminating in fine, membranous, five-toothed apical lobe.
Flowering V—VI]; fruiting VII.

Wormwood or feather-grass steppes.—Soviet Central Asia: Balkhash
Region, Dzhungaria-Tarbagatai (Dzhungarian Alatau); Tien Shan (Tekes).
Endemic. Described from the vicinity of Kopal. Type in Leningrad.

131. J. adenocarpa Schrenk, Enum. pl. nov. I (1841) 46; Ldb. FI.
Ross. II, 2, 765; Boiss. Fl. or. III, 578; Korsh. Izv. Akad. Nauk. 1, 2,
127.—J. chaetocarpa var. adenocarpa Herd. in Bull. Soc. Nat. Mosc.
XL, 167; O. and B. Fedtsch. Perech. Rast. Turk. IV, 303; Iljin in Tr.
Turk. Nauchn. Obshch. II, 23.

Perennial. Plants 15—40 cm high, weakly grayish-green, with densely
crowded and woolly-tomentose caudices at base, many-stemmed. Stem
branched, floccose-arachnoid-hairy. Leaves pinnatipartite or pinnatisect,
petiolate, with oblong or linear, subobtuse segments, often narrowly
sinuate, floccose-arachnoid-hairy on both sides, especially beneath; basal
leaves numerous; upper cauline leaves sessile, linear, undivided,
acuminate. Capitula more or less numerous, small, solitary. Involucre
weakly floccose-arachnoid-hairy, 10-12 mm long and 8-12 mm wide,
cupuliform, roundish at base, involucral bracts broadly lanceolate to
linearly lanceolate, all appressed, green; innermost bracts weakly purple
along margin. Receptacle densely covered with bristles, one-third to
two-fifths as long as involucre. Corolla pink-purple, covered with
glandular hairs, short, 10-14 mm long, narrowed over 3.5—6 mm, with

689

681

short, 2-3 mm long limb. Pappus persistent, snow-white, 8-10 mm long,
barbate-plumose or plumose, with barbules three to six times as long as
thickness of bristles. Achenes 4.5—6.5 mm long, oblong, with dense
sessile glandular hairs, longitudinal discontinuous ridges and apically
appressed cusps, modified into sharp spiniform tubercles above.
Flowering V—VIII; fruiting VII-IX.

Hummocky sands, approaching sandy steppes, less often on gravelly
slopes.—Soviet Central Asia: Balkhash Region, Dzhungaria-Tarbagatai
(Dzhungarian Alatau, rare). Endemic. Described from Balkhash Region,
Ayaguz Region, near Arganaty Mountains.

Note. S.A. Nikitin collected sharply differing specimens of this
species under two numbers from the lower reaches of the Ili River on
hummocky sands in the Randa-Tugai area. I have referred them to var.
latifolia \\jin var. nov.

132. J. tianschanica Rgl. and Schm. in Tr. Peterb. Bot. Sada, VI
(1880) 322; Trautv. ibid., VIII, 2, 513; O. and B. Fedtsch. Perech. Rast.
Turk. IV, 301; Ijin in Tr. Turk. Nauchn. Obshch. II, 24.

Perennial. Plants 15-50 cm high. Stems few, usually erect, angular
in lower half, cylindrical above, branched, with branches straight, upward
spreading in lower half or one-third, ribbed-barbate, grayish arachnoid-
hairy, less often almost green (f. glabrescens Ijin) and covered with
fine, scattered, sessile glandular hairs, with many leaves, except below
capitulum; basal and lowermost cauline leaves narrowed gradually in a
short petiole; cauline leaves sessile, very short-decurrent, with entire
wings, usually narrow-oblong, often appressed, almost undivided and
entire, or usually sparsely sinuate-toothed or broadly sinuate-toothed,
with obtuse apex, sometimes pinnatilobate or pinnatipartite, with short
cuspidate upcurved lobes, narrowly involute, usually green, weakly rugose
above, with fine sessile glandular hairs and depressed midrib, whitish-
grayish beneath due to more or less dense arachnoid-hairy tomentum,
occasionally concolorous, canescent (f. canescens Ijin); uppermost leaves
filiform. Capitula with many florets. Involucre cupuliform, 12-15 mm
long, imbricate, five-rowed, arachnoid-hairy in lower half; outer
involucral bracts somewhat pale, with prominent midrib and dark green
weakly fleshy, subobtuse or short-cuspidate, uncately bent apex,
sometimes with purple tinge; all other bracts entirely appressed; middle
bracts lanceolate, acuminate, dark purple, with three to five prominent
veins, scarcely visible ciliolate along margin, like innermost bracts,
densely covered on outer surface with large sessile glandular hairs;
innermost bracts acuminate with brownish cusp. Receptacle squamose;
scales acuminate, one-fifth to one-fourth as long as involucre. Corolla
raspberry-red, 13-17 mm long, narrowed over 4.5—6 mm, with 3-4 mm

690

682

long lobes of limb, covered with fine sessile glandular hairs; apical
appendages of anthers abruptly narrowed into thin, 2.5—3 mm long, cusp,
basal appendages setaceous, divided, 2.5 mm long; filaments almost
smooth; style glabrous, usually purple; stigma about 1.5 mm long,
divided, almost up to base into divergent papillate branches, with tuft
of short erect bristles at base. Pappus always smoky, slightly longer
than achene, excluding two or three prominent bristles, arranged in few
rows, shorter toward outside and connate into ring, easily shedding on
maturation of achenes, bristles short-barbate, with barbules equal to or
two times as long as thickness of bristles, oblique and dense. Achenes
3—4(5) mm long, obpyramidal, four-angled, with toothed corona at tip,
and longitudinal discontinuous parallel rows of tubercles along edges
below corona modified into one or two, or three toothed tip, densely
covered, especially above, with lustrous sessile glandular hairs, light
brownish, with flask-shaped pappophorus at apex terminating in stalked
nectary exserted from pappus ring. Flowering VII—VIII; fruiting VIII—
IX.

Stony slopes.—Soviet Central Asia: Tien Shan (center), Pamiro-Alai.
Endemic(?). Described from the vicinity of Lake Issyk-Kul. Type in
Leningrad.

133. J. foliosa (Iljin) Ijin comb. nov.—J/. lanipes subsp. foliosa
Ijin in Izv. Glavn. Bot. Sada, XXVII, 1 (1928) 83.
Perennial. Plants 30-60 cm high. Stems usually few, erect, simple,

weakly floccose-arachnoid-hairy, especially above, leafy up to middle

or slightly below. Leaves lanceolate, petiolate, petiole longer in basal
and lower cauline leaves, acuminate, undivided and entire, less often
weakly lyrate, with narrow, acuminate terminal lobe and few, small,
distant lateral lobes, green and subglabrous above, grayish-green beneath
due to very fine arachnoid-tomentum. Capitula solitary. Involucre 15—
25 mm wide, glabrous or subglabrous; involucral bracts almost equal,
linearly lanceolate, acuminate; middle and outer bracts weakly recurved-
sinuate, herbaceous-green. Corolla 15-20 mm long, pink-purple, narrowed
over 5—7 mm, with 4—5 mm long lobes of limb. Pappus entirely persistent
or less often deciduous, snow-white, barbate, with barbules almost as
long as or slightly shorter than thickness of bristles; bristles weakly
connate in a ring, partly persistent on achene. Achenes 6-7 mm long,
toothed-fimbriate, tuberculate-spiny in upper half, with tubercles in
longitudinal rows and connate at base, with broadly cupuliform, sinuate,
whitish, pappophorus at apex inside corona, somewhat dome-shaped in
center and terminating with small nectary that sometimes penetrates
through pappus ring and then fused with ringlet by membranous base.
Flowering VI-VII; fruiting VI-VIII.

69

—_

683

Steppe, meadow-steppe and stony mountain'slopes—Soviet Central
Asia: Tien Shan (eastern). Endemic. Described from Alma-Ata Region,
from the Aleksandrovskii Range, from Shekul Mountain. Type in
Leningrad.

134. J. almaatensis [jin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, VI, 3 (1937) 58.

Perennial. Plants 25-40 cm high, many-stemmed. Stems arising from
caudices covered with remains of petioles of basal leaves, branched
only in lower half, leafless in upper half. Leaves somewhat hard, bright
green and glabrous above, greenish beneath due to somewhat arachnoid-
hairy pubescence, more dense in lower leaves; basal and lower cauline
leaves narrowed into more or less long petiole, lyrate or almost
pinnatisect, with oblong or lanceolate terminal lobe and one to three
pairs of oblong, remote lateral lobes, others entire or with one or two
teeth in lower part, oblong; all leaves narrowed toward base, uppermost
linear. -Capitula solitary. Involucre cupuliform, about 15 mm long and
15-18 mm wide, almost glabrous; outer and middle bracts recurved.
Receptacle squamose; scales almost half as long as involucre. Corolla
about 18-20 mm long, narrowed over 6—7 mm, with 4-5 mm long
limb. Pappus deciduous, barbate, with barbules almost one and one-half
times as long as thickness of bristles; bristles connate into ring at base,
surrounding nectary at base. Achenes 5—6 mm long, covered with sharp
scales (tubercles), especially along ribs, in longitudinal rows, connate
at base; pappophorus narrowly conical, modified at apex into stalk
terminating with nectary.

Rocks.—Soviet ‘Central Asia: Tien Shan. Endemic. Described from
the Trans-Ili Alatau, from the Malaya Almaatinka River. Type in
Leningrad.

135. J. lanipes Rupr. ex Osten-Sacken and Rupr. in Mém. Acad.
Pétersb. sér. VII, XIV, 4 (1869) 58; Korsh. in Bull. Acad. Sc. Pétersb.
V, 1, 126; O. and B. Fedtsch. Perech. Rast. Turk. IV, 301; Trautv. in
Tr. Peterb. Bot. Sada, VIII, 1, 310; Iljin in Tr. Turk. Nauchn. Obshch.
iy 28.

Perennial. Plants 15-60 cm high, with solitary or few erect and
simple branches (exception, single branch at extreme base), floccose-
arachnoid-hairy, almost leafless stems. Leaves usually green above, also
weakly arachnoid-hairy, with depressed veins, sparsely tomentose
beneath, sometimes almost entirely tomentose when young, weakly
sinuate and very narrowly involute, pinnatisect into oblong or oblongly
linear obtuse lobes (f. pinnatipartite Korsh.) or lyrate (sometimes mixed
with undivided leaves) below, short-petiolate; uppermost leaf usually

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684

filiform; basal leaves usually numerous. Capitula solitary, relatively large.
Involucre cupuliform, with depressed base, 15—20 mm long and wide,
green or floccose-arachnoid-hairy with long and finely acuminate,
lanceolately linear, appressed bracts; innermost bracts often with purple
tinge and revolute in upper part. Receptacle densely covered with bristles
about one-third as long as involure. Corolla purple, covered with sessile
glandular hairs, 15—24 mm long, narrowed over 4-9. mm, with 3-6 mm
long lobes of limb, less often shedding entirely, usually with partly
persistent bristles below pappophorus. Pappus white, 8-10 mm long,
densely barbate, with barbules approximately as long as thickness of
bristles. Achenes 5-6 mm long, oblong-obpyramidal, with parallel
discontinuous ridges modified into acute tubercles above, with toothed
corona above, enclosing flask-shaped pappophorus with neck, exserted
from opening of ring and fused with it, encircling tip, and terminating
with five-lobed nectary. Flowering VI-VIII; fruiting IV—IX.

Alpine and subalpine forest meadows, loessy mounds, stony and
rocky habitats—Soviet Central Asia: Tien Shan, Pamiro-Alai Region.
General distribution: Dzhungaria-Kashgaria (Sinkiang). Described from
central Tien Shan. Type in Leningrad.

Note. Korshinsky distinguished the following varieties with
corresponding synonyms: var. typica Korsh. l.c. (J. arachnoidea x var.
lyrata Winkl. in herb. and Korsh. I.c.).—Leaves lyrate, divided,
sometimes mixed with undivided leaves; var. pinnatipartita Korsh. |.c.
(J. chaetocarpa var. macrantha Herder in Rgl. and Herd. Bull. Soc.
Nat. Mosc. XL (1867) 167).—Leaves pinnatisect.

136. J. algida Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada, V, 11-12
(1924) 170; Iljin in Tr. Turk. Nauchn. Obshch. II, 26.—J. depressa O.
and B. Fedtsch. Perech. Rast. Turk. IV, 304, non al.

Perennial. Plants 2-10 cm high, mostly stemless or with very short
stems. Leaves oblong or lanceolate, obtuse or subacute, undivided (f.
integrifolia \ljin), lyrate (f. lyrata Iljin) or pinnatipartite (f. pinnatifolia
Iljin), less often lobed, entire, finely arachnoid-hairy above, gray or
grayish-whitish beneath due to tomentum, gradually narrowed: into
petiole. Capitula usually sessile. Involucre 15-20 mm long and 12-20
mm wide, green, often with purple tinge above; outer and middle
involucral bracts not strongly arachnoid-hairy, distinctly recurved in upper
herbaceous, half or two-thirds, part; innermost bracts almost glabrous,
appressed, acuminate, narrowly membranous-fimbriate along margin.
Receptacle squamose; scales short, connate at base, smooth, acuminate,
one-sixth to one-fifth as long as involucre. Corolla pink or pale pink-
purple, 15—20 mm long, narrowed over 6—8.5 mm, with 3.5—5 mm long
lobes of limb, covered with sessile glandular hairs. Pappus entirely

693

685

deciduous on maturation, snow-white, 13-16 mm long; pappus bristles
short-barbate, barbules distant, all bristles connate into dense ringlet,
surrounding nectary at base, fused with membranous roundish base,
dropping off on maturation. Achenes 6-7 mm long and 2 mm wide,
glabrous, smooth, lustrous, brownish, longitudinally striate, striations.
tuberculate, with toothed corona above along edges enclosing conical
pappophorus modified above columnar structure, terminating with
somewhat five-lobed nectary with recurved lobes. Flowering VII-IX;
fruiting VIII-IX.

Stony, rubbly, gravelly slopes, as well as in juniper zone, in
subalpine and alpine zone——Soviet Central Asia: Tien Shan, Pamiro-
Alai Region. Endemic. Described from Karabulak, on the basis of
collections of O. Fedtschenko (subsp. fetissowii (Winkl.) Iljin) from the
Pamiro-Alai Region, in the Karakol Region (subsp. kuschakewiczii
(Winkl.) I}jin).

~ Note. Two races can be noted: the one, indigenous to the Tien
Shan Mountain system, J. algida subsp. fetissow:i (Winkl.) Iljin (in
Tr. Turk. Nauchn. Obshch. Il (1925) 26).—Pappus bristles scabrous
or scabrous-barbate; and the other, growing in the Pamiro-Alai
Mountain system, J. algida subsp. kuschakewiczii (Winkl.) Iljin 1.c.—
Pappus barbules one and one-half to three times as long as thickness
of bristles.

137. J. darvasica Iljin in Izv. Tadzh. Bazy Akad. Nauk SSSR, 1, |
(1933) 66.

Perennial. Plants caespitose, 10-20 cm high, having caudices covered
with remains of old leaves. Stems usually numerous, arachnoid hairy-
tomentose, simple or weakly branched, erect or often ascending, slightly
exceeding leaves. Leaves lyrately divided, with large oblong-oval or
oblongly obtuse terminal lobe and one to three pairs of lateral, small,
distant lobes, matte-green and glabrous above, sparsely grayish-tomentose
beneath; basal leaves numerous, like lower cauline leaves, petiolate,
other leaves less numerous, reduced, almost undivided, sessile. Capitula
with hemispherical involucre, 15—17 mm long and wide; outer and middle
bracts lanceolate, acuminate, appressed, purple, almost glabrous. Corolla
pink, 20-22 mm long, narrowed over 8-10 mm, with about 4 mm long
lobes of limb; basal appendages of anthers setaceous, 2.5—-3 mm long.
Pappus deciduous, snow-white, barbate, with barbules two to three times
as long as thickness of bristles; bristles, connate into ring. Achenes
spinulose, toothed above, almost with bottle-shaped pappophorus in center
of corona, with five-toothed nectary at tip of column piercing through ~
ring opening.

Stony mountain slopes.—Soviet Central Asia: Pamiro-Alai Region.

694

686

Endemic. Described from Darvaz, from a place between Dashtak and
Kergovat. Type in Leningrad.

138. J. densisquamea [ljin in Bot. Mat. Gerb. Glavn. Bot. Sada, V,
11-12 (1924) 169; Ijin in Tr. Turk. Nauchn. Obshch. 2, 20.

Perennial. Plants short, 10-15 cm high, caespitose with many-headed
caudex, many-stemmed. Previous year’s stems reduced, woolly-
tomentose, densely covered with scales at base, narrowed toward apex,
woody; stems of current year simple or with few branches, slender,
with erasing arachnoid-hairy pubescence, arachnoid hairy-tomentose at
base and below capitulum. Leaves oblongly lanceolate or lanceolate,
pinnatipartite, less often pinnatisect or sometimes mixed with undivided
leaves (usually lower leaves), acute or subobtuse, gradually narrowed
toward base into petiole, broadened at base; uppermost leaves small,
linear, sessile; all leaves greenish, somewhat arachnoid-hairy and rugulose
above, gray or whitish, arachnoid hairy-tomentose beneath. Capitula
solitary, poculiform, about 20 mm long (including florets). Involucre
weakly floccose-arachnoid-hairy, mainly glabrous or subglabrous below,
usually green, up to 15 mm wide above; involucral bracts five- or six-
rowed, reduced toward outside, acuminate, obliquely membranous-
fimbriate along margin; inner bracts usually with pink-purple tinge; all
bracts with almost indistinct veins, appressed. Receptacle densely covered
with more or less short, smooth, bristles. Corolla pale-pink, covered
with scattered glandular hairs, especially in lower part, 17-18 mm long,
narrowed over 5—6 mm, with about 4 mm long limb; anther appendages
setaceous, 2—2.5 mm long; filaments smooth. Pappus snow-white,
deciduous, less often few outermost bristles persistent, barbate, about
15 mm long, with barbules two to three times as long as thickness of
bristles, dense; bristles connate into compact ring or, more precisely,
short tube enclosing urceolate nectary. Achenes 5.5—-6 mm long (not
fully mature), obpyramidal, toothed above, along edges, with
longitudinally arranged tubercles above on outer side, terminating with
very short spinules at apex, decurrent at base and in lower part of achene
modified into discontinuous, longitudinal, fused folds, absent in lower
side, with whitish conical pappophorus at apex, nectary piercing through
pappus ring. Flowering V(?)-VI; fruiting VI-VII.

Stony places.—Soviet Central Asia: Pamiro-Alai Region. Endemic.
Described from the Isfairam River Valley between the mouth of the
Surma-Tash River and Lyanger area. Type in Leningrad.

139 J. nivea Winkl. in Tr. Peterb. Bot. Sada, XI, 2 (1892) 321; O.
and B. Fedtsch. Perech. Rast. Turk. IV, 300; Ijin in Tr. Turk. Nauchn.
Obshch. II, 21.

695

687

Perennial. Plants 8-20 cm high; tap root bearing many-headed
caudex, continued into short vertical rhizomes covered with membranous,
yellow, scaly, remains of old leaves in axils and woolly globules. Stems
few, erect, simple or with isolated branched at base, cylindrical, floccose,
and whitish arachnoid-hairy, finely-sulcate. Basal and lower cauline
leaves and leaves on vegetative shoots oblong-oval, obspatulate or less
often oblong, undivided and entire, occasionally lowermost leaves with
distant, obtuse, short lobes, cuspidate, gradually narrowed into petiole,
broadened at base, rugulose, with somewhat depressed midrib, grayish-
green above due to fine tomentum, whitish beneath due to more dense
arachnoid-hairy tomentum, with prominent yellowish midrib; upper leaves
narrower, sessile, narrowed toward lower side, smaller. Capitula solitary
terminal on branches, many-flowered. Involucre cupuliform, 18-22 mm
long, floccose-arachnoid-hairy; outer bracts green, oblong, short-
cuspidate; middle bracts lanceolate, acuminate, with less prominent veins,
somewhat ciliate, with sessile glandular hairs, purple, like innermost
bracts, gradually narrowed, acuminate, broadly membranous; middle and
usually outer bracts arcuately recurved. Receptacle squamose; scales long,
very broad, acuminate. Corolla pink-purple, covered with sessile glandular
hairs, 14-16 mm long, narrowed over 6 mm, with 3-4 mm long lobes
of limb; apical appendages of anthers 2—2.25 mm long, short-acuminate,
basal appendages 2—2.5 mm long, setaceous; filaments almost smooth;
style covered with sessile glandular hairs from middle; stigma 1.5—2
mm long, bifid, with small papillae, with short erect basal bristles. Pappus
very immature, persistent, snow-white, 6-7 mm long (excluding long
bristles); bristles short-barbate, many-rowed, two or three inner bristles
almost two times as long as others. Immature achenes brownish,
obpyramidal, four-angled, tuberculate-rugose(?), glabrous or with sessile
glandular hairs, with toothed corona above along edges and flattened
pappophorus at tip. Flowering VII.

Ravines and gravelly river beds.—Soviet Central Asia: Pamiro-Alai
Region. Endemic. Described from the Kyzylsu River Valley in Kavuk
area. Type in Leningrad.

140. J. grumosa Iljin in Sched. ad Herb. Fl. As. Med. XIV-XX
(1928) 107—Exs.: H.F.A.M. No. 491.

Perennial. Plants short, densely caespitose, stemless, tap root long, ©
stems undergound, arising from root in tuft and densely covered with
long scaly remains of leaves. Leaves numerous, linear, acute or subacute,
gradually narrowed toward base, only slightly broadened at attachment,
usually remotely pinnatipartite, pinnatilobate or only toothed, less often
almost entire in upper part, narrowly involute, grayish-green above due
to sparse arachnoid-hairy pubescence, whitish-tomentose beneath.

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688

Capitula sessile or less often on short arachnoid-hairy, up to 0.5 cm
long, peduncle, solitary terminal on stem, forming dense mass on surface
of pad. Involucre cupuliform, about 10-15 mm long, glabrous, arachnoid-
hairy at base, only at isolated places, with green appressed bracts, narrow
pale-membranous-fimbriate along margin, narrowly lanceolate, acuminate.
Receptacle densely covered with short, up to 3 mm long, yellowish
bristles. Corolla pale-purple, 12-13 mm long, narrowed over 5—6 mm,
with 2.5-3 mm long limb; basal appendages of anthers setaceous, 2—3
mm long, undivided or almost undivided; filaments smooth. Pappus
deciduous, up to 10 mm long, snow-white, short-barbate, barbules densely
arranged, approximately as long as thickness of bristles; all bristles
connate at base into ring, through which nectary passes. Achenes four-
angled, narrow, 4-6 mm long, with small teeth above along edges,
especially at corners, with scarcely visible scattered spinules and sessile
glandular hairs in upper half, modified in lower half into distinct,
nonuniform folds; pappophorus conical, with short nectary at tip.
Flowering VII(?)-VII; fruiting VITI-IX(?).

Pebbles, gravelly places.—Soviet Central Asia: Tien Shan. Endemic.
Described from a place at the confluence of the Kyzylsu and the Ak-
Sai rivers. Type in Leningrad.

141. J. poacea [jin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XX (1960) 344, Fig. 1.

Perennial. Plants 30-50 cm high, many-stemmed, glabrous, green.
Stems bearing fibrous roots, finely sulcate, simple or with few upward
spreading branches in lower part. Leaves linear, usually 6-8 cm long
and 3—5 mm wide, or still narrower, narrowed at base and usually very
short and narrowly decurrent, short acuminate, undivided and entire,
green and glabrous on both sides, with one whitish vein and scattered
sessile glandular hairs above in deep alveolae; leaves obliquely directed
upward, numerous, solitary in uppermost part of stem, small, petiolate.
Corolla solitary terminal on stem. Involucre cupuliform, with weak
arachnoid-hairy pubescence, 10-15 mm long, with many oblong, ovate,
inner broadly lanceolate, short-acuminate bracts, usually three-veined,
with narrowly membranous margin, often with uneven small teeth,
appressed. Corolla pink (?), 10-12 mm long, narrowed over 4-5 mm,
with 2.5-3 mm long lobes of limb; basal appendages of anthers setaceous,
divided up to 2 mm. Pappus deciduous, 10-12 mm long, whitish,
somewhat yellowish at base, comprising many unequal, somewhat flat
bristles, especially longest, with dense barbules, equal or shorter than
flattened side, connate in a ring. Achenes 2.5—3 mm long, with distinct
markings of discontinuous striations, almost continuous on edges, corona
with unequal teeth, almost membranous, with gentle conical pappophorus

697 Plate LI.

1—Jurinea persimilis Ijin; 2—J. longicorollaris Min;
3—J. chaetocarpa I\jin.

698

690

and nectary above passing through the ring. Flowering VII—VIII(?);
fruiting VIII-IX. (Plate XL, Figs. 6, 7.)

Rocks, stony slopes.—Soviet Central Asia: Tien Shan. Endemic.
Described from the vicinity of Lake Sary-Chilek. Type in Tashkent.

142. J. baldshuanica Winkl. in Tr. Peterb. Bot. Sada, XI, 2 (1892)
283; O. and B. Fedtsch. Perech. Rast. Turk. IV, 302; Iljin in Tr. Turk.
Nauchn. Obshch. 2, 25.—J. hissarica Iljin Ibid. 35.—Ic.: Iljin, Ibid.
Plate VIII.

Perennial. Plants extremely polymorphic, 25-100 cm high or taller;
rhizome short, producing strong funiform adventitious roots. Stems few
or solitary, simple or with isolated long branches originating in lower
half, erect, cylindrical, distinctly nbbed-sulcate, finely arachnoid-hairy,
with scabrous moniliform hairs below, floccose-arachnoid-hairy above,
especially below capitulum, covered with scattered, fine, sessile, glandular
hairs, leafless from middle. Basal and lower cauline leaves gradually
narrowed into long, more or less winged petiole, undivided and entire,
lyrately divided or lyrately toothed, with very large, terminal, usually
acute lobe and small lateral teeth; middle cauline leaves sessile, narrowed
toward base, long-decurrent with undivided or somewhat toothed wing,
with distinct network of veins on both sides, scabrous above due to
short, somewhat stiff, moniliform hairs, grayish beneath due to arachnoid-
hairy pubescence, veins and petioles with large, many-celled, moniliform
hairs. Capitula large, solitary, with many florets. Involucre cupuliform,
four-or five-rowed; outer bracts somewhat thick, green, herbaceous,
acuminate, lax (similar to J. cyanoides), usually somewhat floccose-
arachnoid-hairy and, besides, covered with fine appressed bristles,
recurved in immature fruits, other bracts appressed, gradually acuminate,
with prominent veins, membranous-fimbriate, somewhat ciliate, covered
on outer side with short appressed bristles and scattered sessile glandular
hairs, purple. Receptacle densely squamose, scales smooth, acuminate.
Corolla purple, covered with sessile glandular hairs, 17-20 mm long,
narrowed over 6—8 mm, with 3—5 mm long lobes of limb; apical
appendages of anthers about 3 mm long, abruptly narrowed into fine
cusp, uncately incurved; basal appendages caudate or membranous,
divided, 2-2.5 mm long, filaments somewhat thick, glabrous or slightly
scabrous; style glabrous; stigma about 3 mm long, divided up to base,
finely papillate, with short erect bristles at base. Pappus deciduous, snow-
white or slightly yellowish, 7-10 mm long (excluding three or four
longer bristles), not strongly connate into ring, but shedding easily on
maturity; bristles unequal, short-barbate, with barbules as long as or
one and one-half times (less often two times) as long as thickness of
bristles; bristles connate into ringlet, through which pierces nectary with

699

691

small membranous lobe and surrounded by broadly funnél-shaped
structure. Achenes 6—7 mm long, oblong-obpyramidal, whitish, glabrous,
toothed above along edges, with parallel approximate bands along
longitudinal ridges below upper edge to one-third or half of achene
length; pappophorus conical, inside corona, with nectary on tip of
narrowed part. Flowering V—VII; fruiting VI-IX.

Alpine and subalpine zone, in belts of juniper, rose communities,
tallgrass meadow and alpine steppes, less often reaching into the belt of
woody vegetation, found at 1,800—-3,500 m.—Soviet Central Asia:
Pamiro-Alai Region. Endemic. Described from the vicinity of the village
of Kangurt. Type in Leningrad.

Note. An extremely polymorphic species, highly variable in growth
and morphological characters, varying from the type described by
Winkler as J. baldshuanica Winkl., to the type described by Iljin as J.
hissarica I\jin. Based on the herbarium material, which is still inadequate,
it is difficult to determine whether these are unique races (for example,
J. baldshuanica subsp. hissarica \in) or forms associated with alpine
tallgrass and subalpine steppes. In extreme cases, they sharply differ in
their growth habit, although the individual morphological characters are
found in very different combinations, which requires detailed study under
natural conditions.

This description of the species is proposed taking into account all
the forms studied that are available in the Herbarium of the Botanical
Institute, Academy of Sciences of the USSR, and in the herbarium in
Tashkent.

143. J. kirghisorum Janisch in Tr. Obshch. Estestv. Kaz. Univ.
XLI (1905) 5, Plate 2, 2; Tr. Peterb. Bot. Sada, XXVIII, 4; O. and B.
Fedtsch. Perech. Rast. Turk. IV, 298; Ijin in Tr. Turk. Nauchn. Obshch.
2, 17; Fl. Yugo-Vost. VI, 390.

Perennial. Semishrub, 10—25 cm high, densely covered with leaf
scars, in basal woody part of branches, with numerous strongly branched
reduced nonfloriferous branches; floriferous branches of current year, as
also leaves, grayish due to fine arachnoid-hairy pubescence, simple, with
one to three branches above, leafy up to half or two-thirds. Leaves
spatulate, gradually narrowed toward base, entire or with sparse, often
attenuate teeth, usually subacute; uppermost leaves almost linear, sessile.
Capitula solitary. Involucre 11-13 mm long and 10-12 mm wide (at
fruiting up to 14 mm), ovoidly cupuliform, grayish due to arachnoid-
hairy, roundish at base, comprising appressed weakly obtuse, imbricate
bracts; outer bracts oval, middle oblong, and innermost lanceolately
linear, acute, membranous-fimbriate. Receptacle covered with long
bristles up to half or two-thirds as long as involucre. Corolla pink, about

700

692

15-16 mm long and narrowed over 6—7 mm, with 3.5-4 mm long lobes
of limb. Pappus persistent, snow-white, 5-7 mm long, barbate, with
barbules dense, almost as long as or slightly longer than thickness of
brisltes. Achenes 4—5 mm long, pale-yellowish, sharp tuberculate along
edges and ribs; tubercles in longitudinal rows, terminating with incurved
cusps; pappophorus flask-shaped, with small, cylindrical, five-toothed
nectary at tip. Flowering V—VI; fruiting VII-—VIII.

Chalk beds.—European Part: Lower Volga (eastern part); Soviet
Central Asia: Aralo-Caspian Region. Endemic. Described from the chalky
Akchattau Mountains north of Uilsk. Type in Leningrad.

144. J. spissa Iljin in Izv. Glavn. Bot. Sada, XXVII, 1 (1928) 80;
Fl. Turkm. VII, 220.

Perennial. Semishrub, 5—25 cm high, with long woody root,
caespitose, with numerous caudices bearing rosettes of leaves. Stems 1—
16 cm long, arising singly from apex of each stolon, erect or often
spreading, simple, longitudinally striate-sulcate, sparsely and weakly
arachnoid hairy-pubescent, greenish-yellow or with purple tinge. Basal
leaves very numerous, linear, 2.5—7.5 cm long, involute up to midrib,
grayish due to arachnoid-hairy pubescence, gradually narrowed at base
into flattened petiole and here woolly-tomentose; cauline leaves few,
sessile. Capitula solitary terminal on stem, almost always declinate.
Involucre poculiform, 15—25 mm long and 7.5—12 mm wide; involucral
bracts relatively lax, almost appressed up to anthesis, usually with
divergent apices at full bloom and fruiting, weakly arachnoid-hairy, entire
upper part pink-purple, along border, also in inner bracts, lanceolate,
long-toothed, especially innermost bracts (latter never divergent).
Receptacle densely covered with yellowish, smooth, very short bristles,
often connate at base forming corona around achene. Corolla pink-purple,
20-23 mm long, gradually narrowed over 7-10 mm, with 4-6 mm long
limb, with scattered glandular hairs; anther appendages 3-4 mm long,
thin, setaceous; filaments smooth. Pappus deciduous, snow-white,
plumose-barbate, with barbules three to four times as long as thickness
of bristles, not very dense; pappus bristles connate at base into short
tube. Achenes up to 6 mm long and about 2 mm thick; obiong, four-
angled, with well developed, almost membranous, corona at apex;
somewhat toothed along edges, separated from echene itself; edges of
achene finely tuberculate-scabrous, with scattered glandular hairs, ribs
with toothed, whitish scaly thickenings, with small conical pappophorus
above, in center, enclosing nectary on long stalk. Flowering VI; fruiting |
VI-VII(7). (Plate L, Fig. 1.)

Gravelly, stony mountain slopes, rocks.—Soviet Central Asia:
mountainous Turkmenia (Bolshie Balkans). Endemic. Described from
the Bolshie Balkans from the Mugur Spring. Type in Leningrad.

701

693

145. J. czilikinoana I[ljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, VII, 10 (1937) 246.

Perennial. Semishrub, 20-40 cm high, with relatively dense grayish
compact remains of old leaves and woody lower branches, forming loose
turf; vegetative shoots numerous, erect or ascending in lower part, simple,
finely floccose-arachnoid-hairy, striately striped, densely leafy in lower
part, leafless above. Leaves 2—5 cm long and about 1 mm wide, almost
cylindrical, undivided, strongly involute, finely arachnoid-hairy, obliquely
directed upward. Capitula solitary. Involucre cupuliform, about 15 mm
long, roundish at base, floccose-arachnoid-hairy; involucral bracts
appressed, linearly lanceolate or lanceolate (outer bracts) one-veined,
gradually narrowed into fine cusp, herbaceous-green, later turning gray;
inner bracts linear, narrowed into still thinner and longer, aristate,
membranous, usually purple and somewhat divergent cusp (like other
involucral bracts). Receptacle densely covered with short bristles about
one-third as long as involucre. Pappus persistent, snow-white, 8-10 mm
long, short-barbate, with dense barbules as long as thickness of bristles.
Achenes 5.5—6 mm long, four-angled, obpyramidal, with thickened ribs,
fimbriate above with toothed corona; coronal teeth continued in upper
part of edges and located as parallel dicontinuous ridges at extreme tip,
connate at base and, later smooth; pappophorus somewhat large
cupuliform at achene apex, with five-toothed nectary inside. Flowering
V-VI(?); fruiting VI-VII(?).

Gravelly desert slopes of mountains —Soviet Central Asia: Tien Shan
(Karatau). Endemic. Described from the mountains of Greater Aktau.
Type in Leningrad.

146. J. fedtschenkoana Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada,
V, 4-5 (1924) 7; Iljin in Tr. Turk. Nauchn. Obshch. 2, 17.

Perennial. Semishrub, up to 25 cm high, with short, woody stems,
arising from turf, covered with scales of dead leaves, one-year-old
branches usually simple, floccose-arachnoid-hairy. Leaves linearly-
filiform, strongly involute, sessile; lower leaves somewhat broader at
base, with remote, solitary, linear or toothed lobes along margin. Capitula
solitary. Involucre 1-1.5 cm wide, weakly arachnoid-hairy; involucral
bracts short-acuminate with scarcely visible very short and appressed
bristles, ovate (outer) to oblong-lanceolate. Corolla 13-14 mm long,
narrowed over 5 mm, with 4 mm long lobes of limb. Pappus deciduous,
barbate or barbate-serrate, with dense barbules, almost as long as or
slightly shorter than thickness of bristles. Achenes about 6 mm long,
compressed-four-angled, longitudinally tuberculate, with appressed
spinules above and along edges, connate below into longitudinal ridges,
thickened along main ribs, with thick patellate pappophorus above,

702

694

enclosing cylindrical, milky-white, sinuate, nectary. Flowering V—VI;
fruiting VI-VII. (Plate XLVI, Fig. 2.)

Chalk beds.—Soviet Central Asia: Aralo-Caspian Region. Described
from chalky Akbut mountains south of Temir. Type in Leningrad.

SPECIES OF UNCLEAR POSITION
Section 10. Derderia (Jaub. and Spach) Boiss.

1. J. pteroclada [ljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, VIII, 10 (1940) 195.

Perennial. Plants 20-25 mm high, more or less many-stemmed,
grayish-green, almost glabrous, or somewhat floccose. Stem virgate,
simple or branched in lower part; branches long, like stem, winged,
one-headed. Leaves oblong, undivided and entire, subobtuse and short-
cuspidate, often scarcely sinuate above, one-veined, long and broadly
decurrent, and almost appressed, borne right up to top. Capitula sessile.
Involucre turbinate, 25-30 mm long, slightly narrowed at base, somewhat
arachnoid; involucral bracts imbricate, linear, appressed, narrowed into
straight awn, 3.5-7 mm long.Receptacle densely setose, with up to 2
mm long bristles. Corolla pink, densely glandular, 28-30 mm long, with
10-17 mm long tube and 4-5 mm long lobes of limb; basal appendages
of anthers filiform. Pappus snow-white, up to 20 mm long, with almost
plumose unequal bristles. Achenes immature, glabrous and smooth.
Flowering VIII; fruiting IX(?).

Deserts in the region of varicolored adyrs.—Soviet Central Asia:
Pamiro-Alai Region. Described and known only from the vicinity of the
Kim Springs. Type in Dushanbe.

Section 11. Stechmannia Boiss.

2. J. bracteata Rgl. and Schm. in Tr. Peterb. Bot. Sada, VI, 1
(1880) 323; Trautv. ibid. VIII, 1, 510; O. and B. Fedtsch. Perech. Rast.
Turk. IV, 299; Iljin in Tr. Turk. Nauchn. Obshch. II, 15. ;

Perennial. Semishrub, 7-12 cm high; caudices covered with remains
of old leaves, some bearing rosettes of leaves, others modified into
floriferous shoot. Stems erect or usually weakly ascending, covered up
to tip with small, bracteal leaves, simple or often branched above, bearing
one to three capitula. Leaves on vegetative shoots and basal leaves
oblong-lanceolate, sinuate-toothed or sinuate-pinnatipartite, gradually
narrowed below, pointed like teeth, involute, scabrous and glutinous
above. Involucre turbinate, about 15 mm long, many-rowed, glabrous;

695

involucral bracts appressed, outer significantly shorter, ovate, subobtuse,
gradually elongated inward; inner bracts linear, acuminate; all bracts
with distinct midrib. Receptacle densely covered with hairs, up to three-
fourths as long as involucre. Pappus barbate; immature achenes smooth.
Fruiting V. |

Soviet Central Asia: Balkhash Region. Endemic. Described from
the Ili River Valley near Karachoku. Type in Leningrad.

Note. Until now known only from the original specimen of A. Regel,
which was quite fragmentary. Owing to the lack of achenes and pappus,
it is difficult to assign it to a particular section.

3. J. spiridonovii Iljin, in Addenda XXVI, 724.

Perennial. Semishrub, 8—15 cm high, with many-headed caudex and
short rhizomes covered with reddish brown scales and producing rosettes
of leaves at soil surface. Stems simple. Basal leaves oblong-linear,
gradually narrowed into petiole, somewhat broadened at base, undivided
and entire, hard, with depressed midrib, rugose, and weakly arachnoid-
hairy above, also with scattered sessile glandular hairs, whitish-tomentose
beneath with prominent midrib, glandular under tomentum; cauline leaves
absent. Capitula small. Involucre about 10 mm long, poculiform;
involucral bracts appressed, almost glabrous; outer bracts ovate, middle
oblong-oval, inner exceeding all others, membranous in upper part.
Receptacle covered with long, thin, smooth scales almost as long as
involucre. Corolla, pappus and achenes absent.

Soviet Central Asia: Aralo-Caspian Region. Endemic. Described from
the Selekta River Valley, 1.5 km west of the Tas-Mola Tomb, on the
basis of the collection of M.D. Spiridonov.

Section 15. Chaetocarpa (Korsh.) Iljin.

4. J. mariae Pavl. in Vestn. Akad. Nauk KazSSR, 3(60) (1950)
34.—Ie.: Pavl. Ibid, Fig. 11.

Perennial. Plants 35-45 cm high, loosely turfy, producing numerous,
long, woody caudices. Stems few, more or less ascending, cylindrical,
finely sulcate-ribbed, sparsely floccose-arachnoid-hairy, with scabrous,
short, acute spinules along ribs, weakly branched up to two-thirds height
with long, more or less arcuate branches, almost leafless in upper one-

703 third part. Leaves obovate-oblong, or uppermost and basal leaves
oblanceolate, entire, short-cuspidate, frequently with unequal large teeth
in lower part, with triangular acute teeth; basal and lowermost cauline
leaves narrowed into short narrow-winged petiole; other cauline leaves
sessile, decurrent; all leaves weakly scabrous up to tip due to stiff,
somewhat acute, tubercles, green, without arachnoid-hairy pubescence,

704

696

weakly rugose and with sessile glandular hairs, somewhat grayish beneath
due to sparse-arachnoid-hairy pubescence, with numerous sessile
glandular hairs; uppermost leaves small, somewhat decurrent. Capitula
solitary, with many florets, on long peduncle. Involucre cupuliform, 15—
16 mm long and 8-10 mm wide, roundish at base or slightly depressed,
slightly floccose-arachnoid-hairy in lower part or glabrous; bracts
appressed with prominent veins; outer bracts herbaceous green; middle
and inner bracts purple, membranous and ciliolate along margin, with
scarcely visible, short, appressed bristles on outer surface; all bracts
gradually acuminate. Receptacle densely squamose; scales smooth, acute,
up to half as long as involucre. Corolla pink-purple, glabrous, 15—17
mm long, narrowed over 4—5 mm, with 4-5 mm long lobes of limb;
apical appendages of anthers about 3 mm long, abruptly narrowed into
thin cusp, basal appendages caudate, about 2—2.5 mm long; filaments
slightly tuberculate; style glabrous; stigma about 3 mm long, divided
almost up to base, very small papillate, with longer erect, short bristles
at base, somewhat continued on style. Pappus snow-white, 12-13 mm
long; bristles equal, short-barbate; barbules dense, as long as or shorter
than thickness of bristles. Achenes immature, 4-5 mm long, obpyramidal,
four-angled, smooth, with plicate, discontinuous, parallel longitudinal
ridges, terminating in very short teeth, with toothed corona above along
edges. Flowering VII—VIII.

Stony screes in subalpine region.—Soviet Central Asia: Tien Shan.
Endemic. Described from southern Kazakhstan from Bostandyk in the
upper part of Aksarsai.

Note. According to N.V. Pavlov, it is close to J. capusii Franch.,
but differs in having entire leaves, a glabrous involucre, and a shorter
corolla (7-9 mm long).

5. J. monticola Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada, V, 11-—
12, (1924) 69; Tr. Turk. Nauchn. Obshch. II, 26.—J. abramovii O. and
B. Fedtsch. Perech. Rast. Turk. IV (1911) 303 p. p.

Perennial. Plants 5—15 cm high, usually with ascending stems, one-
year-old stems usually simple, arachnoid-hairy; leaves 1—2.5 cm long,
spatulate or almost oblong, gradually narrowed toward base, narrowly
involute, pinnatipartite or less often pinnatilobate, with very few,
triangular-oblong, weakly obliquely directed upward lobes, grayish-green
above, grayish-tomentose beneath; basal leaves numerous; uppermost
leaves linear or filiform. Capitula solitary. Involucre about 15 mm wide;
involucral bracts linear, acuminate, with scarcely visible teeth, purple,
almost glabrous, appressed. Receptacle squamose; scales one-third as
long as involucre, or longer, but reaching up to half its height. Corolla
pinkish, 13-14 mm long, narrowed over 5—6 mm, with 3.5-4 mm long

697

lobes of limb. Pappus persistent, with barbules almost as long as or
slightly shorter than thickness of bristles. Achenes with patellate, reddish-
brown, toothed, pappogenus at apex, enclosing nectary.

Gravelly slopes.—Soviet Central Asia: Tien Shan (Karatau).
Described from Mindzhelke in the northern part of Karatau Range. Type
in Leningrad.

GENUS 1585. Jurinelia Jaub. and Spach.':?

Jaub. and Spach, Ill. pl. or. II (1844-1845), tab. 183-185.

Capitula with many florets; receptacle alveolate; all florets bisexual;
corolla actinomorphic, long-pedunculate; anthers glabrous with apical
appendages gradually acuminate, basal appendages membranous, caudate-
flexuous; filaments more or less densely covered with fine hairs at
transition of tubular stipule into constriction; style glabrous; stigma bifid
up to base, with tuft of bristles at base, remaining part densely covered
with very short papillae. Pappus fragile, few-rowed, scabrous-barbate or
scabrous, with all outer shorter hairs connate into ring and easily
separated from achene, with tuberculate thickening inside (pappogenus)
and nectary in center, latter roundish, five-toothed along the margin.
Achenes oblong or oblong-oval, slightly obpyramidal, four-angled,
without corona, with smooth margins, modified into a beak in middle,
basal hilum, with longitudinal striations and usually with transverse
depressions. Plants mostly stemless, perennial. Only four species have
been reported so far.

Type species: J. subacaulis (Fisch. and Mey.) Ijin.

—

. Plants stemless or with poorly developed stem...................:006 2.
a Plants wathwl0—30xemmhigh|istemse! 210.3... Me ANS e, PES

pets: well ceeveny. ANE Ss, 3. J. squarrosa (Fisch. and Mey.) Iljin
2. Leaves lyrately divided, with large ovate or roundish, usually entire

lobeseazaeranitoei el oats ee 1. J. moschus (Habl.) Bobr.
+ Leaves uni- or bipinnatisect, with terminal lobe equaling lateral
lobessets: wenn lait 2. J. subacaulis (Fisch. and Mey.) Ijin

1. J. moschus (Habl.) Bobr. in Bot. Zhurn. XLVII, 4 (1962) 580.—
Jurinella aucheri Jaub. and Spach, Ill. pl. or. Il (1844-1846) 183.—
Centaurea moschus Habl. in Neue nord. Beiter. IV (1783) 54 and ‘in
S.G. Gmel. It. TV (1784) 61.—Serratula depressa Stev. in Mém. Soc.

'Treatment by M.M. Iljin.
?Diminutive of generic name Jurinea.

706

698

Nat. Mosc. IV (1813) 101; MB. FI. taur.-cauc. III, 548—Onopordon
lyratum Schulz Bip. in Linnaea, XIX (1847) 329.—Jurinea depressa
C.A.M. Enum. cauc. (1831) 67; Boiss. Fl. or. III, 583; DC. Prodr. VI,
677; Ldb. Fl. Ross. II, 767; Grossh. Fl. Kavk. IV, 170; Sosn. in Zhurn.
Russk. Bot. Obshch. 11, Nos. 1-2, 200.—Jurinea depressa var.
biebersteiniana Trautv. in Tr. Peterb. Bot. Sada, VII, 2 (1880) 474.—
Jurinea moschus (Habl.) Bobr. in Bot. Zhurn. XLII, 11 (1958) 1545.—
Ic.: Trans. Linn. Soc. XI, tab. 38; Tchichatcheff, As. Min. atl. bot.
(1860) tab. 33.

Perennial. Plants stemless or with very short stem, less often up to
15 cm high (f. caulescens Sosn.). Stem simple, or sometimes branched;
roots with musky smell. Leaves usually in basal rosette, lyrate, with
large terminal lobe, broadly oval or oblong-ovate, usually cordate at
base or less often roundish, entire, sometimes irregularly small toothed
or boldly sinuate, and one to five (six) pairs of distinctly separate, much
smaller, roundish or ovate, less often oblong, lateral lobes, usually
narrowed at base, entire or more or less lobate, gradually narrowed into
long, flat petiole amplexicaul on lower side; cauline leaves significantly
reduced, and simple, pale greenish or yellowish-green, rugose above,
weakly arachnoid-hairy, somewhat scabrous due to sparse and fine
papillae, gray or whitish beneath due to arachnoid-hairy tomentum, with
network of veins on both sides, depressed on upper surface and raised
beneath. Capitula large, with many florets, usually in groups of few due
to approximate branches bearing solitary heads. Involucre broadly
cupuliform, 15-30 mm long, many-rowed, arachnoid-hairy, with
herbaceous bracts; outer and middle bracts recurved, innermost row with
herbaceous, membranous, soft, long-acuminate, straight bracts with
scarcely visible appressed bristles and sessile glandular hairs. Receptacle
alveolate, alveolae surrounded by acuminate smooth scales, several times
shorter than involucre. Florets usually fragrant. Corolla pale, white,
or pinkish, less often even yellowish, 22-30 mm long, narrowed at
10-18 mm, with 6-8 mm long lobes of limb, scatteredly covered by
sessile glandular hairs; anthers usually violet; appendages 3—3.5 mm
long, gradually acuminate upward, basal appendages 2.5-3 mm
long, membranous-caudate and sinuate; filaments covered with
very short hairs. Pappus less often white, slightly yellowish or dirty
yellow, smoky, in few rows, easily separated from achene, usually 15—
20 mm long; pappus bristles unequal, but barbate, with very sparse
barbules, fragile, connate into ring. Achenes 5—6 mm long, light or dark
gray, oblong or somewhat obpyramidal, more or less four angled, finely
longitudinally and usually also transversely strongly and scabrously
rugose, roundish at tip, modified into short, whitish, pappophorus.
Flowering VI-VIII; fruiting VI—VIII.

707

699

Screes, rocks in ravines, clayey slopes in alpine and subalpine
zones.—Caucasus: Ciscaucasia, Dagestan, western, eastern, and southern
Transcaucasia. General distribution: Northwestern Iran, northwestern
Turkey. Described from eastern Caucasus between the mountain streams
Khodial and Kussarchai.

Type in London?

Note. An extremely variable species. Plants with fruits of three
different kinds are found: 1) light gray, with fine, close, longitudinal,
threadlike ridges and transverse, deep, oblong-oval depressions; usually
these achenes do not exceed 5 mm in length; 2) longer, to 6 mm, darker
and more narrowly oblong, with only fine longitudinal markings; and 3)
intermediate forms. But are these different developmental stages? The
question remains unanswered for want of enough material and the
absence of studies in nature.

2. J. subacaulis (Fisch. and Mey.) Ijin comb. n—Jurinea subacaulis
Fisch: and Mey. in Ind. sem. H. Petropol. IV (1837) 39; Ldb. Fl. Ross.
II, 767; Sosn. in Zhurn. Russk. Bot. Obshch. XI, 1-2, 200; Grossh. FI.
Kavk. IV, 170.—J. rhizantha DC. Prodr. VI (1837) 677.—J. depressa
a. pinnatisecta Boiss. Fl. or. II] (1875) 583.—Jurinella chamaecynara
Jaub. and Spach, Ill. pl. or. I] (1844-1846) 184.—Ie.: Jaub. and Spach
Ibid tab. 184.

Perennial. Plants stemless or less often with 15 cm high stems (f.
caulescens Boiss.), with single large capitulum up to 6-8 cm wide or
with two to four aggregated capitula; rhizome virgate or less often few,
merging into very short underground stem, covered with fibrous remains
of old leaves. Leaves basal, numerous, petiolate, with somewhat flat,
usually dirty purple petioles, broadened at base, lamina usually pinnatisect
or bipinnatisect, with terminal lobe equal or almost equal to lateral lobes,
or distinctly larger, but not roundish as in previous species (f. lyrata
(Trautv.) Iljin = J. lyrata Trautv.); lateral lobes oblong or oval, usually
subobtuse, sometimes acute, somewhat narrowed toward base, often more
or less approximate, on narrow-winged rachis, with entire margin,
toothed, lobate or divided, less often entire (f. subintegra Sosn.), glabrous,
green, or weakly arachnoid hairy-pubescent, smooth, often slightly rugose,
occasionally somewhat scabrous, and involute above, whitish beneath
due to more dense tomentum; rosette flat or arcuately raised, and
surrounding capitulum. Capitula with many florets. Involucre broadly
cupuliform; outer bracts recurved, cuspidate and floccose-arachnoid-hairy
and erect; inner bracts thin-membranous, gradually acuminate, more or
less dirty-purple, with small teeth, often with scattered sessile glandular

hairs. Receptacle alveolate, covered with short (about 5-7 mm long),

smooth, acuminate scales, connate at base around alveoles. Corolla 20— -

708

700

25 mm long, narrowed at 10-15 mm, with 5—8 mm long lobes of limb
bearing scattered sessile glandular hairs; anthers usually violet; apical
appendages attenuate, about 3 mm long, basal appendags 2-2.5 mm
long, caudate, divided below; stigma divided up to base, acute, 2—2.25
mm long, with papillae; style glabrous. Pappus 12—20(25) mm long,
fragile, snow-white, later turning yellow, somewhat serrate, with
serrations shorter than thickness of bristles, less often almost as long;
pappus bristles in few rows, somewhat unequal, all connate into ring,
shedding easily. Achenes 4—6 mm long, almost obpyramidal, four-angled,
blackish-brown or dark gray, strongly transversely rugose, glabrous, more
or less truncate at apex, with roundish edges, terminating with dome-
shaped, whitish pappophorus in center covered with scales. Flowering
V(end)-VII; fruiting VII—VIII.

Alpine and subalpine belts, on moraines, along screes, slopes of
ravines, in alpine tapetum and subalpine meadows on clayey-gravelly
and sandy-gravelly soils; on limestone debris reaches permanent snow
line and to edges of glaciers, 2,000 to 3,500 m.—Caucasus: Ciscaucasia,
western, eastern, and southern Transcaucasia. General distribution: Asia
Minor, northwestern Iran. Described from Azerbaidzhan, from the peaks
of Arekligeduk Mountain, as Jurinea subacaulis, in the same year, from
the same place, and on the basis of the same specimen as Jurinea
rhizantha. Type in Leningrad.

Note. An extremely polymorphic species, close to Jurinella depressa
(Stev.) Iljin. It is possible that after detailed study in nature and under
cultivation a series of separate races can be recognized. The Karabakh-
Nakhichevan race, subsp. rhizantha (DC.) Iljin, with fine, usually
bipinnatisect, arcuately bent leaves with acute lobes, is quite distinct, as
well as race segregated by Trautvetter as a separate species: Jurinea
lyrata Trautv., which in typical form, is widespread in Balkaria (for
example, on the Shtulu Pass) and is described by him on the basis of
Radde’s collection from Neshkadag (Jnd. sem. H. Petropol. (1865) 37).

3. J. squarrosa (Fisch. and Mey.) Iljin comb. n.—Jurinea squarrosa
(Fisch. and Mey.). Ind. sem. horti. Petropol. XI (1845) 75; Grossh. FI.
Kavk. IV, 170; Sosn. in Zhurn. Russk. Bot. Obshch. 11, 22, 200.—J/.
coelestis Sosn. in Vestn. Tifl. Bot. Sada, 49: (1920) 34.

Perennial. Plants mostly (8)10-30 cm high; roots vertical, merging
with a short rhizome densely covered with remains of old leaves. Stems
solitary or few, simple, relatively thick, distinctly sulcate-striate, strongly
arachnoid-hairy due to thin long hairs. Leaves in basal rosette many,
petiolate, broader toward base, narrow-winged; lower cauline and middle
leaves on shorter petioles, imparipinnatisect and pinnatisect into narrowly
lanceolate segments with longer terminal and oblique lateral lobes, often

709

701

geminate at base and sometimes short, entire, green above, usually
scabrous due to more or less short papillae, lustrous, densely white-
tomentose beneath, involute almost up to midrib; uppermost leaves entire,
linear, usually arachnoid-hairy. Capitula 2-5 cm wide, solitary terminal
on stems, with many florets. Involucre broadly cupuliform, 20-25(30) |
mm long, with depressed base; outer bracts arachnoid-hairy, squarrose,
hard, prickly, with prominent midrib, bracts green, purple. at ends;
innermost bracts erect, membranous, ciliate, mostly purple, with whitish
or yellowish cusp, at fruiting all bracts recurved. Receptacle alveolate-
pitted, strongly squamose; scales equal, smooth, acuminate, surrounding
alveoles and usually connate at base, slightly shorter than half of
involucral length. Corolla 17-23 mm long, narrowed over 8-12 mm,
with 5-7 mm long lobes of limb; apical appendages of anthers more or
less gradually acuminate, 3—-3.5 mm long, basal appendages 2-3 mm
long, divided below, often caudate; filaments finely tuberculate; stigmas
22.5 mm long, divided up to base, surrounded by tuft of short bristles,
often gradually modified into small papillae on outer side; style glabrous.
Pappus fragile, snow-white, serrate-barbate or somewhat serrate, with
serrations shorter than thickness of bristles, comprising unequal bristles
arranged in irregular rows, innermost bristles longest, all connate into
ring, easily detached; ring inside pappus modified into broad area
terminating with pentagonal base of corolla. Achenes glabrous, oblong-
obpyramidal, four-angled, 4.5-5.5 mm long, dark gray, with dark
transverse spots, finely striate; mature achenes strongly rugose
transversely, without corona, terminating into whitish tuberculate
pappophorus covered with scales. Flowering VI-VII; fruiting VII—VIII.

Subalpine zone, mostly on rocks and sandy slopes.—Caucasus:
Southern Transcaucasia (Trialeti Range, Nor-Bayazet, Sevan, Alagoz).
Endemic. Described from Alagyoz. Type in Leningrad.

GENUS 1586. Perplexia Iljin':?

Iljin in Addenda XXVI, 727.

Plants stemless; capitula sessile, more or less with few florets;
receptacle comprising few, polygonal, small, glabrous areas, slightly
convex, surrounded by very short, smooth, acuminate scales; all florets
bisexual, corolla gradually narrowed toward base; apical appendages of
anthers attenuate above, basal appendages setaceous-caudate, flexuous;
pappus bristles connate at base into ring; pappus entirely shedding easily,

'Treatmen by M.M. Ijin.
*From the Latin word perplexus—unclear, confused, with double meaning.

710

702

some lower bristles above base abruptly deflexed and slightly entangled,
half of inner bristles erect; achenes oblong, turbinate, glabrous and
smooth, without corona, only with sparse, fine tubercles, surrounding
pappophorus, even when mature.

Two species of this genus are known, the one in the USSR, and the
other, P. frigida (Boiss.) Ijin, in Iran.

1. P. microcephala (Boiss.) Iljin in Addenda XXVI, 727.—Jurinea
microcephala Boiss. Fl. or. III (1825) 584; Iljin in Tr. Turk. Nauchn.
Obshch. UI, 27.—Ie.: Iljin ibid. X, Fig. 1; Fl. Turkm. VII, Plate XXXIV.

Perennial. Plants 2-8 cm high, stemless, forming relatively loose
pads, with woody, short, numerous, caudices covered with remains of
old leaves, terminating with rosette of short leaves, 5—10(15) mm long,.
linear, somewhat narrowed at base, entire, involute up to midrib, grayish
due to arachnoid-hairy pubescence. Capitula sessile, solitary. Involucre
cylindrical, 10—-12(15) mm long; outer bracts sparsely arachnoid-hairy,
tapering from oblong-ovate base toward apex and with curved tip, broadly
membranous along margin; inner bracts lanceolate, glabrous,
membranous, acuminate and usually weakly recurved at tip or even erect;
all bracts without distinct veins. Receptacle squamose; scales few,
smooth, several times shorter than involucre. Corolla pink, with sessile
glandular hairs, 12-17 mm long, narrowed over 5—8 mm and with 4.5—
5 mm long limb; anther appendages setose, longitudinally parted. Pappus
snow-white,- 10-12 mm long, comprising thin, lax, very finely serrate
bristles. Achenes 4.5—6 mm long and 1.5—2 mm wide, oblong-turbinate,
with scarcely visible ribs, glabrous and smooth, matte, gray. Flowering
VU; fruiting VII-VIII(?).

Stony habitats in subalpine mountain zone.—Soviet Central Asia:
mountainous Turkmenia. General distribution: Iran. Described from
Khorasan, between Nishapur and Mashhad. Type in Geneva.

GENUS 1587. Pilostemon Ijin’: ?

Ijin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XXI (1961)
39%:

Capitula cyathiform. Receptacle punctate-alveolate, with alveoli
surrounded by short, smooth, acuminate scales, connate at base. All
florets tubular, bisexual; corolla glandular; anthers covered with long,
thin, crisped, fragile, easily shedding hairs; filaments glabrous; style
glabrous; stigma divided up to base into arcuate branches, scarcely

'Treatment by M.M. Iljin.
*From Greek words pilos—hair; and stemon—stamen.

71

—

703

enlarged at tip, grooved above, with tuft of short bristles at base. Pappus
snow-white, of numerous, many-rowed, almost equal plumose bristles;
outer bristles shorter, adherent to achene, others weakly connate into
ting, always without two to four prominent long bristles (as in Jurinea).
Achenes weakly obpyramidal, oblong, three- or four-angled, densely
covered with sessile and short-stalked glandular hairs; hilum basal. Plants
herbaceous, perennial. A West Asian and Soviet Central Asian species.
Type species: Pilostemon karategini (Lipsky) Iljin.

1: Leaves oblong; corolla 10-13 mm long, narrowed over 3-5 mm,
lobes of limb 1.25—2 mm long; achenes smooth ..............cceceee
Bsc wie alta adh negate sadiggnei onset Plase scores 1. P. karategini (Lipsky) Iljin

+ Leaves filiform; corolla 16-20 mm long, narrowed over 7-8 mm,
lobes of limb 3-4 mm long; achene longitudinally sulcate..........
Be Diao hh cick eras Leth, «timate ica Maat 2. P. filifolia (Winkl.) Ijin

1. P. karategini (Lipsky) Iljin in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 391.—Jurinea karategini O. Fedtsch. in O.
and B. Fedtsch. Perech. Rast. Turk. IV (1911) 297; Iljin in Tr. Turk.
Nauchn. Obshch. 2, 15.—Saussurea karategini Lipsky in Tr. Peterb.
Bot. Sada, XXVI (1909) 323.—Ie.: Iljin, op. cit. (1961) 392.

Perennial. Plants 25-80 cm high, more or less divaricately branched
from base almost up to top, canescent due to flocculose arachnoid-
hairy pubescence, less often almost green (f. glabrescens Iljin). Tap
root long and fibrous. Stems several, finely sulcate-ribbed, cylindrical,
with scattered sessile glandular hairs. Basal leaves, sometimes also
lowermost cauline leaves oblong, up to 6-10 cm long and 10-15 mm
wide, undivided and entire, usually with short cusp, grayish above, and
with depressed veins, whitish beneath due to dense velutinous
pubescence, withering before anthesis; cauline leaves linear, with
prominent midrib beneath, grayish, narrowly revolute, cuspidate, reduced

_ upward. Capitula numerous, solitary terminal on stem and branches.

Involucre poculiform, 12-15 mm long, almost entirely covered with
fine white tomentum; involucral bracts mostly five-rowed; outer bracts
small, middle oblong, with prominent midrib and pair of less distinct
veins on each side, yellowish-golden, often with violet-purple tinge,

with short, brownish cusp; innermost bracts often only. slightly

arachnoid-hairy or almost glabrous, broadly membranous, lanceolate,
tapering into longer spine, long ciliate. Receptacle alveolate, margins of
alveoli covered with yellowish basally connate scales, several times
shorter than involucre. Corolla pinkish-purple, 10-13 mm long, narrowed
over 3-5 mm, with 1.25—2 mm long lobes of limb, usually with scattered
glandular hairs, apical appendages of anthers 2.25-2.5 mm long, abruptly

704

narrowed above; basal appendages about 2 mm long, caudate, flexuous;
anthers sparsely arachnoid hairy-pubescent; filaments glabrous; style
glabrous; stigma 2.5—2.75 mm long, divided, with very short hairs at
base, divided up to base in arcuately divergent branches with scarcely
visible papilliform hairs. Pappus snow-white, 10-15 mm long, of almost
equal bristles connate into ring, at base, less often reduced toward outside;
outer pappus shorter row of bristles with sparse and shorter barbules,
often adherent to achene; inner pappus of long, many-rowed, plumose
bristles connate with not dense ring. Achenes four-angled, obpyramidal,
4-5 mm long, yellowish, pale, smooth, covered with scattered glandular
hairs, with toothed corona above. Flowering V—VIII; fruiting VII—X.
Sandy-pebbled deposits in river valleys, stony river beds in mountain
regions, wormwood—saltwort deserts, on stony soils.—Soviet Central
Asia: Tien Shan, Syr-Darya, Pamiro-Alai Region. General distribution:
Iran (Afghanistan), Dzhungaria-Kashgaria (Sinkiang). Described from
Karategin, from the Surkhob River Valley. Type in Leningrad.

2. P. filifolia (Winkl.) Iljin in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 393.—Jurinea filifolia Winkl. in Tr. Peterb.
Bot. Sada, XI, 1 (1890) 170; O. and B. Fedtsch. Perech. Rast. Turk. IV,
296; Iljin in Tr. Turk. Nauch. Obshch. 2, 15.—Saussurea filifolia Rgl.
and Schm. in Tr. Peterb. Bot. Sada, VI, 1 (1890) 170.—TIe.: Ijin op.
cit. (1961).

Perennial. Plants 25-50 cm high, canescent due to dense arachnoid-
hairy tomentum, less often grayish green. Stems numerous, divaricately
branched from base, simple at tip, finely sulcate, with dense globules in
axils of scaly leaves at base. Leaves sessile, linear, involute up to middle,
entire, subobtuse, gradually reduced upward to distinct, bract-like, very
small leaflets; lowermost leaves up to 10 cm long, somewhat flat, often
with isolated and distant teeth, usually withering at anthesis. Capitula
numerous, solitary terminal on branches, with few florets. Involucre
poculiform, more or less densely covered with crisped arachnoid-hairy
pubescence, five- or six-rowed; outermost bracts short, others lanceolate,

- with violet-purple tinge, with prominent midrib and one or two pairs of

712

lateral veins, with scarcely visible appressed bristles under flocculose
pubescence along margin, with small teeth, terminating into short,
yellowish spine; innermost bracts cuspidate, with serrations along margin
modified into hairs. Corolla 16-20 mm long, pink-purple, covered with

sessile and short-stalked glandular hairs, gradually narrowed over 7—8

mm, with 3-4 mm long lobes of limb; apical appendages of anthers
whitish-pinkish, about 2 mm long, abruptly attenuate above, basal
appendages caudate, fiexuous, 2-3 mm long; filaments glabrous, initially
whitish, on maturation brownish-yellowish; style glabrous, with visibly

713

705

projecting bristles only below bifurcation; stigma 1.5—2 mm long, divided
and divergent, somewhat enlarged at apex, with scarcely visible hairs
outside, inner side ciliolate. Receptacle densely squamose; scales small,
smooth, acuminate, several times shorter than involucre. Pappus snow-
white, about 12—15 mm long, pappus bristles deciduous, plumose, central
bristles almost equal, four prominent brisltes absent as in Jurinea,
outermost bristles short. Achenes oblong, about 5 mm long, four- or
usually three-angled, whitish or olive green, longitudinally sulcate,
covered with sessile or very short-stalked, glandular, hairs, especially
above, toothed above, teeth translucent, more or less divided, with short
pappophorus in center bearing nectary. Flowering VII—VIII; fruiting
VIII-X.

Pebbles of rivers, dry gullies, gorges——Soviet Central Asia: Tien
Shan (central), Dzhungaria-Tarbagatai. General distribution: Dzhungaria-
Kashgaria. Described from the Khorgos River Valley from the Dzharkent
Mountains. Type in Leningrad.

GENUS 1588. Nikitinia Iljin':?

Ijin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX (1960)
356.

All florets bisexual; corolla tube long, with five-partite limb; anthers
dark brown, with membranous-setose basal appendages; filaments distinct,
densely barbate; style and stigma prominent, stigma with short tuft of
bristles at base, divided at apex, with erect or weakly divergent branches,
enlarged at tip, obtuse, covered with scarcely visible papillae; receptacle
almost cylindrical, convex, with pit, covered with long, smooth, glabrous
scales; pappus of equal bristles; achenes with basal hilum; involucre
poculiform with five- or six-rowed involucral bracts.

A monotypic genus from Soviet Central Asia.

1. N. leptoclada (Bornm. and Sint.) Iljin in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XX (1960) 356; Iljin in Fl. Turkm. VII, 235.—
Jurinea leptoclada Bornm. and Sint. in Journ. Russ. Bot. (1911) 5; O.
and B. Fedtsch. Perech. Rast. Turk. IV, 296; Iljin in Tr. Turk. Nauchn.
Obshch. 2, 14—Exs.: H.F.A.M. No. 493.—Iec.: Iljin op. cit. pp. 357,
358.

Perennial. Plants 50-100 cm high, almost semishrubs, glabrous in
all parts, somewhat woody, many-stemmed, strongly virgate, branched

'Treatment by M.M. Iljin.
*Named after V.V. Nikitin, student of the Turkmenian flora.

706

from base. Stems numerous, smooth, not always sharply sulcate, densely
branched in lower half, with few long branches above. Leaves numerous
in lower and middle part, very dense, narrowly linear, sessile, short-
cuspidate to whitish cartilaginous at apex, 20-40 mm long and 1.0—2.0
mm wide, entire, gradually reduced upward, so that capitula appear long-
pedunculate; basal leaves withering before anthesis. Capitula solitary
terminal on main stem and branches, numerous, narrow-poculiform.
Involucre 18-25 mm long and about 5—6 mm wide in middle, glabrous;
involucral bracts almost coriaceous, imbricate, glabrous; outermost bracts
finely scaly, others oblong-ovate, yellowish-brown, gently convex, with
three dark brown striations above, compactly appressed, subobtuse, very
short cartilaginous-cuspidate; innermost bracts prominent, linearly-oblong,
membranous, acuminate. Receptacle convex, covered with long, smooth,
whitish, membranous scales with fine scattered barbules visible under
high magnification, almost equaling involucre. Florets bisexual, corolla
pink, 17—19(20) mm long, narrowed over 6-8 mm, with acuminate lobes
of limb, 4—5 mm long, eglandular, connate along margin, and further
along narrower part, borders continued into darker, thin, lines; staminal
tube slightly longer than corolla with apical appendages of anthers white-
pinkish, 2.5—-3 mm long; anthers brownish, about 5 mm long, with basal
appendages, colorless, membranous-caudate, discontinuous in lower half,
0.5—1.2 mm long; filaments distinctly papillate. Pappus 8-12 mm long,
ferruginous; pappus bristles many, equal, serrate, connate into ring at
base. Achenes at maturity 6-7 mm long, narrowly oblong, glabrous,
smooth, with basal hilum and slightly narrowed here, with small apical
corona. Flowering V-—IX; fruiting VII—IX.

Stony, gravelly mountain slopes, rocks in the steppes and juniper
zone.—Soviet Central Asia: Mountainous Turkmenia. General distribution:
Iran. Described from the Iran border, from the vicinity of the village of
Sulyuklyu (Saratovka). Type in Leningrad.

714

707

ADDENDA XXVI*

DIAGNOSES PLANTARUM NOVARUM IN TOMO XXVII
FLORAE URSS COMMEMORATARUM
(Identification of New Species Included in Vol. XXVID

August, 1962

ECHINOPS L.

1. E. kafirniganus Bobr. n. sp. (sect. Ritrodes -Bge.).

Radix palaris 3—5-cephala; caules in numero duae-tres, 60-80 cm
alt. superne ramosi, non manifeste sulcati albo-arachnoidei,. sub tomento
glabri eglandulosi; folia inferiora 15-20 cm long., 6—8 cm lat. superne
pinnatisecta lobis 3—4-paris suboppositis uno terminali, lobi superi 3-4
cm long., 2-3 cm lat. margine fissi spinoso-dentati remoti breviter
descendentes; lobi inferi—1—2 cm long. late remoti; folia basi semi-
amplexicaulia spinosa; folia superiora lanceolata profunde pinnatifida
lobis spinosodentatis; folia supra albo cinereo-viridia eglandulosa
arachnoidea, subtus propter tomento denso arachnoideo feré alba,
eglandulosa. Calathidia duo-tria 3.5—-4 cv diam., periclinium universale
e setis membranaceis 6-8 mm long. reflexum, cauli adpressum;
receptaculum pyriforme vel obovatum 4—6 mm long.; capitula sive
calathidia partialia 14-15 mm long., penicilli setae plures planiusculae
8—9 mm long.; involucelli squamae 16—20 tenuissime scabridae pallidae,
extimae—9-10 mm long. membranaceae subfiliformes apice
cochleariformes dilatatae, fuscae, dentatae, utrinque 2-3 cilias gerentes;
squamae duarum serierum sequentiun 12-14 mm long. carinatae, superne
acuminatae margine ciliis adpressis, interne fuscae; squamae intimae
inferne triente coalitae, interne atrofuscae vel nigricantes apice liberae
acuminatae fimbriatae; tubus corollae superne minus glandulosus;
achenium fusiforme dense pilosum, pilis flavidis appressis pappum
superantibus.

Affinis est E. albicauli Kar. et Kir. sed notae singulares habet: folia
inferiora pinnatisecta lobis 3—4-paris suboppositis; calathidia partialia

14-15 mm long., penicillus 8—9 mm long; tubus corollae superne minus

glandulosus.

“Reproduced from the Russian Original—General Editor.

715

708

Habitat in pistaceis semidesertis.

Area geographica: Asia media, Tadzhikistania australis.

Typus: Kafirnigan, in decliviis inter fontes Ajchatyn et Czilczik
5 x 1931 leg. A. Butriakov, conservatur in Leningrad. .

2. E. tjanschanicus Bobr. n. sp. (sect. Ritrodes Bge.).

Radix 2—3-cephala; caules in numero 1-3 basi petiolis vetustis vestiti
usque ad apicem dense arachnoidei vel laxe albotomentosi, superne
ramulo uno vel duobus; folia coriacea superne incana vel incano-viridia
laxe arachnoidea glandulis sessilibus minutis praedita, petiolis stipitati-
glandulosis, subtus propter pubescentiam densam arachnoideam fere
albotomentosa foliola inferiora petiolata lanceolata usque ad 20 cm long.,
3-4 cm lat. in parte superiore profunda pinnatipartita lobis lanceolatis -
spinoso-dentatis rachide angusti sinuati, lobi foliorum in parte inferiore
laminae deminuti fere aculeiformes; folia caulina sessilia amplexicaulia,
basi spinosa, superiora lanceolata acute spinoso-dentata. Calathidia 3-4
cm diam. caesia; capitula seu calathidia partialia 14-15 mm long.;
penicilli setae inaequales, 3-4 mm long.; involucri phylla (involucelli
squamae) in numero 20—24 libera caesia scabra superne paulo reslinata;
plylla extima setas duplo longiora lanceolata acuminata carinata ciliata
ad intimas sensim elongata plus attenuata; phylla intima longior ciliata,
medio cohaerentia atrofusca; corolla coerulescens profunde partita tubo
glanduloso pilis brevibus non numerosis tecto; achenium pilis appressis
dense tectum; pappi setae scabrae inaequales superne liberae. VII—VIII.

Affinitas. Ab E. dasyantho Rgl. et Schm. differt foliis superne
laxe arachnoideis insuper glandulis minutis sessilibus tectis, petiolis
glandulis stipitatis praeditis, tubo corollae pilis brevibus non numerosis
tecto.

Habitat in steppis montanis et in Juniperetis.

Area geographica: Asia media, in jugis australibus Tjanschanicis.

Typus: in decliviis australibus jugi Kuramensis, in fissura Babijjab
superior, junipereta, 2200 m s. m. 27 VII 1935 n° 573, leg. Emme-
Markovskaja, conservatur in Leningrad.

JURINEA CASS.
Sectio Cyanoides (Korsh.) Ijin.
3. Sec. Cyanoides (Korsh.) Iljin sect. nov.—Jurinea §2. Longifoliae
DC. Prodr. VI (1837) 674 p. p., §3. Decurrentes DC. Ibid. 675 p. p. and

§4. Pinnatilobatae DC. Ibid, 675 p. p.—Sect. Platycephalae Benth. and
Hook. f. Gen. Pl. II, 1 (1873) 473 p. p—Jurinea §3. Pinnatae Boiss.

716

709

Fl. or. III (1875) 572 p. p.; Hoffm. in Pflanzenfam. IV, 5 (1894) 321 p.
p.—Stirps J. cyanoides Korsh. in Izv. Akad. Nauk, 1, 2 (1894) 8; Iljin
in Izv. Bot. Sada Akad. Nauk SSSR, 30, 3-4 (1932) 348.—Group
Cyanoides Klok. (non Iljin) in Bot. Zhurn. URSR, 8, 1 (1951) 47.—
Sect. Platycephalae Benth. and Hook. subser. Cyanoides Sosn. in Zhurn.
Russk. Bot. Obshch. 11 (1926) 196.

Pappus nunquam deciduus, setis serrulatis vel serrulato-barbellatis,
basi in anulum non connatis. Achenium obpyramidatum, quadrangulum,
ad basin attenuatum, glabrum, sublaeve vel paralleliter striatum,
coronulam veram non formans, attamen supra margine denticulatum,
pappogeno composito, setis pappi incumbentis, nectario quinquedentato
coronato instructum. Plantae semper herbaceae, perennes, caulibus.
foliatis, saepe alatis foliis diversiformiter pinnatipartitis, interdum integris,
supra plus minusve viridibus, subtus canescenti-albotomentosis.

Sectio nova ad sectionem Bellae Iljin approximata est, attamen
Species omnes sectionis omnes sectionis Bellae statura humiliore, caulibus
semper scapiosis, eramosis sat distant.

Typus sectionis: Carduus cyanoides a. monoclonos L.

Area geographica: Europa, Asia occid.

4. J. schischkiniana Iljin sp. nov.—J. cyanoides Schischk. (non
DC.) in Krylov, Fl. Zap. Sib. XI (1949) 293 p. p.—J. pollichii Ldb.
(non DC.) FI. Ross. II (1844-1846) 764.—Serratula cyanoides Bge. (non
DC.) in Ldb. FI. alt. TV (1833) 42 p. p.—dJ. albicaulis x J. cyanoides
Schischk. op. cit. 2933.

Perennis, 30-75 cm, caulibus erectis, pauciramosis vel simplicibus,
foliis radicaulibus et caulinis inferioribus petiolatis, reliquis caulinis
sessilibus, supremis diminutis integris, omnibus non decurrentibus,
pinnatisectis, segmentis oblongo-lanceolatis vel linearibus per rachidem
decurrentibus et basi confluentibus. Capitula grandiuscula, 20-35 mm
in diam. involucro 15-20 mm alt., manifeste arachnoideo, squamis
omnibus depressis. Corolla roseo-purpurea, 14—21 mm lg. limbi laciniis
5—7 mm lg. Pappus e setis serrulato-barbellatis constans. Achenia 5—
6 mm lg.

Area geographica: Sibiria altaica et Kazachstania altaica.

Typus: Dit. Barnaul. Inter pagum Solonovka ad ripam fl. Kamsala
et Malyschevy Log. in pinetis, 23 VI 1913, leg. V. Reverdatto. In Herb.
Inst. Bot. Ac. Sc. URSS (Leningrad) conservatur.

5. J. xerophytica [ljin sp. nov.

Perennis, 20-60 cm alt., rhizomate verticali longo (in arenis) vel
valde abbreviato, saepe ramoso, squamis brunneis obtecto, in axillis
glomerulis gossypinis instructo; caules basi atque petioli foliorum

717

710

inferiorum dense albo-tomentosi, solitarii vel numerosi, reliquorum
floccoso-arachnoidei vel fere glabri (f. glabrescens Iljin), tenuiter striato-
sulcati, striis flavidis et purpureis, rarius fere ex toto purpurascentes,
sulcis viridulis, caules plus minusve flexuosi vel erecti, a basi vel supra
basin, interdum in parte superiore ramosi; folia rigida, radicalia et caulina
inferiora petiolata pinnatipartita vel pinnatisecta, media et superiora
sessilia, segmentis oblongis vel linearibus acuminatis seu acutis,
integerrimis, interdum lyratopartita vel raro segmentis basi lobatis vel
dentatis, omnia vel inferiora tantum integra et integerrima, superiora
diminuta, supra omnia virescentia, tenuiter arachnoidea vel glabra,
rugulosa, asperiuscula, glandulis sessilibus instructa, margine anguste,
interdum ad nervos primarios revoluta, subtus albo-tomentosa, sub
tomento glandulosa; capitula mediocria vel parva, multiflora 15-25 mm
diam., involucro 10-15 mm alt. plus minusve arachnoideo, phyllis ut in
J. cyanoide dispositis, corolla roseo-purpurascens, pauciglandulosa (11—
12) 16-20 mm alt., laciniis limbi (4) 5—6 mm lg.; pappus niveus (10)
12-15 mm alt. setis longioribus breviter barbellatis, barbellis plus-minus
dense dispositis; achenia (4-5) 6 mm lg., ferruginea, opaca
longitudinaliter striata, superne margine denticulata, apice pappophoro
instructa.

Area geographica: Partes meridionales et orientales Kazachstaniae
centralis.

Typus: In dit. Turgai, in valle sicca (log) Aksaj in systemate fl.
Sarysu, 9 VI 1941, leg. M. Spiridonov. In Herb. Bot. Inst. Ac. Sc.
URSS (Leningrad) conservatur.

6. J. gorodkovii Iljin sp. nov.

Perennis, 20—50 cm alt., caulibus solitariis vel paucis, tenuiter
arachnoideis, simplicibus vel paulo ramosis, foliis oblongo-linearibus,
acuminatis, integris, integerrimis, raro segmentis solitariis instructis,
supra viridibus, subtus albo-tomentosis, radicalibus usque ad basin
gradatim attenuatis, caulinis sessilibus, semiamplexicaulibus, anguste
decurrentibus; capitula parva vel media raro 20 mm in diam., involucro
10-15 mm lg., tenuiter arachnoideo, phyllis omnibus vel solum
exterioribus apice herbaceis, tenuiter linearibus, laxis vel solum basi
appresis, internis erectis, acuminatis; corolla rosea vel roseo-purpurea,
13-15 mm lg., laciniis limbi 4-4.5 mm lg. Pappus niveus, setis serrulato-
barbellatis; achenia 4.5-5 mm lg., oblonga, obpyramidata, margine
denticulata.

Area geographica: Sibiria occid. (in ditione Tobol).

Typus: Sibiria occidentalis in ditione Jalutorovsk, pineta cladinosa
in Padunskaja 18 VIII 1912, n° 1850 leg. Sviticz. In Herb. Inst. Bot.
Ac. Sc. URSS (Leningrad) conservatur.

718

711

7. J. creticola [ljin sp. nov.

Perennis, 15-30 cm alt.; caules erecti, rarissime decumbentes,
floccoso arachnoidei, striati, ramosi, raro fere simplices; folia radicalia
atque caulina inferiora petiolata petiolis albo-tomentosis, laminis
simpliciter pinnatisectis, rarissime fere bipinnatisectis, segmentis anguste
linearibus acuminatis vel acutis, remotis, margine manifeste, saepe ad
nervos medianos impressos revolutis paucis, caulina media et superiora
sessilia, alatim vel angustissime alatim decurrentia, supra rugulosa
glandulis sessilibus inter rugas dispersis, vix exasperata subtus
albotomentosa, sub tomentulo glandulosa, superiora et suprema integra
et integerrima linearia et filiformia; capitula apice caulis et ramorum
solitaria, mediocria, 20-30 mm in diam., multiflora, involucro ca. 12—
15 mm alt., phyllis exterioribus et mediis arachnoideis, acuminatis, laxe
et perplexe dispositis ut in Jurinea cyanoide intimis rectis in parte
superiore purpurascentibus, paleaceis, acuminatis, margine ciliatulis, a
tergo vix conspicue hirsutiusculis; corolla roseo-purpurea 16-18 mm alt.,
laciniis limbi 5-6 mm lg.; pappus niveus, setis breviter barbellatis,
barbellis densissime dispositis; achenia immatura striata.

Typus: In vicinitate Chvalynsk (distr. Saratov), arenae cretarum
“Pesczanaja Gora”, prope “Sosnovaja Myza” 25 VI 1926 n° 1044 leg.
K. Grosset. In Herb. Bot. Inst. Ac. Sc. URSS (Leningrad.) conservatur.

8. J. altaica [jin sp. nov.

Planta 25-40 cm alt., caulibus fere a basi ramosis, floccoso-
arachnoideis; folia radicalia et caulina inferiora in petiolum attenuata,
pinnatisecta, segmentis oblongo-linearibus, acuminatis vel-mucronatis,
cauline reliqua sessilia, sat late et longe decurrentia, superiora integra
et integerrima diminuta, supra canescenti-virescentia, tenuiter arachnoidea
subtus albo-tomentosa; capitula mediocria inflorescentiam corymbosam
racemosam formantia involucro arachnoideo, plyllis exterioribus

- retreflexis canescentibus; corolla purpurascentia 14-15 mm lg., lacintis
-limbi circiter 4 mm lg.; achenia supra marginem vix denticulata.

Area geographica: Sibiria altaica.
Typus: Sanatorium Aul (Krasnyi Aul), arenae ad marginem pineti.
prope stationem viae ferreae-Aul. 19 VIII 1932, leg. V. J. Veretschagin.

In Herb. Inst. Bot. Ac. Sc. (Leningrad) conservatur.

9. J. transuralenis [Ijin sp. nov.

Perennis, 15—70 cm alt., caulibus erectis, floccoso-arachnoideis vel
albo tomentosis, interrupte alatis, ramosis, rarius simplicibus; folia
radicalia numerosa, petiolata, pinnatisecta, segmentis linearibus
acuminatus vel acutis, integerrimus margine anguste revolutis, rarissime
integra (f. integrifolia), caulina sessilia basi dilatata, auriculata et

WAS

712

manifeste decurrentia atque pinnatisecta, apicem versus diminuentia
integra, omnia supra viridia vel vix canescentia, glabra vel tenuiter
arachnoidea, subtus albo-tomentosa; capitula parva vel mediocria,
involucro 10-15 mm alt., glabro vel arachnoideo, phyllis externis
gradatim in aristas elongatas, glabras, coloratas saepissime virides
abeuntibus horizontaliter arcuato-inclinatis, internis erectis, apice
palaeaceo-acuminatis; corolla roseo-purpurascens, 12—15 mm alt., laciniis
limbi 4-5 mm lg.; pappus niveus (10) 12-15 mm lg., setis serrulato-
barbellatis; achenia vix striata, costulata, margine superiore denticulata,
apice pappophoro composito instructa.

Area geographica: Kazachstania transuralensis.

Typus: Dit. Aktjubinsk. In systemate fl. Uil, prope stationem
Aschkinau (via ferrea Gurjev-Kandagacz), prope urbem Temir, steppa
arenosa, stipeta agropyreta, 24 VI 1955, n° 61, leg. V Grubov et N.
Lubarsky. In Herb. Bot. Inst. Ac. Sc. URSS (Leningrad) conservatur.

10. J. kazachstanica [ljin sp. nov.

Perennis, 20-60 cm alt., rhizomate longo, squamis brunneis obtecto.
caulibus erectis, raro simplicibus, plerumque supra medium ramosis,
floccoso-arachnoideis, interrupte alatis; folia radicalia numerosa, atque
caulina inferiora petiolata, pinnatisecta, segmentis linearibus, elongatis,
acuminatis, integerrimis, remotis, laminis rarius integris (f. integrifolia),
reliqua caulina sessilia, late vel anguste decurrentia, suprema diminuta
integra, omina supra viridia vel canescenti-virescentia, tenuiter
arachnoidea, subtus albo tomentosa; capitula numerosa parvula, 10—12
mm in diam., multiflora, involucro 10-15 mm alt., arachnoideo, floccoso,
phyllis externis et mediis apice vix bamatis, internis erectis, apice
paleaceis, corolla rosea vel roseo-purpurea (10) 12-16 mm alt., laciniis
limbi 3-6 mm lg.; pappus niveus 10-12 mm alt., setis serrulato-

barbellatis, dense dispositis; achenia 5-6 mm lg.

Area geographica: Kazachstania occid., arenae mobiles (Tajsugan,
Kok-dzida, Barsuki et al.).

Typus: Dit. Uralsk, in reg. Gurjev. Arenae grumosae Tajsugan, n°
297, 15 VI 1927, leg. M.M. Iljin et J.S. Grigorjev.

11. J. pineticola Iljin sp. nov.

J. pollichii Ldb. (non DC) FI. Ross. II, 2 (1845-1846) 764 p. p.—
Serratula cyanoides Bge. (non DC.) in Ldb. FI. alt. IV (1833) 4 p. p.—
Jurinea polyclonos Korsh. (non DC.) in Izv. Akad. Nauk 1, 2 (1894)
119; J. cyanoides x polyclonos Korsh. Ibid. 121 (specim. e Loktjewsk,
Irtysch and Semipalatinsk).—J. albicaulis Iljin (non Bge.) Obz. Turk.
Vidov Roda Jurinea Cass. (1925) 20 p. p.—J. albicaulis x cyanoides
Schischk, in Krylov, Fl. Zap. Sib. XI (1949) 2933.

720

713

Perennis, 15—35 cm alt., radice dura in rhizomata horizontalia vel
verticalia squamis brunneis et reliquis foliorum vetustorum obtecta
transeunte; caules albo-tomentosi, rarius floccoso-arachnoidei, erecti vel
decumbentes, plerumque ramosi; folia radicalia numerosa, integra et
integerrima, acuminata, usque ad basin sensim in petiolum brevem
attenuata vel epetiolata, interdum nonnula-pinnatipartita, vel pinnatisecta,
lobis bijugis vel trijugis, caulina integra, sessilia non decurrentia vel vix
decurrentia vel solum semiamplexicaula et auriculata, raro manifeste
decurrentia, suprema diminuta, omnia supra viridia et virescentia, tenuiter
arachnoidea, subtus albo-tomentosa; capitula parva, 10-12 mm in diam.,
solitaria vel apice caulis et ramorum bina-terna, involucro 10-12 mm
alt. arachnoideo phyllis exterioribus appressis vel apice praecipue
fructificatione horizontaliter retrofractis, internis erectis. acuminatis, in
parte superiore paleaceis; corolla rosea vel roseo-purpurascens, interdum
ochroleuca (10) 13—15 mm alt., laciniis limbi 3-5 mm lg.; pappus 10—
12 mm lg., niveus barbellatus; achenia (4.5) 5—6 mm lg.

Area geographica: Kazachstania, pineta distr. Semipalatinsk.

Typus: Prope pagum Schulbinskaja, arenae mobiles, 7 VII 1930, n°
23, leg. U. Winter. In Herb. Bot. Inst. Ac. Sc. URSS (Leningrad)
conservatur.

12. J. tanaitica Klok. sp. n.

Planta perennis rhizomate brevi et surculis sterilibus evolutis. Caules
saepissime solitarii ca. 35-55 cm alti, inferne ob pubescentiam albidi,
superne flocculoso-arachnoidei, apice ramosi. Folia radicalia petiolata
ca. 10-25 cm longa, petiola lamina breviore, pinnatisecta, segmentis
lateralibus utroque 3—5, subtus albitomentosa, supra laete viridia,
glabriuscula et laevia; foliorum segmenta linearia oblongaque acutata,
margine revoluta et manifeste crispula; segmentum terminale usque ad
10 cm longum et 7 mm latum, lateralia ca. 2-6 cm longa et 2-5 mm
lata; folia caulina inferiora radicalibus minora, usque ad 10 cm longa;
media trifida segmentis angustis, superiora anguste linearia, diminuta;
caulina exceptis inferis decurrentia, in caule alas usque ad 5—6 cm longas
et 5—7 mm latam formantia. Calathidia numerosa (15-26) in pedunculis
usque ad 6—7 cm longis, inflorescentiam paniculato-corymbosam
formantia; calathidium floribuscum ca. 17 mm longum; involucrum
crateriforme ca. 10-11 mm longum, inferne ob pubescentiam tenuiter
tomentoso-arachnoideam in phyllis externis cinereum, superne sordido-
purpureun ob phylla interna colorata; involucri phylla oblongo-vel
lineari-lanceolata, ca. 2-11 mm longa et 0.7—1.4 mm lata, acuminata,
omnia adpressa; corollae flosculorum ca. 13-15 mm longae, roseo-
purpureae. Achenia cuneata, 4-5 mm longa, ca. 2.5 mm lata, laete
brunne 1; pappus ca. 10 mm longus, setis longissimis 5, albidus.

721

714

Habitat in sabulosis tanaiticis.

Affinitas. Ab J. amplexicauli (S. G. Gmel.) Bobr. phyllis involucri
adpressis, a J. thyrsiflora Klok. caulibus late alatis primo aspectu
abhorret.

Typus: Ditio Kamenskiensis, prope pag. Oblivskaja, in sabulosis ad
fl. Czir, 25 VIII 1928 fl., A. Gael. In Herbaria Instituti botanici Ac.
Sci. RSS Ucr. Kioviae conservatur.

13. Sectio Bellae Iljin sect. nov—§5. Subacaules Boiss. (non Benth.)
FI. or. Ill (1875) 582 p. p.; praeter J. frigidam Boiss. and J.
microcephalam Boiss.

Perennis, scaposa, non raro capitulis sessilibus, foliis rosulatis; setis
pappi denticulatis vel barbellato-denticulatis in anulum non concretis;
achenia parvula 2.5-3 (4-5) mm lg., glabra, + laevia, plerumque
longitudinaliter striata, supra margine denticulata, apice pappophoro
composito instructa.

Plantae alpigenae in Caucaso et Asia occidentali distributae.

Typus sectionis: J. cadmea Boiss.

14. J. venusta Iljin sp. nova.

Planta 25—30 cm alt., caulis scapiformis, solitarius, tenuiter striatus,
vix laxe arachnoideus. Folia rosulata in petiolum sensim attenuata,
pinnatisecta, segmentis oblongo-linearibus, obtusatis, interdum vix
mucronatis, integerrimis 2—3 mm lat., basi dentibus vel lobis brevibus
et obtusatis instructis, margine anguste revolutis, rachide aequilatis,

oblique sursum vel rarius fere horizontaliter vergentibus, sinulis segmentis

triplo-quadruplo latioribus, omnia utrinque pallide viridia, glabra, supra
scabrida, nervis impressis, parce glandulosa, subtus dense glandulosa,
nervo medio pallido, prominente. Capitula p. m. magna, circiter 30-35
mm diam., mutiflora; involucrum cupuliforme, 18-22 mm alt., basi vix
arachnoideum vel glabrum, phyllis subaequilongis, laxe dispositis,
flexuosis intenis + erectis. Corolla rosea. purpurascens, 12-13 mm lg.
5—6 mm lat., laciniis limbi 4—5 mm lg. Pappus circiter 10-12 mm alt.,
fumosus, setis scabrido-barbellatis. Achaenia circiter 5 mm lg., oblonga
compresso obpyramidata, pallida, glabra, paralleliter striata, in parte
superiore denticulato-marginata dentibus inaequalibus, apice pappophoro.

Area geographica: Caucasus septentrionali-occidentalis (in regione
Krasnaja Poljana).

Haec species ad Jurineam Iljinii Grossh. valde accedit, attamen J.
Iljinii Grossh. rosula foliorum medium vel supra medium scapi attingente,
foliis angustioribus, 5-15 mm lat., segmentis 3-10 mm lg., omnibus
integerrimis + acuminatis vel acutis, manifeste sursum vergentibus,
phyllis involucri appressis sat distat.

715

Typus: in parte superiore fluminis Malaja Laba. In pratis lapidosis,
18 VIII 1936, leg. L. Vassiljeva. In Herb. Inst. bot. Acad. Sc. URSS
(Leningrad) conservatur.

15. Sectio Integrae Iljin sect. n.

Caules plus minus alati, folia integra integerrima nonnulla segregata
rarissime teriata, decurrentia; achenia glabra laevia, saepe striata vel
jugata laeviterque striata, sub coronula denticulata; pappus cum
pappogeno caducus (J. kokanica, J. rhizomatoidea et J. trifurcata
exclusae).

Plantae praecipue tjanschanicae; in flora nostra species 12 adsunt.

Typus sectionis: J. Abolinii Ijin.

16. Sectio Suffrutices [ljin sect. n.

Suffrutices foliis integris integerrimisque linearibus vel filiformibus;
involucri phylla appressa, apice tantum interdumque reclinata; pappus
crenatus vel crenato-barbullatus non caducus (J. Sintenisii exclusa);
achenia glabra laevia saepe longitudinaliter striata vel plicata;
pappogenum semper adest; pappophorum tantum J. Sintenisii habet.

Typus sectionis: J. mugodsharica jin.

Sectio Penduliflorae Iljin

17. Sectio Penduliflorae [ljin sect. n.

Flores marginales nutantes; corolla basin sensim angustata, rosea
vel purpurea glandulis sparsis sessilibus praedita. Pappus non caducus
(J. Komarovii, J. elegantissima, J. Kamelinii exclusae). Achenium
magnum glabrum costis teretibus parallelis parum conspicuis praeditum.
Pappophorum vulgo non evolutum, pappogenum calycinum, nectarium
satis magnum, manubrio sat breve.

Sectio habet quindecim species in flora URSR.

Typus sectionis: J. robusta Schrenk.

18. J. kamelinii [jin sp. n.

Planta 25—40 cm. alta; caules solitarii vel pauci canaliculati costis
albescentibus glandulis sessilibus dense et pilis -longiusculis flexuosis
tectis, apice arachnoidei; folia in dimidio inferiore aggregata, radicalia
ad basin angustata anguste-lanceolata, superiora linearia deminuta sessila
omnia integerrima acuminata utrinque viridia dense pilosa glandulis
sessilibus plurimis tecta supra venula impressa, folia suprema auriculata.

Calathidia solitaria multiflora 25-35 mm in diam.; involucrum 15-20
mm alt., involucri phylla appressa oblonga vel anguste lanceolata

VED

716

acuminata dorso glandulis dispersis munita albo-marginata, phylla interna

_longe acuminata purpurascentia dense glandulosa margine ciliata; corolla

24-27 mm long., rosea lobis 5-6 mm long.; pappus albus nectario
quinquelobato; achenium maturum 7 mm long. glabrum longitudinaliter
lineatum apice spinellosum, coronula inaequidentata pappophoro praedita.

Habitat in montibus Pamiroalaicis.

Affinitas. Haec species J. Komarovii valde affinis differt tamen pappo
partim caduco, pubescentia pilosa nec papillosa.

Typus: Jugum Seravschanicum, in decursu medio fluminis Seravschan
18 VI 1960 n° 568 leg. R. Kamelin; typus in herbario Leninopolitano
conservatur.

19. J. elegantissima [ljin sp. n.

Planta 50-60 cm alta usque a basi ramosa viridis parum arachnoidea
glandulis numerosis sessilibus ramis tenuibus; folia linearia integerrima
sessilia non decurrentia utroque latere unicolora reclinata. Calathida
solitaria, involucrum 20-25 mm long., phyllariis appressis acuminatis
glabris viridibus externis valde deminutis internis longe attenuatis; corolla
roseola 25—27 mm long., lobis 6-7 mm long.; pappus caducus; achenium
oblongum 9-10 mm long. costis elongatis apice denticulatum.

Habitat in gypsaceis pamiroalaicis.

Affinitas. Ab J. robusta Schrenk foliis linearibus integerrimis non
decurrentibus differt.

Typus: Pamir-Alaj, fontes fluminis Jakkabag, in vicinitate pagi
Taschkurgan, in gypsaceis 4 VII 1936 n° 624, leg. V. Botschantzev et
A. Butkov; typus in herbario Leninopolitano conservatur.

20. J. botschantzevii Ijin sp. n.

Planta vulgo 25—40 cm alt., caulibus furcatis arachnoideo-tomentosis
ad dimidium longitudinis ramosis foliosisque, ramis basi foliatis; folia
pinnatipartita lobis elongatis obtusis caulinis decrescentibus utrinque
arachnoideis glandulosisque, folia inferiora petiolata, superiora sessila.
Calathidia solitaria multiflora involucrum acetabuliforme 20-25 mm alt.
pubescens glandulis dispersis praeditum, phylla interna elongata
acuminata, media—lanceolata, interna—longe acuminata glandulosa et
paulo setosa; flosculis roseis disperse glandulosis 22-30 mm long. lobis
5—7 mm long.; pappus non caducus, pappo setis barbellatis tenuissimis;
achenium vix lutcolum tetragonum obpyramidatum longitudinaliter
striatum 6 mm long. apice dense dentatum; pappogenum quinquelobatum,
nectarlum magnum minute quinquedentatum.

Habitat in montibus Pamiroalaicis in declivibus cretaceis.

Affinitas. Ab J. bucharica Winkl. differt caulibus magis robustioribus
foliis caulinis sessilibus.

723

VAD

Typus: Tadshikistania australis, montes Kojkitau in vicinitate pagi
Ajvadzh, in decliviis schistosis 17 V 1960 n° 46, leg. V. Botschantzev
et T. Egorova; typus in herbario Leninopolitano conservatur.

21. J. winkleri [jin sp. n.

Haec species e serie Pinnatae Iljin in primis ab J. Botschantzevii
Iljin differt foliis non undulatis integris vel lyratis, ab J. tadshikistanica
Iljin differt pubescentia evanescente, laminis foliorum minus rugosis
densius glandulosis, glandulis non impressis, venula minus impressa.

Habitat in promontoriis regionis Ferganensi.

Typus: Prov. Fergana distr. Margelan, prope pagum Assake, 4 V
1913, n° 426, leg. N. Dessjatova; typus in herbario Leninopolitano
conservatur.

22. J. tadshikistanica Iljin sp. n.

— A praecedente J. Winkleri I\jin differt pubescentia crispula regulari
laminis foliorum alde et minute rugosis, glandulis dispersis profunde
dispositis, venula distincte impressa, pappi setis raro breviterque
barbellatis setulis semiappressis.

Habitat in promontoriis vallis Vachsch.

Typus: Tadshikistania, in valle fluminis Vachsch, in decliviis
schistosis montis Buritau, 3 V 1937, n° 519, leg. V. Nikitin; typus in
herbario Leninopolitano conservatur.

23. Sectio Insculptae

Folia varie incisa decurrentia, pappus ex toto cum pappogeno et
nectario deciduus (J. asperifolia, J. serratuloides, J. hamulosa et J.
litophila exclusae); achenia glabra vel sub coronula denticulata apice
nectario mitriformi.

Typus sectionis: J. maxima Winkl.

24. J. zakirovii Iljin sp. n.

Planta ad 50 cm alt., rhizomata squamis dense obtecta, in axillis
superioribus globulis gossypino-villosis instructa. Caules pauci, erecti,
ramis paucis elongatis, pro more solitariis, cylindrici, manifeste sulcato-
costati, glandulis sessilibus parvulis et pilis crispatis brevioribus -albidis
obtecti, sub capitulo solum vix arachnoidei in dimidio et triente superiore
aphylli. Folia radicalia 10—15 cm longa in petiolum basi dilatatum sensim
transeuntia, oblonga, pinnatilobata vel pinnatidentata, lobis et dentibus
oblique oppositis et sursum vergentibus, terminalibus manifeste elongatis,
gradatim acuminatis, apice cartilagineo-mucronatis, margine anguste
revolutis; caulina sessilia, decurrentia, integerrima vel rarius dentibus
triangularibus solitariis et binis instructa, omnia utrinque concolora,

724

718

opaco-viridio, supra nervo medio. impresso instructa, breviter crispule
hirta, rugulosa et glandulis sessilibus minimis obsita, subtus nervo medio
prominenti et nervis lateralibus conspicuis craspedodromis donata, pilis
crispatis longioribus et glandulis sessilibus obtecta. Capitula apice caulis
et ramorum singularia, multiflora; involucrum 15-20 mm alt., basi ad
caulem impressum, flavido-viride, phyllis quadriseriatis externis et mediis
oblongo-lanceolatis, in parte superiore acuminatis et vix arcuatis, nervis
manifestis percursis, intimis in arista longa abeuntibus; omnibus margine
ciliatis, extus glandulis sessilibus numerosis parvulis cum papillis minimis
intermixtis obsitis; receptaculum paleis involucro triplo brevioribus.
Corolla rosea, 16-18 mm alt. laciniis limbi 4-5 mm lg., dense glandulosa;
filamenta antherarum vix tuberculata. Pappus deciduus, setis in anulum
connatis, barbellatis. Achaenia obpyramidata, quadrangula, ca. 5 mm
Ig., glabra et laevia, longitudinaliter striata, coronula aequaliter dentata,
cuius in parte intériore pappogenum conoideum apice nectario coronatum,
in anulum pappi penetrans observatur.

Affinitas: Specis haec nova ad J. helichrysifolia M. Pop. appropinquat
sed foliis decurrentibus, concoloribus differt.

Typus: Asia Media, Usbekistania, jugum Nuratau, in circumjectus
Jangokly, prope Langar, 19-21 V 1941, leg. K. Zakirov. In Herb.

Univers. Asiae Mediae (Taschkent) conservatur.

25. J. helichrysifolia M. Pop. in Tr. Uzbeksk. Univers., Nov. Ser.
27 (1941) 105 (nomen nudum).

Planta 20-40 cm alt., radice valde fibrosa in rhizomate verticali
transeunte, apice polycephala; tenuiter tomentosa. Caules pauci, erecti
simplices, flavescenti-virescentes, cylindrici, striati, pilis brevibus crispatis
et glandulis sessilibus obtecti, non arachnoidei, sub capitulo solum vix
arachnoidei. Folia radicalia in petiolum longum, basi dilatatum sensim
transeuntia, oblongo-linearia, ad 10-20 cm lg., ca. 5 mm It., margine
vix revoluta, caulina sessilia, anguste decurrentia, usque ad medium caulis
vel vix supra vel infra disposita, omnia linearia, integra et integerrima,
cauli semiappressa, mucronata, mucrone cartilagineo, supra nervo medio
impresso percursa, manifeste rugulosa, punctato-foveolata, propier papillas
brevissimas asperula, vix floccose-arachnoidea, margine ‘valde revoluta
plerumque ad nervum medium, glandulis in foveolis assidentibus
instructa, subtus magis arachnoidea, suprema diminuta, filiformia.
Capitula apice caulis solitaria, mutiflora; involucrum cupuliforme, ca.
20 mm alt., phyllis imbricatis appressis, externis oblongis, flavescentibus,
apice obscure viridibus, breviter mucronatis, nervis vix conspicuis mediis
lanceolatis longe acuminatis, in parte superiore sordide purpurascentibus
nervis prominentibus, intimis in aristam gradatim elongatis, omnibus
extus glandulis sessilibus instructis et vix araneosis, margine minime

TS

ciliolatis. Corolla roseo-purpurea, glandulosa, plerumque 17-18 mm alt.,
laciniis limbi 4-5 mm lg. Pappus niveus, barbellatus. Achaenia immatura
glabra longitudinaliter vix plicatula, coronula denticulata.

Affinitas: Haec species ad J. Zakirovii Iljin accedit, attamen HONS

* discoloribus, necnon phyllis externis involucri sat recedit.

725

Typus: Asia Media. Ad radices montium jugi Hissar in ditione
Kschtut, in ripa fl. Negnot, leg. M.G. Popov. In Herb. Univers. Asiae
Mediae (Taschkent) conservatur.

Sectio Stechmannia Boiss.

26. J. spiridonovii [Ijin sp. n.

Species ut videtur e sectione Stechmannia; suffrutex 8-15 cm alt:
caudice polycephalo rhizomate brevi; caules simplices; folia in fasciculos
congesta, inferiora elongato-linearia in petiolo angustata ima basi vix
dilatato, integerrima rigida rugosa supra venula impressa, infra
albotomentosa sub tomento glandulosa, folia caulina adsunt. Calathidia
parva, involucrum ca 10 mm alt. acetabuliforme, phyllaria appressa
subglabra, externa—ovata, media—elongato-ovata, interna—longiora
apice membrancea; receptaculum paleis tenuibus longis laevibusque
tectum; corolla pappi et achenia desunt.

Typus: Kasachastania in valle fluminis Selecta non procul a sepulcro
Tassmola 7 VIII 1929, n° 2280 leg. M. Spiridonov; planta in herbario
Leninopolitano conservatur.

Sectio Nanae [lyin

27. Sectio Nanae Iljin sect. n.

Suffrutices montani humiles acaules vel caulibus nanis, foliis integris
vel tenuiter fissis, praesertim rosulatis, pappus barbatus crenatusque,
semper caducus, pappogenum pappophorumque semper evoluta; achenium
glabrum saepe costis laevibus.

Typus sectionis: J. Abramovii Rgl. et Herd.

28. J. bobrovii Iljin sp. n.

Suffrutex ramulis prostratis apice dense foliatis; folia in fasciculos
congesta; caules 15—25 cm alt. striati flocculoso-arachnoidei stricti inferne
foliati, folia inferiora fasciculataque lanceolata vel elongata emarginata
vel dentibus raris praedita, basi angustata apice acuminata; folia caulina
angustata sessilia; omnia supra cinereo-viridia rugosa glandulis profunde
sessilibus venula prominula. Calathidia solitaria, involucrum patelliforme

726

720

inferne arachnoideum, phyllaria externa acuminata apice arcuatim reflexa,
media—minus reflexa, interna—erecta membranacea longe acuminata,
omnia venula prominula, margine serrulata; corolla 14-15 mm long.
lobis 4—5 mm long. glandulis dispersis; pappus caducus setis coalitis
crenulatis pappogenum patelliforme lobulatum inferne subalbidum interne
rubrum in centro pappophorum gerens, achenia 2—2.25 mm long.
longitudinaliter lineata apice coronata e dentibus membranaceis
aequalibusque.

Habitat in decliviis montosis Turcomaniae occidentalis.

Affinitas: Ab J. Abramovii Rgl. et Herd. foliis integris
subintegerrimisque, ab J. Knorringiana Iljin foliis lanceolatis inferne
albotomentosis, pappo setis crenulatis, achenio minore differt.

Typus: Montes Kopetdagh in declivi occidentali montis Chossardagh
VII 1928 leg. E.G. Bobrov; planta in herbario Leninopolitano asservatur.

Sectio Molles Korsh.

29. J. kasakorum [jin sp.nov.

Planta ad 50 cm alt. Caules pauci et verosimiliter singulares, erect,
simplices, vel breviter et paulo ramosi, longitudinaliter striati, virescentes
vel sordide purpurascentes, floccoso-arachnoidei, glandulis sessilibus
parvulis instructi. Folia fere omnia in parte inferiore caulis contracta,
radicalia numerosa brevipetiolata, caulina sessilia, loco insertionis dilatata,
semiamplexicaulia et denticulato-decurrentia, omnia pinnatisecta, lobis
linearibus, rarius oblongis, remotis, emarginato-dentatis, acuminatis,
margine valde revolutis et semper crispatis, cum rachide folii anguste
alato, partim denticulato confluentibus, superiora simpliciuscula et
diminuta, omnia supra canescenti-arachnoidea, vix rugulosa, non asperula,
glandulosa, subtus laxe albo-tomentosa, sub tomento glandulosa. Capitula
apice caulis solitaria multiflora; involucrum late cupuliforme, basi
impressum, ca. 20 mm alt., phyllis extimis et mediis appressis,
triangulato-lanceolatis valde arachnoideis, glandulosis, apice breviter
calloso-acuminatis e tomento vix emergentibus, intimis aristato-elongatis,
non arachnoideis, extus glandulis sessilibus densis et setulis minimis
appressis obtectis, margine breviter ciliatis; receptaculum paleis
acuminatis laevibus involucro duplo triplo brevioribus obtectum. Corolla
rosea ca. 12-13 mm alt., laciniis limbi 4-5 mm lg. Pappus caducus
niveus, 6-8 mm alt., setis breviter barbellatis, basi in anulum concretis,
barbellis dense dispositis, anulo fundiculo paleaceo margine breviter
inflexo aperto est, apice cum nectario centrale cum pappo adjecto.
Achaenia ca. 3—3.5 mm lg. pallide flavida, mox brunnescentia, per costam
crassiusculo cartilagineam, transverse dense plicatam interdum interrupto

727

721

tuberculato foveola ornata, in lateribus 3-4 paralleliter et dense costulata,
costulis interrupto-foveolatis et glandulis sessilibus minimis disperse
obsita, apice margine dentato coronato, intra coronulam pappophoro
calyptriformi instructa.

Affinitas: J. cretaceae Bge. proxima est, sed phyllis involucri externis
et mediis appressis, corolla minore, achaeniis densius costatis et
validioribus ex toto ornatis, dentibus coronulae erectis, paleaceis, mox.
cartilagineis sat bene distat.

Typus: Dit. Volgograd, pagus Klekaja. In creta 16 VII 1887, leg. D.
Litvinov. Typus in Herb. Int. Bot. Ac. Sc. URSS (Leningrad) conservatur.

30. J. michelsonii [jin sp. n.

Caulibus ca. 50-60 cm alt., erectis, striatis, molliter arachnoideis,
glandulosis, striatis, a basi usque ad tertiam partem caulis longe ramosis;
foliis radicalibus et infimis longe petiolatis, petiolis planis, alatis, usque
ad basin gradatim dilatatis et semiamplexicaulibus, caulina media breviter
petiolata vel sessilia, basi auriculata et decurrentia, omnia pinnatisecta,
sectionibus linearibus, acuminatis, margine anguste recurvatis, remotis
opposite vel vix oblique opposite dispositis, cum rachide alata
confluentibus, supra rugulosa, viridia, glandulosa, subtus canescentia,
laxe arachnoideo-tomentosa; folia suprema angustissima linearia,
simplicia. Capitula majora ca. 3.5—4 cm in diam., involucro late cupulato
20-25 mm alt., phyllis extremis et mediis longe arcuatis, sensim
acuminatis, arachnoideis, partibus arcuatis atrato-violaceo-purpureis,
internis erectis acuminatis, apice longe aristatis, margine longe ciliatis
et valde involutis, extus hirsutiusculis et glandulosis. Corollae
purpurascentes, glandulosae, 15-16 mm lg., laciniis 5-6 mm lg. Pappus
breviter barbellatus. Achenia immatura.

Area geographica. Ucrainia occidentalis.

Affinitas: species nova ad J. mollissimam Klok. et J. calcaream
Klok. appropinquat, sed phyllis externis et mediis valde arcuatis recedit.

Typus: statio Maximovka—Bogdanovka, prope pag. Romanovka, 9
VI 1916, leg. A. Michelson. In Herb. Inst. Bot. Ac. Sc. (Leningrad)
conservatur.

31. J. demetrii Iljin sp. n.

Perennis, 15-60 cm alt., flavescenti-virescens, caulibus comparate
tenuibus, striatis, a basi usque ad partem mediam longe ramosis, abunde
glandulosis et laxe et tenuiter arachnoideis, felia radicalia et caulina
infima brevipetiolata, petiolis alatis, semiamplexicaulibus, caulina media
sessilia, basi auriculata et vix anguste et breviter decurrentia, pinnatisecta,
segmentis linearibus, integerrimis acutis, margine revolutis supra viridia
scabrella, rugulosa, subtus albotomentosa. Capitula mediocria, involucro

122

cupuliformi, 12—18(20) mm alt., basi leviter arachnoideo, phyllis externis
apice breviter deorsum recurvatis, apicibus atro-viridibus vel atro-
purpurascentibus et mucronatis; internis erectis, sensim acuminatis, dense
glandulosis. Corollae 13-18 mm alt., purpurascentes, laciniis 5-6 mm
lg.; pappus facile ex toto deciduus, albotomentosus, serruiato-barbellatus;
achenia 2.5—3 mm lg., vix scrobiculata, tota et glandulis sessilibus,
breviter pedicellatis structuram achenti occultantibus densissime obtecta.

Area geographica: Turcia ad finem USSR.

Affinitas: Haec species J. anatolicae Boiss. et speciebus affinibus
proxima est.

Typus: Distr. Kars, Olty, inter Bardu et Promezshutoczny, 26 VI
1911, leg. Demetrius Sosnovsky. In Herb. Inst. Bot. Ac. Sc. (Leningrad)
conservatur.

32. Pereplexia Ijin gen. n.

Planta scaposa; capitula sessilia + pauciflora; receptaculum
polygonale, polygonis leviter convexis, nudis, inter ea paleis acuminatis
laevibus brevissimis donatum; flosculi omnes hermaphroditi, corolla
usque ad basin plus minusve gradatim attenuata, appendices superiores
antherarum sursum attenuatae, inferiores setaceo-caudatae, flexiles, setae
pappi basi in annulum concretae, omnes facile deciduae externae supra
basin reflexae et perplexae, internae erectae; achaenia oblonga, obconica,
glabra, laevia, non coronulifera vel perfecte matura tuberculis minimis
disjunctis pappophorum cingentibus instructa.

A genere Jurinea Cass. genus nostrum structura achenii et pappi
‘distinctissimum est.

E genere hoc duae tantum species adhuc cognitae sunt.

33. P. fragida (Boiss.) Iljin comb. n.—Jurinea frigida Boiss. Diagn.
I, 6 (1845) 112; Boiss. Fl. or. II] (1875) 582.—Jurinella absinthifolia
Jaub. and Spach. Ill. pl. or. I] (1844-1846) 185.—Ic.: Jaub. and
Spach. tab. 185.—Iran septr.-occid, in alpibus (mons Demavend, mons
Totschal).

34. P. microcephala (Boiss) Iljin comb. n.—Jurinea microcephala
Boiss. FI. or. III (1875) 584; Iljin in Tr. Turk. Nauchn. Obshch. 2 (1925)
Die

735

INDEX ALPHABETICUS *

nominum specierum atque synonymorum plantarum
in tomo XXVII Florae URSS commemoratarum

Abolinia Tschern., sect. ................05 218
Acantholepis Less. .................ee eee 54
— orientalis LESS. ............:::cceseceeeeeeee 54
— — var. songorico (Trautv.) O.
and B. Fedtsch. ...............::0006 54
Acanthotoma Juz., sect. ................66 334
Acarna chinensis auct..........cccccee 89
— cyanoides BESS. ..........ccccesseeeees 546
— ovata WiIId. .............cccccccceesesseseeeees 89
Acarnaceae Link. .............c1cccceeceeeeeees 56
Actinia Bge., S€CE. ...........::eeeeeseeeeee 168
Alatae Sosn., subsect. ...............0:00006 672
Alboviodoxa Woron. ex Grossh. ... 75, 76
— elegans (Alb.) Woron. ex Grossh. ....
Cer is dene pure, py nent 75, 76
Alfredia talassica Korov. .................. 360
Alpinae Bge., Sect. ..............00+ 278, 334
Amberbopsis Tschern., sect. ............. 249
Amphilaena involucrata Lipsch. ...... 383
Amphoricarpus Vis. .................... 57, 74
eiclegans Albsveccie..cscssnces-1c ee 75
— kunetzowi Winkl. ex Lipsky.......... 75
— neumayeranus (Vis.) Vis.... 74, 75, 77
Amphorocarpus Post. and Kize. ......... 74
AnuraWuzscsect: 4.0.8... a eee 107
Anura (Juz.) Tschem. ...............0 107
— pallidivirens (Kult.) Tschern. ...... 107
Aplotaxis DEAE AAs. Le 361
— sect. Frolovia DC. ............c0:.:0085 384
SF rOlOVit DE Hs... occ SO 384
— gnaphalodes Royle. ...............0+++ 390
— involucrata Kar. and Kit. ............ 383
— sorocephala Schrenk. .................. 390
Appendiculatae Winkl., sect. ............ 259
VAN CULMIM GEE: ctccissassecsenesceessecseccceeestecstens 93
— aciculare O. Ktze. ...........:0000:00 284
— affine O. KtZe. .......ccccceceeteetees 153
— albicaule O. Ktze. ............0000000- 260
— alpinum O. KUZE. ..........cccesceeees 284
— x ambiguum Nym. ....................: 105
—— anomalum O. Kize. ...... noc eee eee 182
— antonowi O. KtzZe. ............000..00006 158
— arctioides O. KtZe. ..........cccceees 177

— aureum O. KUZEe. .........0:ccceeceeees 244
— bardana MLB. ...........yeccccceceeteeeeees 103
— bonvaloti O. Ktze. 0.0.0... 354
— bucharicum O. Ktze. .............5..... 357
— bungeanum O. Ktze. ............00 272
— buphthalmoides O. Ktze. ............. 287
SI CAPUSIIONKIZOR tt eee 166
— carduiforme O. Ktze.................... 196
— carduncelloideum O. Ktze. ......... 285
— chlorocephalum O. Ktze. ............ 188
— coronatum O. KtzZe. ............::0- 279
— cynaraoides O. Ktze. ...........0006 262
— dissectum O. KUZEe. ...........:::00c 160
— dolicholepis O. Ktze. ...............2: 197
— eriophorum O. Kize. ...........:0.0 361
— eryngioides O. Ktze. ............000- 186
— franchetianum O. Kize. ............... 221
— hohenackeri O. Ktze. .................. 195
— hystrix O. KtZe. oe 276
— integrifolium O. Ktze. ............0... 310
— intermedium Lge. .............:.2+++ Be: 99
aS =O; . Rtzeie see ah ot asl )
— karatavicum O. Ktze. ...............-- 246
— karelini O. Ktze. «0.0... eee 260
— kokanicum O. Ktze. .............:0-++ 328
— komarofii O. Ktze. .............0...06 250
— korolkovii O. Ktze. .........0:c0e0 248
— krauseanum O. Ktze. ............0.06+ 324
= MAD PANAUCL ypseeecn. Petes. ctst-eeeetee 98
eS tlappay Lie wtee...c8 Ro a ee 97
— — x A. tomentosum Beck. .......... 105
— — ssp. majus Aréne. ............000 97
— — ssp. majus x A. minus ssp.
eu-minus ATréne. ............-.00- 102
— — ssp. platylepis Aréne. ............... 95
ovat. Priby...L 3) eee eee: 104
— lappaceum O. KiZe. ...........:000005. 180
— x leiobardana Juz. and C. Serg. .. 105
— leiospermum Juz. and C. Serg. ..... 98
— — x A. tomentosum Mill. ........... 105
— lyratum O. KtZe. «0.00... ceee eee 264
— macrocephalum O. Ktze.............. 263
736 Arctium macropterum O. Ktze. ........ 277

"Reproduced from the Russian Original. Russian page numbers appear in the left-hand

margin of the text—General Editor.

724

— macrospermum Hayek. ............:. 99
—majus Bembe ren. ee ee 97°
-~— microcarpum O. Ktze. ..............-+ 317
— microcephalum O. Ktze. ............. 196
— minus Bermmh..............ccccceceeseeeees 100
— minus ssp. eu-minus Syme. ......... 100
— x A. tomtentosum Aréne.............. 106
— minus ssp. nemorosum Boswell. ... 99
— — var. nemorosum Th. Dur.......... 99
— — 3) macrocephalum Rupr. ....... 105
— minus Bernh. x A. nemorosum

| Wi) Ren py Re ee ce 102
— minutum O. KUtZe. .......cceeeereee 324
— x mixtum Nym. .............c.ccccees 106
— molle O. Ktze. ..0...ceccceeecssccceeese 185
— multilobum O. Ktze. ............000005- 350
— nemorosum Le}j. ...........:.ccccceeeeeee 99
— nemorosum Lej. x A. tomento-

sum’ Mill. erie ee 105
— x nothum (Riihm) Weiss. ............ 102
— obovatum O. KtZe. .........::cc0c00 277
— olgae O. Ktze. ....c....cceciecsseseseeees 166
— outichaschense O. Ktze. .............. 286
— palladinii (Marc.) Grossh. ........... 103
— pentacanthum O. Ktze................. 239
— platylepis (Boiss. and Bal.) Sosn. .. 95
— platylepis O. Ktze. ...........cccccccceess 318
Ses OSM rhs cas ees ke ee 106
— polyocanthum O. KtZe. ................ 260
— polycephalum O. Ktze. ...............: 206
= princeps O. Ktze.. ........204.2..c 30s: 300
— pulchellum O. KtZe. ......0...0cc00008 314
— pulchrum O. Ktze. ........00.cccccce 280
— purpureum O. KtZe. .......... eee 264
— pusillum O. KtZe. ...........cccccceeceees 273
— pygmaeum O. KitZe. ........:ccceee 167
— raddeanum O. KtZe. ........00:c0000 176
— radians O. Ktze. ................ Sosa 278
— radula Juz. and C. Serg. ............. 106
— schmalhausenii O. Ktze. ............. 246
— semenowii O. Ktze. ...............00006- 197
— sewertzowii O. KtZe. ............c000 313
— smirnowii O. KtZe. ...........00000000 266
— stenolepis O. KtzZe. .0.......ccccceee 278
— stshurowskianum O. Kize............ 298
— submuticum O. Ktze. ..........0.0.0.-- 325
— subracemousum Nym. ...........0...+- 102
— tenellum O. KtZe......cccceeceees 333
— tomentellum O. KtZe. ............00... 179
— tOMENCOSUM AUCK. ......0.-ccececceeeereees 103
— tomentosum Mil. ..............0000c00006- 104
— tomentosum var. glabrum Aréne. .. 103
— transcaucasicum SOs. .............+.. 103

— triflorum O. KtZe. «20... 212

— turcomanicum O. Kize. ............... 145
— umbrosum O. Ktze. .........0..c000008 178
— uncinatum O. Ktze. .........000cc00 178
— verticillare O. Ktze. ........0...:c0008+- 353
— wolgense O. KtZe. ........:cccscceseees 152
Athamus; NCC kigss.creeusiys ete st shade 77
Atractylis japonica Koidz. .................. 89
— lyrata var. ternata (Kom.)

Kiordz 2th. ee (Ve eae 89
— ovata Thumb. ...........cccccccceescceeeneees 89
Atractylodes DC. 0.0.0.0... eee 88
— lancea (Thunb.) DC. .........000000000... 88
— ovata (Thunb.) DC. .........0..000..0.. 89
— chinensis (Bge.) Koidz. ................. 88
— Aucklandia Fale. ..............c00000000. 361
Badghysia Tschern., sect. ................. 144
BanbeyavADly senses ee 75, 76
Bardana Tiillpes-cssse:.- ee ee 93
Bellae Ijin, sect. ...........ccccceeee 567, 720
Bennetia S. F. Gray. .........cccccccceeceeee 361
Brachyacanthae Winkl., sect............ 146
Calcitrapae Boiss., sect. .. 186, 277, 223
Cardopateae DC. .........:ccccccseseeseseeeees 57
Cardopatium JUSS. .........0:cc0c0c0e 58, 59
—_ eigen toldy Winkle 1, ans 58
— boryi Spach. 0.0... ccccceccseeseeees 59
Carduaceae BeSS€Y. ...........c0sccsceeerees 56
Fe CASSvececasseslecssagaessioceseteniee. ROE 89
CarduéesyBaill..:28. See ees. 56
Carduiformes Bge., Sect. ...........00... 187
Carduinae O. Hoffm., subtr. ............... 89
Carduinede CASS. ...........cccccccceceseseseeees 89
SST LIESS as, psckess2. ee 57, 89
Carduncellus Juz., subsect. ............... 284
Carduus astracanicus Spreng. ......... 152
— horridus B. Fedtsch. .................... 361
== Haxas DISChs msn cncsoe he eee 556
== mollis Mi: Bok. Sees eee. 667
— orientalis Adams. ...............0......+- 196
— cyanoides Pall. .........ccccccccceeeees 558
— — a. polyclonos L. ........cccccceeees 546
— — B. polyclonos Ly w....sceeeeeees 558
— inermis...J. G. Gmel................ 558
— polyclonos M. B. ........00cccceeeeeeeeee 559
— wolgensis M. B. .........:cccccceeeeeeees 152
GCarlina Ls gakccccete RAee 57, 77
— acanthifolia All... eee 85
— — B. Cynara DC... 85
— — acanthifolia auct. ..........00:c 84
— — B. Cynara auct. ......... eee 84
= acaulis..3.: ae 85, 86, 87

737 —— var. simplex Waldst. and Kit. .. 87

— — B. caulescens (Lam.)

Schimalleeetc cases sasveseseess 86, 87
pat ID SAIC USE ves esse cae cegeneccs vgesies stessee= 87
a ACE itesececk coc cncauansecesasece 80, 82, 87
— biebersteinit Bernh. ex Hornem. ... 82
— caulescens GIN. .........00:ccccceeereeeees 80
Sess 22) LEY TG Aaa 86
— cirsioides Klok. .............:ccccsseeeeeees 87
— intermedia Schur. ................::000+ 82
HI CUICLL AAU G ee c ccs snes suse es feacicweeastee 80
Se [7715 1H2).) La ett Ra a Aa on ta ai 83
— — B. monocephala DC. ................ 83
— — y. prolifera DC, ...... ieee 83
— longicaulis Somm. and Lev. ......... 82
— longifolia Rehb. ............ceeeees 82
— — B. pontica BOISS. ..............00 82
— nebrodensis auct. ..........ccccccccseeeceeee 82
— onopordifolia Bess. ex Szafer,
Kenlez VBE Pawl.. vos scsseeccsosseecee 84
— FACEMOSA AUC. .........cceceeeeceeeeeeteeeees 82
== ASEIMPICXHAUC Ute nce ncn ce ees ch eos scuassenses 87
— x szaferi A. Jas. and B. Pawl. ...... 85
=U ULC GESSANIC bathers cease eos <sesecnsn ie 82
=e AV ULC AMISHIE. Se eattcceesscccs sadeessessoes 78, 79
— — ssp. intermedia (Schur)
Fla. Cig neee ete ceenectar te keseneeren a 82
— — var. araneosa K.-Pol. and
| FARSYO) 0) cite ab ts ae a I PA REI 80
— — var. intermedia (Schur) Lindb. f.
ob sc aais aa RE AIL ty Ween CURA 82
— — var. longifolia Korsh. ............... 82
— — var. nebrodensis Kty]............... 82
— — B. sublanata DC. ..............:000+ 80
— — y. microcephala Ldb. ...................
G@arlineae Cassy. .......6.0....cces..00000 57, 89
ND) Cee RMN coe esos sv aestieastt 57, 90
Carlininae ©. Hoffm. subtr................. 57
Carlowitzia Moench. .................c000008 77
Carphephorus baicalensis DC. ........ 393
Carthamus cynaroides M. B. ........... 262
— pterocaulos C. A.M. ........::0-++ 276
Centaurea amplexicaulis S. G.
(Ciel Bess tet cee hea ate 559
— dubia S. G. Gmel. .............00:0cc0 62
— moschus Hab. ..........0.....cccc00cccees 705
Céntauriede LESS. ...........ccceseeceeeeeeeseees 90
Chaetocarpae Korsh.., stirps. ............ 677
Chaetocarpae (Korsh.) Iljin, sect. ..... 677
Chamaechinops Bge., sect. ................. 48
Chardinia Desf. .......0000000000.. ee 57, 69
— macrocarpa C. Koch.................0 71
— orientalis (L.) O. Ktze. ................ 70
— — var erivanica Tamamsch. ......... 70
— xeranthemoides Dee. .................... 70

125

Chodatella Alb., sect. ..............:0:::eeeee 5
Chromatolepis Dulac. ............0:000000- 77
Chrysantha Tscherm., sect. ...............- 167
CTY SISMIMZ SCC Ug beret reece een ee 244
Chrysoptera Tschern., sect. .............. 160
Cinarocephalae JUSS. ...........c0000c00c00e 56
Cirsium nidulans Rg). ..........:0:0c1000 361
Cnicus altaicus Steph. ex DC. ......... 384
Congestae Bge., SCCt. ......eeeeeeeeeeeeeees
ee: cemeeeeres LOO, MOM 2a Sere
Constrictae Winkl., sect. .................000000
EO. ene 259, 269, 276, 323
GOrdifoliaenO Gy. Re ek eect 643
Coronophora Juz., subsect. ............... 278
GGUSINIAC ASS. 2 ccssncsseecessceesass 108
SSE CLAN rues senor aetna deere deRuvas susati 187
— abbreviata Tscherm. ....................- 347
— abolinii Kult. .....00.. eee 218
— acanthophylla Kult. ...........00+ 338
— acicularis Franch. ..........:.0:c0e10 284
— acrodroma Tscherm. ................026 348
— actinia BOISS. ...........:cceeeeeeeeeeeees 168
— adenophora Juz. ..........0.:eee 337
— affinis Schrenk..................0.. 153, 154
— affinis var. talassica Kult. ........... 154
— agelocephala Tschern. ...............- 216
= HAL CAWNIZ Wee ces io cis easice sn ecusiges veces 254
== ALA AUC UM chit. estas Moun Spats eels 163
= Al ALAN SCNTEN Kage ee Sarees. ete. 163
— alberti Rg]. and Schmalh............. 239
— alberti ssp. horridula Bornm. ..... 240
— albertoregelia Wink. .................. 168
— albicaulis Boiss. and Buhse. ....... 260
— albiflora (Bornm. and Sint.)
JBXOV AAV S sccasageeasacec eee memset acne 147
— allolepis Tschern. and Vved........ 343
— alpestris Bornm. ................:-...005 282
— alpina Bge. .......... eee eeeceeeeeeeeeee 284
— ambigens JuZ. ............esceeeeees 209
— AMOENA AUCH. ooo... eeeeeeeeeteeeeeteeees 227
— amoena WinKl. ...........e eee 228
— amplissima auct. ...........0cceew 179
— androssovil JuZ. ............:0sceeeee Dili
VAMP REMMI 5, cn. cect ences eas ssnuaesstosuanl 199
— angusticeps JuZ. «0.0.0.0... ATS)
— annua Wink. ...........::cceeecceeeeeees 358
— anomala Franch. ................:0:00 182
— antonowii Winkl. ................-.0 158
— apiculata Tscherm.................0.05 174
= APLETA ANC. .... cc... vececsnsnsnrcenaeteness 166
— arachnoidea Fisch. and Mey. ...... 316
— arachnoidea var. spinosior Rgl. ... 316
— — var. typica Rgl....... ee 316

== ATAIENSIS BRE: se. sccigec cose sccesseseessss 324

726

— araxena Takht. .................... ees et 192 — chejrobadensis Kult. .................... 150
— arctioides Schrenk. .........0.......0.. 177 — chionophila Rech. f. and Koie. ... 222
— armena Takht. ..............eeseeeeeeeee 189 — chlorantha Kull. «0.0.0... eee 247
— astracanica (Spreng.) — chlorocephala C. A. M................ 188
Mamamsch)ie--2,.22-22005----- 152, 154 — chrysantha Kult. 00.00.00... eee 340
738 Cousinia atripurpurea Juz. ........ 280, 318 — cisdarvasica V. Nikit.................. 214
— aurea Winkl. 000.0... eeeeeeeeeeeeee 244 — coerulea Kult. 20.0.0... ceeeeeeeereees 235
— aurea B. refracta Bornm. ............ 245 — commixta Kult. ............c:ceeeeeee 286
— auriculata auct. .......:cccccceeseeeeeeeee 222 — congesta Bge. ..........eceeeeeeeeteees 216
— badghysi Kull. .0....... eee 144 — COFONALA AUC. ..........ceseccceeereeeetees 278
— baranovil Juz. oe. eee eeeeeeeeeeee 255 — coronata Franch. ...........::ccceeeee 279
=D ALQININAUC geen. «. ccscasecsstsecessessees 294 — corymbosa Winkl. ..................0 235
— batalinii Wink. 00.0.0... eee 295 — corymbosa f. ramosissima Winkl......
— batalinii a. latisecta Bornm. ........ Od) Sea Wie PER eee tae ee eon oy 297
— — f. tenuisecta Bornm. .............. 295 — cryptadena Juz. ............ceeereeees 157
— — B. virescens Bornm. ............... 296 — CUYVIlObA SUZ. «00.0... cere cere 338
—— DECKEM pl TAUty..<5.2..ceseeczcsceesccnncocs 265 — cynaroides (M. B.) C. A. M........ 262
— bipinnata auct. .........eceeeee 163, 165 — daralaghezica Takht..................... 189
— bobrovii Juz. ....0. eee eee eee 169 — darwasica Winkl. ............:::ee 309
— boissieri Buhse. .......................000 152 — decurrens auct. ...........000000+ 205, 210
— bonvalotii Franch. ........................ 354 — — var. congesta auct. ......... 206, 210
— bonvalotii var. poliortrichodes — — var. scabriseta Winkl......212, 213
BONIS es peetes ees ceecee 352 — — f. oligocephala auct. ...... 206, 207
— botschantzevil Juz.................0 267 — — y. oocephala Bornm. .............. 208
— brachyptera DC. ........ eee 192 — — y. turkestanica Rgl.................. 203
— bucharica Wink. .................000 357 — — f. tschimganica O. Fedtsch..... 198
— Dungeana auct. ...........cccceecceeneeees 274 — decurrentifolia Juz. ............0.:00 257
— — Rgl. and Schmalh. .................. 272 — dichacantha Lipsky. ..............-..-+. 358
— buphthalmoides auct. .................+ 286 — dichotoma auct. ........ ee atc 272
— buphthalmoides Rgl..................... 287 — dichotoma Be. ............ eee: 269
— butkovii Tschern. and Vved........ 304 — dichromata Kull. .......0......e eee 214
— caesia Winkl. ...........eeeeeeeeereeees 264 — dilatata JUZ. .......cecccesceeeeteeeee 183
— caespitosa Winkl. ..........0...:ceee 356 — dimoana Kull. ..........ccccceeeeeeeeeeeee 309
— calcitrapaeformis Jaub. and — dissecta Kar. and Kir................... 160
Spach? ties... oa ccsre teense 324 — dissecta var. genuina Trautv....... 160

= CAV aE pe eo oes veesen suo senesas 295 — — var. macrocephala Kgl.
— campylacantha JuZ. ............c0000+- 279 and Schmalh. ...............::c0ee 161
— campyloraphis Tscherm. ............... 228 — — var. microcephala Trautv. ...... 165
== CATA UI Ziaceece sete ee cewesanstees anos senate 313 — — var. perovskiensis Bornm. .... 161
— candicans JuzZ. ...........ccccceseeeeeeees 230 — — var. simplicior Trautv............. 165
— CANESCENS AUCC. ........cceeeeeeereeeeteees 221 — dissectifolia Kult. 0.00.00... 257
— capusii Franch. .............:::ceeeceee 166 — divaricata Wink. .................: 310
— carduiformis Cass. ..........c00cccce 196 — dolichoclada Juz. ...,.........:.::ee 162
— carduncelloidea auct. ................. 286 — dolicholepis auct. ..............+++ 199, 205
— carduncelloidea Rgl. and Schmalh. — dolicholepis Schrenk. .................. 197
Cieguborspenes td erties, cotagunsesseteaht 185 — dolichophylla Kult. .................2.219
— carduncelloidea B. sonchifolia — dshisakensis Kult. ................ 299, 300
BROMINE 2 Rt oieecces cc nees reer 287 == AUIS SUZ Ac cgecncen se vek cease cesar ese 179
— carlinoides Aauct. ........ccccceccceee 277 — dubia M. Pop. ..........e::ceeceeeeeeeees 327
— catenata Rech. ff. «..........--..c-----00-+ 170 — echinopifolia Bornm. ................... 360
— centauroides Auct. ........:cccceecee 316 a CPONS NUL bi sae cuca tome n teen wee 215
— centauroides Fisch. and Mey. ..... 312 (Fu eye Wi OV Asean ecceecs wea se oncosechenee 218
— ceratophora Kult. ...0... ee 355 — erectispina Tscherm. ...................+. 269

— chaetocephala Kult. ..................... 151 — eriophora Rg}. and Schmalh. ...... 361

727

= EG] Ot Ch a IUZ 4 -26052.004- eos seSbectecnent- 352 — hystrix C. A.M. ..eccccccecceceeeseeeeeees 276
— erivanensis Bornm. ................00: 193 —ailjiniiwakhtject eee al tel sete 194
— eryngioides Boiss. ..................00 186 — integrifolia Franch. ................2-. 310
= eupenii) Kultac...2....-ssccs canoes oatee es 275 —— integrifolia B. heterophylla
= fallaxs WANK aise csoucas tess osewoa 255 BOM. cece as tees 310
— failax var. armata Wink............. 255 — intermedia C. A. M. .......:::0eee 188
— fascicularis Juz. ............cc cesses 184 — interrupta WinKl. ............:ccce 278
— fedorovii Takht. ..........0...... ee 190 -—AIntersitay UZ eee cca POE ee 242
— fedtschenkoana Bomm. ............... 224 — iskanderi Bornm. ................. 287, 326
739 — ferganensis Bornm....................... 320 — jacea Rech. f. and Edelberg. ...... 342
— ferruginea Kult. 00. eee 308 — JASSYENSIS AUCH. «00... ee eeeeeteeereeee 327
— fetissowii Winkl. ..................2.04. 293 — jassyensis Winkl. ............0....c0 332
= MAMTA TZ etic scsvancese sucess 207 — juzepezukii Tscherm. .................... 203
— flava Bornm. ..............::ceseceeeseeeeeee 355 — karatavica Rgl. and Schmalh. ..... 246
— franchetii auct. .........:0ceseeereeereees 220 — karelini LESS. ............:ccceceeeeeeeeeees 260
— franchetii Winkl. .........000.0.. 221 — kazachorum Juz. ...........::ecesee 212
— franchetii B. drepanophora — knorringiae Bornm. ..................... 289
BONN 55.5. cs cere ave 220 — kokanica Rgl. and Schmalh. ....... 328
— Freynii Bornm. and Sint.............. 265 — kokanica var. virgata Wink. ...... 328
= SOTIUIR OCIS rete ve-cecnoskse ae tie esvestcnene 184 — — a. genuina Bornm. ................. 328
— glabriseta Kult... eee eee 292 — — y. perapressa: Bornm. ............. 332
— glandulosa Kult. «0.0000. 158 — — f. nowobadensis Bornm. ........ 325
— glaphyrocephala Juz. ................... 334 — komarowii auct. ..........0cccceeeeeee 256
— glaucifolia Kult. ............::::1:ceee 286 — komarowii (O. Ktze.) Winkl. ...... 250
— glochidiata Kult. .....000.. eee 175 — komarowii var. patula Bornm. .... 252
_ —gnezdilloi Tscherm. ...................... 233 — korolkovii Rg]. and Schmalh. ..... 248
— gomolitZkii Juz. .......... eee 238 — korshinskyi Winkl. ..................+: 225
— gontscharowil Juz. ...............:::00 170 — krauseana auct. ..........:.ccsccceeeeees 325
— grandifolia Kult. ......0.... ee 249 — krauseana Rgl. and Schmalh. ............
— grigoriewii Juz. ............:eeeee eee 290). Rar ye eees eee ee alate 324, 325
e—POTISCA KOU ones cptenae Pe cave eoncaneteodsne 252 — krauseana var. macrocephala
— gulczensis Kult. .............ceeeeeeeee 297 Wine 2a sasenee crane 326
— gundelia C. Koch. ............::e1e 192 — genuina f. elata Bormm. .............. 325
— haesitabunda Juz.................:200+ 247 — — — f. humilior Bornm. ............ 325
— hamadae Juz. ...........ceeeeeeeeeeeeees 155 — — B. submutica Bornm. .............. 325
— hastifolia auct. ...............cseeeeeee: 325 — —y. longispinella Bornm. .......... 326
— hastifolia Winkl. «0.0... 328 — kuekenthalii Bornm. .................... 256
— heliantha Bge. ..............::ccccseeeeee: 268 = tKULTASOWLE MUZo le. ..s sateoe he aSerecetat 307
— heterogenetos Bornm. ......... 287, 315 — kuschkensis Kull. ................000006 163
— heterogenetos a. pulchelliceps — lachnopoda Ju. ..........1..:10000000+ 342
BB OENIN sos caastoe ote de oa coset este cesses 287 — laetevirens aUct. ............ cess 353
— heterogenetos B. majoriceps — laetevirens Wink. ...................06++ 352
ROM iss oooh eiscsacet ch oces oes 287 — laetevirens var. flava Wink. ........ 355 °
— heteromorpha Bornm................... 282 — — var. monocephala Winkl........ 352
ppp AMICUS OUD ods snr ncn ooResauehe oass 288 seplanata Winkle ce. cct2.c cece. cccset: pees: 335
— hissarica Wink. ................00:00000 353 — lancifolia Wink. .............01010+ 1325
— hohenackeri Fisch. and Mey. ...... 195 == lAMiGEPS SUZ soe. oo oc steeaeetsceseepesads 302
— hoplophylla Tscherm. ................... 234 — lappacea Schrenk. ...........0.0.0 180.
= SHOMESCENSIUZ: cee eccce- bathe sanccsec 240 — lasiandra Bge. ..:...........-...- et ey 259
— horridula Juz. ...........cceeeeeeeeneees 212 — lasiosiphon Juz. ...........e eee 259
— hypopolia auct. .........2:cecceeeeeeees 150 — leiocephala (Rgl.) Juz. ................. 314
— hypopolia Bornm. and Sint. ........ 148 —- lepida AUct. .........cccecccsceeseeeeeereeees 147
— hypopolia B. albiflora Bornm. and — leptacantha (Bornm.) Juz. ........... 184

SUIS cicesc A ca enc eet wee eee 147 — leptacma Tscherm. .............::0-+2+ 227

728

— leptocampyla Bornm. .................. 283 — orientalis (Adams) C. Koch. ....... 196
— leptocephala Fisch. and Mey. ..... 146 — orientalis Grossh. .............::000000 196
— leptoclada Kult. «0.0... eee 234 — orthacantha Tscherm. ................... 273
— leptocladoides Tscherm. ............... 232 — ortholepis JuZ. .....0.....eccceeeceteeees 319
— leucantha Bornm. and Sint.......... 252 — outichaschensis Franch. ............... 285
— leucocitrina Kull. .............0eccee 157 — ovezinnikovii Tscherm. ................ 296
— linczewskii Juz. .0...0.. eee 145 — oxiana Tschem. ............:eeeeeseees 163
— lipskyi Wink. ...........eececeeceeeeeee 337 — oxytoma Rech. f. ............:c:eeee 296
— — var. approximata Wink. ........ 340 — pallidivirens Kult. .......0..0.::1000 107
— litvinovii Kult. oe 227 — palmatiloba Jaub. and Spach. ..... 350
— lomakinii Wink. ..............0.....0. 194 — pamirica Bornm. ..............0cc000 222
= lyrata, Bocaire cn. meee 263 —pannosa Winkl. ...........0..cee 220
— lyratifolia Wink. ...........:cccceeee 284 — pannosa var. microcephala
740 Cousinia macilenta Winkl. ............... 226 Wink seers e neers 220
— macrocephala C. A. M. ............... 263 — — var. subviridis Winkl.............. 221
— macrocephala 8. spinulosa — pannosiformis Tscherm. ............... 222
Dorrialkevets eect Ey 263 — patentispina Tscherm. ................... 270
— macroptera C. A. M. .......... eee 277 — pauciramosa Kult. .............0.... 315
— macroptera var. obovata Winkl.... 277 — peduncularis Juz. ......0...... 297
— magnifica Juz... eee eeeeeeeeee 334 — pentacantha Rgl. and Schmalh. .... 239
— maracandica Juz. ......... eee ANS) — pentacanthoides Juz. .................... 241
— margaritae Kull. ...000 ee 330 — perovskiensis (Bornm.) Juz. ........ 161
— margiana JUZ. ...........eeeeeeeeeees 187 — platylepis Schrenk. .................0... 318
=MMedians UZ asec tcc ee eee 245 — platystegia Tscherm. .................... 214
— meghrica Takht. ............:cecceeeeeee 193 — podophylla Tscherm. .................... 232
— microcarpa BOISS. ............::cccee 317 — polycephala auct. ..........ccccccceee 198
— microcarpa var. parviceps — polycephala Rupt................-.0024- 206
Omnia sted See eect 318 — poliothrix Wink. ................. 352, 353
— microcephala C. A. M................. 196 — — var. robusta auct. ..........e 353
— mindshelkensis B. Fedtsch. ......... 237 — — f. albiflora Bornm. ................. 353
— minkwitziae Bornm. .................... 200 — polytimetica Tscherm. .................. 270
— minuta BOSS. ..........c:cceeeeeeeeeeeeees 324 — popovii Kult. .........:ceeceeeeeeeeeeeeeee 320
— mogoltavica Tschern. and Vved. ...... — praestans Tschern. and Vved. ..... 305
ond eee ELEM ARSC T 303 — princeps Franch. ...................-+.---- 300
— mollis Schrenk. .............eeeeeeeee 185 — proxima JUZ. ........ ec eeeeeeeeeeeeeeees 229
— mollis var. latifolia Wink. .......... 185 — psammophila Kult. .........0............ 164
.— — B. squarrosa Bormm. .............. 185 — pseudarctium Bornm. .................. 170
— mucida Kull. .0..... eee eeeeeeeeeeees 150 — pseudarctium f. leiocephala
— mulgediifolia Bornm. .................. 282 “BOMNM soo cccrscctoces tee eee 179
— multiloba DC. .......... eee 350 — pseudoaffinis Kult. ........0... 155
— murgabica Tscherm. ..................4.- 274 — pseudaffinis var. bucharica Kult. ......
= NeglectarIUzases san een eee DAS) : ERM PIE ian enh AA ia an reliance li 155
— newesskiana auct. .........:cccccccceees 346 — pseudo-bonvalotii Juz. ................. 353
— newesskiana Winkl. ....................- 344 — pseudodshisakensis Tschern. and
== nine JUZ seis, el NS See 318 Vivedyrntrasisarad nip nt eran eranettees 299
— nobilis Rech. f. ........cececeeeeeeee 176 — pseudolanata M. Pop. .................. 336
— obovata BOISS. ...........cccccccccccceeeees 277 — pseudomollis Winkl............. 183, 184
— olgae Rgl. and Schmalh. ............. 166 — pseudomollis var. platyacantha Bom.
— olgae var. cordata Wink. ........... 166 ITS RENE SRN ead SANUS Ste we Seen Tener 183
— omphalodes Tscherm. ................... 326 — B. leptacantha Bornm. ................ 184
— onopordioides Ldb. ..................... 260 — pseudostenolepis Rech. f. and
— OOPOdA JUZ. oes eeeeeteneneceee 170 Edelbot en crete teectre 280
— oreodoxa Bornm. and Sint. ......... 174 — pskemensis JUZ. ...........10cccccccee 198

— oreoxerophila Kult... 150 — pterolepida Kult. .........0.....0.....8. 225

729

— pugionifera Spach. ...............0c00 277 — smirnowii var. armata Wink. ..... 266
——puilchellay ge: 2ctss0¥ sas hicensbacckyexcsens 314 — — var. globosa Kull. 0... 266
— pulchella var. latifolia Winkl. ..... 314 — sogdiana Bornm. .............cc000c0000 161
— pulchra Wink. .............ccceeeeees 280 — sonchifolia Bornm. ...... WS ccasiesenceuats 287
— pungens JUuZ. 00... eeeeeeeeeeeeenees 205 = SOTOMA MUZE ccsccces thet Seradtees 346
— purpurea C. A. M. ........eeeeeeeeeee 264 — speciosa WinkKl. ............cccceeeee 294
— pusilla Wink]. «0.0.0... eeeees 273 — spiridonovii Juz. ...........0:cccee 208
— PYEMAEA AUCL. ........eeccceescceseeereees 274 — splendida Winkl. ..:........:csc.00.::. 337
— pygmaea Winkl. .......... cece 167 — sporadocephala Juz. ...............00... 207
— raddeana Winkl. ................::0008 176 — spryginii Kull. ..............0.ccccseene 268
— radians Be. ............:cceceeseseeeeeeees 278 ae SDUNIGUZs 2. eee eee 179
— radians §. paradoxa Bornm. ...... 279 =I SQUATTOSA AUC te aes crs ees 192
— — y. alpestris Bornm. ................. 282 — stahliana Bornm. and Gauba. ...... 150
— ramulosa Rech. f,. ...........:ccceee 348 — stellaris Born. .............::cccc00e 287
— rava Wink... eee ceeeeeeneees 290 — stenolepis Rgl. and Schmalh. ...... 278
=== TehractanS UZ ik. hes, WL dee a, 245 — stenophylla Kult. ...............cc 251
= STE GEL WWATIKIE betes cserettceevent acess 316 —— Stenop tera VUZ. 02. cccs. 209. ere eee 165
<A TESINOSA UZ .2 keseiet Moc ccteceraseasdenss 210 — stephanophora Winkl................... 340
= TESUPINAtA UZ. 8.228 ekeessoaeente 220 — stricta Tschem. ..:..........:ccc:cc0ecceees 172
— rhodantha Kull. .......... eee 307 — strobilocephala Tschern. and
741 —rigida Kult. .........cc.ccecceseceesersees 236 Vivediins 5 bgks SE eho sacl 305
=—TOSeat Kult ee Aas ca ocdens 306 — subappendiculata Kult. ................ 320
— rotundifolia auct. .........0cccceccees 287 — subcandicans Tscherm. ................. 230
— rotundifolia Winkl. ...................0. 307 — subinermiceps JUZ. ......c.:cccccc0ccce 161
— rubiginosa Kull. ............ccceseseees 349 — subinermis Tscherm. ..........:.....000+ 235
— rubrarenosa Kull. ..............cccee 307 — submutica Franch. ...........:cc0 325
—— WF USSCOLAUIUZS eee at Se eee sepclicke 266 =—FSUD ULES WZ se netes: eee 280, 284
— sarawschanica Winkl. .................. 355 — sylvicola Bge. ........ccccceceessvesseenes 272
— sarawschanica §. elegantula — syrdariensis Kult. ................:0000 202
SONI Bee. costed aoe cesses 355 — talassica (Kult.) Juz. .......0.000000.... 154
— Seabrida JuZ. .........::c.sscccecceceeseeeeee 199 — taldykensis JZ. ........c.sccccccesseceeess 328
— schischkinii Juz. .........ccccceeeeees 294 — tamarae JUZ. ..........ceccetcceseeeseeeseeees 326
— schistoptera Juz. .............ccceccccee 165 — taschkentica Juz. ........ccecceeeeees 178
— schmalhausenii Winkl. ................ 246 — tedshenica Tscherm. ...............005- 274
— stshurowskiana auct. ........... 199, 300 — tenella Fisch. and Mey. ............... 333
— schtschurowskiana Rgl. and — tenuifolia auct. .........ccccceecceeseeees 189
Schmalhis sc. 8ae ene ee 298 — tenuisecta JUZ. ....... cc eeeeseeneeeees 202
— schtschurowskiana a. laniceps — tianschanica Kult. ...........0.....00 303
ES OTIS ec cc cl 302 — tomentella Wink: ...................0. 179
— — f. polylepis Bornm. ................ 306 — trachylepis var. integrifolia
— — y. subadprssa Bornm. ............ 300 OOR¢Z GPa ee Bea ected 357
— schugnanica Juz... 346 — trachyphylla Juz... 339
— scleracantha Kull. ........00.0....c. 205 — transcaspicad JUZ. ........00.ccccceeeees 152
— sclerophylla Juz. ..........0..ccccee 173 — transiliensis JUuzZ. ...........:cccccceeee 329
— semenovii Rgl...............cccceeseeeees 197 — transoxana Tscherm. ..............::0000 208
— semidecurrens Wink. .................. 222) — triacantha Kull. ........... ee al79,
— semilacera JUZ...............:..ess0000000: 342 triceps Kult. 202), nec. ee 292
— SEVETIZOVII Rg... ceeeseceseeneees 313 — trichophora Kult. ............ccccee 345
— severtzowii var. leiocephala — triflora Schrenk. ...............0ccece 242
RG Bee ead to teen, 314 — PYUNCALA SUZ. oo. .eecescceccceseetseeteeeees 308
— simplex Kull. ........cccccccccccscscereereees 252 — turcomanica Wink. .................... 145,
— SIMpLiCiO“ SUZ. .....c..00cce000cc0eeeeneee 165 — turkestanica (Rgl.) Juz. ................ 203
— simulatrix Wink. ..................000 322 — turkmenorum Bornm. .................. 165

— smimowii Trautv. ............cc00.cc0ee. 266 — ulotoma Bornm. .......................000- 258

742

730

— umbilicata Juz. ..........c.c0:ccccceeeeeees 204
— umbrosa Be. .............ceecesceeteeeeee 178
— umbrosa f. virescens Bormm. ..... 178
— uncinata Rl. «0.00... ceeeeeseeeeeeeee 178
— vavilovii Kult. ............... eae oe a 249
— verticillaris Bge. .............: ese 353
— verticillaris var. leiocophala
BOmMmeancscn te Se 353
— — Val. MAJOFr AUC. .........c0 cece 353
— vicaria Kull. ...... ce eeeeeeeeeneeees 198
— victoris Tschern. ...........:..:ceeeeee 156
— vvedenskyi Tscherm. .................... 323
— waldheimiana Bornm. .................. 330-
— wolgensis C. A. M.........:ccccesee 152
— — var. offinis Rgl.............. Serta 153
— xanthina Bornm. ..................eee 338
— xanthina . subvestita auct.......... 340
— — B. Bormm. ...... eee 340
— — y. approximata Bornm. .......... 340
— xanthiocephala Tscherm. .............. 156
— xeranthemoides Bege. ................... 317
— xiphiolepis Boiss. ................ 276, 357
Cousiniopsis Nevski. .................::0006 57
— atractyloides (Winkl.) Nevski........ 58
Ctenarctium Juz., sect. ...........cceee 182
Cyathidium Lind. .................0000000000 361
Cyanoides Korsh., stirps. .................. 541
Cyanoides (Korsh.) Iljin, sect. ... 541, 715
Cynaraceae Lind). ..............cccccccceeees 56
CYNAREAE Less., trib. ............ sehen 56
Cynareae subtr. Echinopsideae Less. .. |
‘— — Echinopsidinae O. Hoffm. ......... 1
Cynaroideae Benth. ..............cc00cc0000 56
Cynaroideae Bge., sect. ................65 259
Cynaroideae subtr. Echinopsideae
Benthi. ney... ee ]
Derderia Jaub. and Spach. ............... 643
Derderia (Jaub. and Spach) Boiss.,
SOCIAL MISOIAE Lt As 643
Derderia cheirifolia Jaub. and Spach. ...
Te each SN 644
Dichacantha Winkl. subgen. ............ 357
— — Lipsky, sect. .......0. eee 357
Dichotomae Bge. sect. ....................-- 269
Dichotomae Juz., series. .................. 269
== Be sect. hice Se 323
Distemma Spach, subgen. .................. 59
Drepanophorae Winkl., sect. .................
TE AGS ae 152, 160, 166
Echinanthus NeCK. ..........1.cc0cc0ccceseereeees 2
Echinops' 2). 22222 eae 2
= SECH IB GE ROT eee 40

— abstersibilis Ijin. ........0..00... 46
— acantholepis Jaub. and Sp. ........... 54
— — var. songorica Trautv............... 54
— albicaulis Kar. and Kir. ................. 18
— araxinus Mulk. ..0.....:.cccccecccestseeseeeeee
— argyrocomus Bornm. ................00+- 1]
— armenus Grossh. ..........c0ccesccceeeeee 14
— babatagensis Tscherm. .................... 24
— bannaticus ssp. quercifolius Freyn.
TOLER TR LR es 43
— Dipinnatus auct. ..........cccccccesseeeeenes 26
— brevipenicillatus Tschem. .............. 12
——"Caulerw* Gmelin totes cesoces 30
— — subunifloro. .. Gmel. ............... 45
— chantavicus Trautv. ..................00.5 22
— cirstifolius C. Koch. ...............c0006 40
— colchicus Sos. ............c::ccsceeseeees 43
— commutatus Juratzka. .................... 47
— connatus C. Koch. ............ eee 27
— conrathi Freyn. ............c:ccecceeseeees 34
— daghestanicus Iljin. ....................... 41
— dahuricus Fisch. ..:..........::ccccceeee 44
— — a. latilobus DC. ...................4. 45
— dasyanthus Rg}. and Schmalh. ...... 21
— Aavaricus THeVIN. ..........0cccccccereees 50
— dissectus Kitag. ..........ceeeeseeseee 45
— dubjanskyi Iljin. .......00.... 19
— erevanensis Mulk. .................00000 4]
— exaltatus AUCL. ........ccccceceeescceeeeees 46
— exaltatus Schrad. ...............eeeeee 47
— fedtschenkoi Ijin. ...........0... ee 25
— foliis integris Gmel. ............0...0.00-- 52
— foliosus Somm. and Lev. .............. 35
— galaticus Freyn. ...............ccceeeees 43
— gmelini AUC. ..........ceeccceeseeseeeseeees 45
— gmelini Turcz. ............eeseeseeesseeeee 52
— gordgaginii Keller. .................:0 51
— griffitianus auct. ......... cece 14
=erossheimit Ijiny sic ee 14
— horridus auct. ........ccccccccececesseeeseeees 25
— humilis M. B......... cece eeeeeeeeeneeeeee 50
— — var. elatus Kryl.........00. eee 50
— hypoleucus Ruff. ..............c0c00eee 22
== hyintiMulkewieircn- ee reer 37
— integrifolius Kar. and Kir. ............. 48 |
— JAXATICUS BE. .........ccccccccceteteeeeees 18
— jesdianus Boiss. and Buhse. .......... 28
— kafirniganus Bobr. ................. 18, 714
— karabachensis Mulk. ....................-- 38
— karatavicus Rgl. and Schmalh. ...... 23
— kasakorum Pav. .........::cccccceseeeee 18
— knorringianus Iljin. ...........0.... 53
— latifolius Tausch. ..........0... eee 44

— leiopolyceras Bornm. ....................- 14

743

— leucographus Bee. .................000 14
= lipskyi Ijin: -..:..5:....c. nie 16
— macrophyllus Boiss. and Hausskn. ...
ly SORTS ee heat beter: 28

— macrophyllus var. glabricaulis
Bomimen 200 ae ees 27, 30
— maracandicus Be. ...........:ceee 1]
— maximus SieVETS. ...........ccceeeeeeees 40
— multicaulis Nevski. ................0:0 7)
— nanus Be. ...0.0.....eeeeeeeseeeenreeeeeeees $2
— obliquilobus Iljin. 0.0.0.0... 23
— olivieri Jaub. and Sp. .................... 54
— opacifolius Iljin ex Grossh. ........... 26
— orlentalis Trautyv. ........... cece 25
— ossicus C. Koch. .........ce eee 44
— persicus Stev. and Fisch................ 33
— polyacanthus Iljin. ......0.0.. 34
— polygraphus Tscherm. ..................-- 1]
— praetermissus NevsKi. .................... 15
— pubisquameus Iljin. .............0.000.. 20
— pungens Trautv. ......0. cece 33
— raddeanus Somm. and Lev. .......... 33
== TO! Le. ets... Ai ees Peed rime. 30
— — var. latilobus Krryl. ................... 31
— — a. vulgaris DC. ....... ee 31
— — B. tenuifolius DC. ................. 30
— — y. glandulosus Korsh. ............... 82)
— — y. meyeri DC... eee 32
— — 6. litwinowii Korsh................... 31
— ritrodes Bge. ........:cccsccesccceseeseeee 16
— ruthenicus auct. .......:ccccceccceceeereeees 30
— ruthenicus M. B. ...........eeeeeceereeeee 31
— saissanicus (Keller) Bobr. ............. 51
— sevanensis Mulk. ...2........::e:eeceeeeee 38
— sphaerocephalus L. ...............000 40
— spiniger Ijin. 2.0.0... ceeeeeeeeeeee 32
— Strictus Fisch. .............:0ccccsssseeseeees 47
— subglaber Schrenk. ..................0006 20
— szovitsii Fisch. and Mey. .............. 36
— tauricus WiIId. .........:.ccccceeeeneeeees 30
_ — tenuifolius Fisch. ............0cccccccceeeees 30
— tjanschanicus Bobr. ............... 22, 717
— tournefortii Ldb. .............cceee 39
— transcaspicus Bornm. ..................... 10
— transcaucasicus Iljin. ...................... 36
— tricholepis Schrenk. .............:00.00- 46
— tschimganicus B. Fedtsch. ............. 50
— turczaninovii Ldb. ......... ke aetna bin 52
— viridifolius Hjin. ............ eee 42
— viscosus DC. ........:ccccccsecssssssseeseeess 27
— xanthocanthus Rg). and Schmalh. ....
PUES a hae NE: Sb cinrelevmerinnna ae: 11
Echinopodeae Cass. .........::ccsccesneeseeeees 1

Echinopsede Cass. ........:.ccccscccescessecssees 1

731

ECHINOPSIDEAE Cass., trib. ............. 1
Echinopus ‘Adans. .........00000000e00e Bot 2
Eglandulosa Aréne, Sect. .................+- 95
Eriocousinia Tschern., sect. .............. 220
Eriostemon LESS. ........0ccccccccccesceeeeseees 361
Euxeranthemum DC., sect. ................- 62
Floccosae (Sosn.) Iljin, sect. ............: 600
Frolovia (DC.) Lipsch. ............. 361, 384
— asbukinii (Ijin) Lipsch. ............... 387
— formosana (Hayata) Lipsch. ..:..... 386
— ledebouriana Lipsch. ................-.. 384
Ea) 72 417010) Wt |v Nate Mr a ae 384
— sulcata (Iljin) Lipsch. .................. 388
Glaphyrocephalae Tschem., sect. ..... 333
Homatae Boiss. sect. .................0006+- 152
Haplotoxis Endlich. ...............:000 361
— SeCt. FrOlOvia. .......0...ccccccccseeceeees 384
== frolovil db. seni. Re 384
— involucrata (Kar. and Kir.) Ldb. .. 383
— sorocephala Schrenk. ................-- 390
lHarrisonia Necks cc....020:-ccecetee eee 59
Hellanthae Bge., sect. «0.0.0.0... 268
Heteracanthae auct., sect. ........ 168, 220
Wink ssecte.... Sant Seana 152
Heterotrichum M. B. .1...........0.....00-- 361
— arcticum M. B. ..2.......c0000ccccceeeeeees 484
— atriplicifolium M. B.................-.-- 472
— dissectum M. B. .............00000ceeceees 529
— glabellum M. B................++ 472, 474
— leptophyllum M. B.............:. 410, 412
— pulchellum Fisch. .............00.00000+ 523
= salsum! MB aeet) a2 eee Se 504
— schanginii M. B. ..........:0ccccceeees 397
— squalidum M. B. ........1..c.0000000000 526
— Stelleri M. B......ccccccccccccceeseseeeeeeee 444
Homalochaetae Winkl., sect. ....225, 278
Homalochaete Juz. sect. .........0......... 226
Hoplophylla Tschern., sect. .............. 175
Hypacanthium Juz. ..................08 358
— echinopifolium (Bornm.) Juz. ..... 360
Imbricatae Boiss., sect. ................000-+ ae
Inermes Winkl., Sect. ..............:cccc0cceeleeee
Jalen ule Ae 167, 183,. 250, 333
Insculptae Iljin, sect. ................. 624, 723
Integrae iljin, sect. ........5.....0 $83, 721
Irano-armeniacae Takht., subsect. .... 188
Irano-turkestanicae Takht., subsect. .......
Bet EEE Crees 195
Jurimea Cass. ........c..ccccccccccccsesececeeees 538
Jurinea sect. 1:0. ak ee eee eee 672 -:

132

Jurinea § 1. Linearifoliae Boiss. ..... 578 — bucharica Wink. ............0......00. 611
-— § 2. Involucro cylindrico Ldb..... 600 — bucharica var. integrifolia Wink. ....
=~.§.3..PinnataesBoissM eee eee sn RR DR eee 616
ies 578, 600, 672, 677 — caespitans Iljin. ...............0.0 587
— § 4. Pinnatilobatae DC. .............. 600 — caespitosa Winkl..................00 619
— abolinii [Hyjin. oes 584 — calcarea Klok. :........... cece 665
— abramovii O. and B. Fedtsch. ..... 703 == Capusii-Franch.22n) .2e2 ene 588
— abramovii Rgl. and Herd. ........... 652 — carduicephala Iljin. .................... 382
— adenocarpa Scherenk. .................. 687 — cartaliniana BOiss. .................006 577
— alata (Desf.) Cass. ......cceeeseeseeee 673 — cataonica Boiss. and Hausskn. .... 643
— albicaulis Bge. ............ccceeeeeeee 554 — cephalopoda Iljin. ..................00.. 610
— albicaulis x J. cyanoides Schischk. ... — chaetocarpa Ldb. ...................000 680
Sih ae IM seas OA 548, 563 — chaetocarpa subsp. dzhungarica
— — B. longifolia Ldb. ................... 556 Rubtzi0esc03... SR eee 683
— almaatensis Iljin. .........0080.00 690 — — var. adenocarpa Herd. ........... 687
— algida Ijin... cee eeecsseeseeeees 691 — — var. macrantha Herd.............. 691
—— ssp. fetissowii (Winkl.) Iljin. .... 692 — — var. typica Herd. .................... 680
— — ssp. kuschakewieczii (Winkl.) — charcoviensis Klok. ..................... 550
Wy iNhspssecodteieree RR 692 — chondrilloides (Winkl.) O.
— alpigena C. Koch. ...........:cccccceee 668 Fedtschie..20h) Seater 512
— altaica IIjin. ....... eee 559, 71 — ciscaucasica (Sosn.) Iljin. ............ 561
— AMDI_UA AUCE. ..........cceceseeseeeteeees 556 — coelestis Sosn. ...............00 675, 707
Be ee Cnc Sesecuastnd cack RO 579 — coronopifolia Somm. and Lev. ... 576
— amplexicaulis (S. G. Gmel.) Bobr..... — cretacéa Bge. ........cecceeesesseeessees 661
ees desacthas eta Nm A erate 558 — creticola Iljin. .......................555, 717
— androssovii Iljin. ................:0:e 590 — cyanoides Kryl..........:cccccescesseeees 554
— angustifolia Bge. ..........ccccccccccenes 554 — cyanoides (L.) Rehb. ................... 546
— annae SOSM. ...........cccccesssestseseesees 574 — cyanoides Schmalh. ...............0806 547
— antoninae IIjin. .............ceeeeseeeee 656 — — Schischk. 0000.0... cccceeeeseeeeees 548
— antonovii Wink. ................... REN 648 — — X J. eversmannii Korsh. ......... 550
— apoda Ijin. .............ccccesccceseseeeeees 619 — — x J. polyclonos Korsh. .... 554, 563
— arachnoidea Aauct. ..........cccccccceees 663 — — 8B. pollichiana DC. ................. 547
SS as BOISS) Seiccet. A, eee eas 667 — czilikinoana Mljin. ..............:ccee 700
744 Jurinea arachnoidea Bege................... 659 — darvasica Ijin. .......... eee eee 692
— arachnoidea LAD. ..............0.00006+- 668 — demetrii Ijin. 0.0.00... 671, 727
— — var. atropurpurea Winkl........ 640 — densisquamea Iljin. ...................... 693
— — var. caucasica Sosn. .............. 668 — depressa C. A.M. ......eccecceeseeseees 705
— — var. dubia Korsh. ................:. 659 — — var. biebersteiniana Trautv. ... 705
— — var. exuberans Trautv. ........... 670 — — O. and B. Fedtsch. ................. 691
— — var. lyrata Wink. ................... 691 — — a. pinnatisecta Boiss.............. 706
— armeniaca SOSM. ...........cecccsseeseees 600 — — x J. spectabilis Fisch. and
— asperifolia Ijin. ................0... wees 631 Mey nese os See 675
— atripurpurea Winkl....................... 640 — derderioides Wink. ..................... 645
— auscheriana DC. ...................000 644 — dshungarica (Rubtz.) Iljin............ 683
— baissunensis Iljin. ....................005 642 — dubia Ijin. occ 659
— baldshuanica Wink. .................... 696 — dumulosa Boiss. ............::::cee 648
— ssp. hissarica Iljin. .................05 698 — eduardi-regelii Ijin. ..................... 587
— bellidioides Boiss. ...................: eaes70} — elegans (Stev.) DC. .............0.... 601
— bipinnatifida Winkl...................... 628 — elegantissima Iljin. ............... 608, 722
— blanda (M. B.) C. A. M. ............. 668 — eriobasis DC. ............ccccceeeeeeees 648
— bobrovii Ijin. ...... ee 655, 725 — eversmannii Be. ...............ee 549
— botschantzevii Iljin. ............. 615, 722 — eversmannii Korsh. ..................06.- 550
— bracteata Rgl. and Schmalh. ....... 701 —=exiMia Pekewnus. Mie. ae 646

— brachycephala Klok. .................... 581 — exuberans (Trautv.) Sosn. ccees sotee 670

733

— fedtschenkoana Iljin. ................+- 700 — macrocephala DC. ................:00+ 643
— ferganica (Iljin) Iljin. ................... 621 — margalensis Iljin. ..............:.000 682
=— filicifolia BOISS: <s2.44-2.-csceces-orameess 577 == mariae Pavilee sc. sedcieean tenets 702
— — var. akinfiewii Winkl.............. 576 — maxima Winkl. .............:ceeeeees 627
=f L Ay WANK et ceeer eset ee eae tees anes 711 — michelsonii Iljin. .................. 666, 726
— foliosa (Iljin) Ijin. ................ee 689 — microcephala BOIs.. .................-+ 709
— gorodkovii Ijin. .................. 554, 717 — mollis DC. ...........e0 663, 664, 667
— gracilis Ijin. ............ceeeeeeeeeeeeees 609 pe LAD) ceases se csctt st, sors eeeaes «es 665, 668 |
— granitica Klok. .............ceceeseeeees 552 ——— —— Schmalht -.....-<. cessdsur -secas ees ODD)
— grossheimii Sosn................0.000 675 — — var. arachnoidea Ldb. ........... 668
— grumosa IIjin. ..............:c0seeeeeeee 694 — — var. cretacea (Bge.) Fedtsch. ... 661
— hamulosa Rubtz. .............. eee 636 — — f. dubia Korsh. .............00cccceceee
— helichrysifolia M. Pop. ........ (Ceo ee: VEE > MRE cece reer rere © 659, 663, 665, 667 .
— heterophylla auct. ............0100eee 587 — — y. arachnoidea Korsh..............-..-.
—heterophylla (Jaub. and Spach) eve sesetersserentens 663, 664, 665, 667
BOIS ie: ee iea sede aee els rasta sy 579 — mollissima Klok.................::0:000 664
— hissarica Win. ..............0scceeeees 696 — monticola Ijin. ............eeeeeeeee 703
— horrida Rup. ............::cc0cceeceeeeees 361 — moschus (Habl.) Bobr. ................ 705
— iljinii Grossh. ................:eceeeeeeeees 571 — mugodsharica Iljin. ................0..5 596
— impressinervis Iljin. .................... 618 — multicaulis DC. .............. eee 643
— kamelinii Ijin. ..................... 607, 721 — multiceps Ijin. ............. eee 596
— kapelkinii O. Fedtsch. ................. 582 — multiloba (L.) B. Fedtsch. ........... 579
— karatavica Ijin. «0.0.0.0... eens 615 Sa IM, cccse sere oessdesrecasen tee. ete 682
— karategini Lipsky. ................:00++ 710 — — ssp. aulieatensis Iljin. ............. 683
— karabugasica Iljin. ..................0 647 — — var. aulieatensis lljin. ............. 683
— kasakorum Ijin. .................-. 662, 725 — — var. longicorolla lljin. ............ 684
— kazachstanica Iljin. .............. 562, 718 — nargalensis Iljin. ..............:::c0e 682
— kirghisorum Janisch. ................... 698 — NeUMAYEFIANA VIS. ..........c.00e0ceeeee 74
— knorringiana Ijin. ................00 656 — nivead Wink. ..........c ce ceeeeeeereeeee 694
— kokanica Ijin. .....0... eee eee 592 — ablongifolia Boiss. and Hausskn. .....
— komarovii IIjin. «2.0.2.0... eee G06, | RR dc Re et oes 644
— krascheninnikovii Ijin. ................ 686 — olgae Rgl. and Schmalh. ............. 620
— kultiassovii [jin. .........0........ eee 649 — — var. ferganica l\jin. ................ 621
— kuraminensis Ijin. .................0..0. 622 — orientalis Ijin... eee 598
745 — lanipes Rupl.............. ee eeeeeeeeee 690 — paczoskiana Ijin. .................e 594
— ssp. foliosa Nin. «0.0.0.0... eee 689 =sspersimilis, Uji it). stcttetn’..mkea as 680
— var. pinnatipartita Korsh. ............. 691 — pineticola Ijin. .................... 563, 719
— var. typica Korsh. .............. eee 691 — pjataieviae Iljin. .........0. 629
— lasiopoda Trautv. ..............:ccee 614 — poacea Njim. eee eeeeeeeeneenee 695
— — f. integrifolia lljin................... 608 — pollichii Koch. ..............:ceeeee 547
eo AAS NIS CD 8h saanehceonan Ropes cae 556 — pollichii Ldb. .............. 547, 548, 563
= IEGEDOUT BPC. .::. nce, sbae-seaateoae 663 — — var. suidunensis Wink\........... 681
— ledebouri Stev. ............:cccceeeeeeeee 667 — polyclonos autct........... 561, 566, 567
— leptoclada Bornm. and Sint. ....... 712 — — B. Fedtsch. ...........:cesceseeeeeee 562
AME VISTI AND eace cows cont nence sh 575 => — Clausds.2 doce: hen B. weiss 58
Fi | FCTGYEN VHT eM ee vetene ees een 650 Se DC, ccscccevizisscle ees 558
— linearifolia DC. ............::::cscceee 579 ar Kosh: .sac:ccacsnteered.pesteuseriaects 563
SMM Sky lalpiny os tocsresees ac ocn, ons nes cae 687 === Ktyl....2...0..cntth eo eee 554
— lithophila Rubtz. ................... 637 eS D,.. on. sasce scans eeeeceed acest 559
— litwinowil Zing. ............. cee 661 — — Schmnalh. ....... cee eceeseeseeeeeeees 564
— longicorollaris Ijin. .................4 684 — — ssp. ciscaucasica Sosn. .......... 561
— ludimilae Tjin. ........ eee 651 — polytaxis var. longipedunculata
— lyrata Trautv. .0...... eee eeeeeeeee 707 Winky. csscycorceessseee te ees 537
— macranthodia Ijin. .................0 591 ==popovii Ijin. ...:.:.......aceet eee 610

734

— propinqua IIjin. ...... eee 632
— psammophila Iljin. .................0 612°
— pseudocyanoides Klok................. 547
— pseudomollis Klok. ................000++ 663
— pteroclada Ijin... 701
— pulchella (Fisch. and Mey.) DC. ... 581
2S pumila Albuerceest Resa 569
— — var. major Alb...........::ccce 569
— pungens BOSS. .............:ccsseeseees 648
— ramosissima DC. ............:ecceeeeeeee 648
— rhizantha DC, .........cccccscceeseeeteeees 706
— robusta Schrenk. .............e eee 605
— roegneri C. Koch. ..........ccc:cce0 667
— ruprechtii Boiss. ........c.:..cceeeeees 676
— salicifolia Gruner. ................0005 552
— sangardensis Iljin. ........................ 641
— schachimardanica Ijin. ................ 585
— schischkiniana Iljin. ............. $48, 716
— semenovii Winkl................:.:0 635
— serpenticaulis Ijin. .........0..0..... 630
— serratuloides Iljin. .............0...0..0 638
— sintenisii Bornm......................005 597
— sordida Stev. .......:cceecceeeseeseeens 667
— sosnovskyi Grossh. ..............::05 574
— spectabilis Fisch. and Mey. ......... 674
— — var. monocephala Trautv. ...... 675
— spiridonovii Iljin. ................. 702, 724
— spissa Tjim. ....... eee eeeeeeteeteeeee 699
— stachelinae DC. ......0..0..0..cccee 648
— stenophylla Ijin. ........0.. eee 586
— stoechadifolia (M. B.) DC........... 580
— squarrosa Fisch. and Mey. ......... 707
— subacaulis Fisch. and Mey.......... 706
— suffruticosa Fedtsch. ................0.- 598
746 Jurinea suffruticosa Rgl.................... 595
— — var. latifolia Winkl. ................ 597
— — var. major Wink. ................... 595
— suidunensis (Winkl.) Korsh. ........ 681
— tadshikistanica Iljin. ............. 617, 732
— talijevii Klok. ......... cee cece 660 ©
— tanaitica Klok. .......0.0..... 566, 719
— tapetodes Ijin... eee 654
— tenuiloba Be. ............:cccceeseeeee 565
— tianschanica auct. ..........0cccccccccees 635
— tianschanica Rgl. and Schmalh. .. 688
— thyrsiflora Klok. ........0..00....cc 567
— tortisquamea Iljin. ..........2......e 590
— transhyrcanica Iljin. .................0... 645
— transuralensis [ljin................ 559, 718
— trautvetteriana Rgl. and Schmalh. .....
Began URNA ENN EIN STD AEN 608
— trifurcata Ijin... eee 593
— venusta I]jin. ............ cece 572, 720

Swinklent [jinn AEs 616, 722

— woronowii IIjin. «0.0.0... eee 570
— xeranthemoides Ijin. ................... 623
— xerophytica Iljin. ................. 549, 716
— zakirovii Iljin. ..................- 634, 723
Jurinella Jaub. and Spach................ 704
— aucheri Jaub. and Spach. ............ 705
— chamaecynara Jaub. and Spach. ... 706
— moschus (Habl.) Bobr. ................ 705
— squarrosa (Fisch. and Mey.) Iljin. .....
Dodane nasa MRE Mie Cea ae 707
— subacaulis (Fisch. and Mey.) Iljin. ....
pinnae algint ae eI ese ecereases tect 706
Jurineopsis Juz. subsect. .................. 324
Jurineopsis (Juz.) Tschern. sect. ....... 324
Kokuria Juz. subsect. ..............::00 313
Kopetdagia Tscherm. sect. ................ 266
Kuekenthalia Juz. sect. ...........:.0.006 256
Kusnetzowia Caucasica Winkl. ex Lipsky.
Hoth a EB ammcy Mins ait 75, 76
Lagurostemon Cass. .............00-++ 361, 392
— liatroides Fisch. ex Wyd.. ........... 393
— pygmaeum 8. Schanginianum Wydl.
teastatog: NUR a SNM corcate eeeaceeme 397
Lampocarpae Wink\. sect. ................ 278
Lappar Adansy ern cre Sener etetes teers 93
— ambigua Celak...............c.cc00000 105
— glabra y. caucasica C. Koch. ..... 102
— intermedia Rchb. .............:--21000000+ 99
Sat SC Ure Reh tasteless 102
— janczewskii DyD. ..........:.:0:cc0cecee 102
— macrosperma Wallt. ...............000+ 99
—— MAjOr GaeIt. .........cecceecesceeseeees 97
— — X L. tomentosa Hausskn. ....... 105
—SMINGH Hie ines esceeecooences 100
— minor x L. tomentosa Potc. ........ 106
— minor var nemorosa Crép. ............ 99
— mixta Simomk. ............:ecedceeereeeees 106
— nemorosa Koern. ......... ss NSW at 99
— notha RUNM. .............éccceeceeseeenees 102
— palladinii Mate. ............::00c.0cc0e 103
— platylepis Boiss. and Bal............... 98
— ritschiana ASCHETS. ...........00000000+- 106
— subracemosa Simonk. ................. 102
— tenella Fisch. and Mey................ 333
— vulgaris Hill. c...ccccciecceeenceeceeeeeeeeee 97
— fomentOSa AUct. ..........-.cecceeeeeeees 103
Ss earlier SEE ee creeces te 104
Lappaceae Bge. sect. ................ 177, 180
Lasiandrae Bge. sect. ...............00 258
Leiocaules Bge. sect. .............00c00 152
Leptocephala Tschern. sect. ............. 146
Leucocau!on Tschern. sect. .............. 144

747

Liatris baicalensis Adame. ................ 393
Lipskyella Juz. ..............ceescseeeeeeees 357
— annua (Winkl.) Juz. .......... ee 353
Longipapposa Chrtek, sect. ............... 60
Lopholepis Tschern. sect. ................. 236
Lyratae Sosn. subsect. .............:.006 672
Macrocephalae Bge., sect. ............... 259
Macropterae Takht., sect. ................. 277
Malacoloma JuzZ., sect. ...........00...000+ 183
Microcarpa JuZ., S€Ct. .......:.cccceeee 278
Microcarpae Bge., sect. ...............2 278
Microcousinia Tschern., sect. ........... 323
Minutae JuZ., Series. ................0000000 323
Mitina Adan. ..........::ccccceesceeeeeeeeeeeee Oil
Molles Bge., S€Ct. .........cecceeeceseeeeeeee 183
a WOrsh:esectycs.) ee Bee 657
Nanae Iljin, sect. ......... eee 651, 725
Nanarctium Juz., sect. .........:..0c0.00 176
Nanechinops Bge., sect. ................00-- 52
Nikitinia Ijin. «0.0.0.0... cece 712
— leptoclada (Bornm. and Sint.) Iljin.
Lene cel} ean ee 712
Nudicaules Wink, .......0...ccccccccccccceceseeee
sect, <2 152, 175, 244, 321
Odontocarpae Winkl. sect. ................-++-
PE BBs as 220, 266, 278, 334
Olgaea Iljin, Sect. ....... cee eeeeeeeeeeeees 620
Olgaeanthe Tschern. sect. ................: 165
Oligantha Juz. Sect. ..........:ceeeseeeees 242
Oligolepis Bge. SeCt. <...........ceceseeeeee 10
Onobroma cynaroides Spreng. ......... 262
— macrocephalum C. A. M............. 263
Onopordon lyratum Schulz. ............. 705
— polyacanthum Eichw. .............0+.. 260
Orthacanthae Wink. sect. ........ 144, 187
Papilligerae Iljin, subsect. ................. 606
-Pectinatae Winkl., sect. ..............00000. 238
Penduliflorae Iljin, sect. ............ 602, 721
Perplexia Ijin. .............c:.eeee 709, 727
— microcephala (Boiss.) Ijin. ...............
A REC og een ae 709, 727
Persistentes Ijin, SUDSECE. oo. secssceeeeees 608
Pilostemon Iljin. ..................0ccee 709
— filifolia (Winkl.) Iljin. ......0000.0.. 711
— karategini (Lipsky) Iljin. ............. 710
Planta salsa ambigui generis Pall..... 504
- 'Platycephalae Benth. ...........00.0c0000000- 643
— subsect. Floccosae Sosn. ............. 600
— Benth. and Hook. ........................ 672
Pleacme Bornm. S€Ct. ............:.000c0c0000 39

Polytaxis Bge. ......... eens 536

735

— lehmannii Be. ..........ccccceeeeee 536
— — var. longipedunculata Wink. ......
AS ANS son cleduricineine ses 537
= winkleriAlpin Baek eae 537
Pseudactinia Tschern., sect. .............. 173
Pseudarctium Juz., Sect. .........:.:000 177
Psilacanthae Boiss., sect. ......... 278, 333
Pugioniferae Bge., sect............. 187, 277
Pygmaeae Juz., Semies. ............:0 167
Rectispina Boiss. S€Ct. ...........cc0000006 187
Recurvatae Bge. sect. .........:::.000 160
IRetrovkEndl:-vsecte earner 30
(RIO BOSS Ue. Ee 40
Ritrodes Bgewisect +2 ee ee. 16
Robustae Ijin, subsect. .................... 605
Saussurea DC. ............:0:cccsceeeseeeeee 361
— subgenus Amphilaena (Stschegl.)
Tipschi se ees AR 383
— — eriocoryne Wall. ..............0.... 389
— — eusaussurea Hook. f. ............. 392
— — frolovia (DC.) Lipsch. ............ 384
— — SAUSSUTEA. ....... ee eeeeeeeeeeeeeeeeeeee 392
— — stephanodontos Lipsch........... 382
— — theodorea (Cass.) Lipsch. ....... 520
— sect. Amphilaena Stscheegl. .......... 383
— — benedictia DC..............0.0 428
— — subsect. Hololepis Lipsch. ..... 429
— — subsect. Odontolepis Kitam. ... 428
— — bracteatae Clarke. .................. 383
— — caulescens Hook. f................. 392
— — chaetocline Franch. ................ 428
— — congestae Clarke. .............2.... 389
— — depressae Clarke. ........... Sess
— — elatae Hook. ...........ceee 520
— — eriocoryne Wall. ............:000008 383
— — eusaussurea C. A. M............. 428
— — frolovia (DC.) Kitam. ............. 384
— — gymnoclyne Franch. ............... 515
— — jacea Lipsch. ........... ee 513
— — jurineiformes Lipsch............... 512
— — laguranthera (C. A. M.) Lipsch.
BAe a SN I ER eves 403
— — — subsect. Laguranthera C.A.M.
IYER SOREN ek eI 403
—_—— Luptivastemol (Cass.) BOA a
MP di N, AEE SO 403, 428
— — Obvallatae Maxim.......x......... 383
— — Pycnocephala Lipsch. ............ 392
— — Strictae Clarke. ............0...0 392
— — Theodorea (Cass.) DC. .......... $20
— — subsect. Theodorea-vera Kitam. ~
Ss a, Shere 5205 522

736

— acuminata TUrcZ. .........ceeecceseeeeee 454 — — f. monocephala Lipsch. .......... 521
— acuminata Turcz. y. gracilis Herd. ... — — amara f. glomerata Trautv......... 520
SRE, ES oc satlsessowes 474 — amaurolepis FreyN. .................... 456
— — subsp. sachalinensis Kitam. ... 455 — ambigua Kryl.............ccce 491, 496
— — B. sachalinensis Herd. ............ 455 — — var. ramosa Lipsch. ................ 497
— ajanensis (Rgl.) Lipsch. ............... 44] — — var. subglabra Serg. ............... 496
alata ACh. Mises cot ee Rie 474 — americana Eaton. .............c:cccc00 478
= alata’ DC hee eats) 525 — amoena Kar. and Kir. ................: 423
— — var. pinnatifida (Schischk.) — amurensis TurcZ. ...........00:::.c000 456
Tei pseh wen ce. cdcovesiecest dees ecaece $27 — AMUFENSIS AUC. .........0ccccececereceeeeee 495
— alata B. aspera Ldb. ........c.c0000000 526 — — subsp. duiensis (Fr. Schm.)
— alata DC. a. genuina Herd......... 526 Kitamn:....:. us, att ueriacee 458
— — Q. laciniata Herd. ................... 529 — — subsp. stenophylla Kitam. ...... 456
— — y. pygmaea Fisch.............00000 528 Ea aval Nakai... pene eee 456
— — 6. runcinata Herd. .................. 528 — ongarensis Lipsch..............0.0.006 496
— alberti Rgl. and Winkl................. 426 — angustifolia AQQTl. .........ccceceescceees 500
— alpina auct. ..........ccccccccceeeees 493, 495 — angustifolia auct. ............c000c0000 $18
— alpina (L.) DC. ....... ec eeeeseeeee 489 — angustifolia (Willd.) DC......491, 498
— — x §. tilesti Ldb. ..........cccce 491 — aphylla Rech. f. ........c.ccccccessecseees 512
— alpina subsp. esthonica (Bear ex — approximata Schischk. and Serg. .....
Rupr.) Kupff. ............:ecceceesee G5 AN APE PCE Mi nce ero a. 389
— alpina subsp. Stubendorffii — asbukinii Ijin. ........... ce eeeeeeeees 387
(Herd.) Lipsch. .............:0cc000 494 — atriplicifolia Fisch. ..............0.0006. 472
— alpina foliosa Herd. .................... 479 — baicalensis (Adams) Robinson. ... 393
— — Sibirica C. A.M. .....ccceecccecceee 490 — blanda Schrenk. .................000008 407
— — var. angustifolia Rgl. ............. 501 — — f. umbrosa Iljin....................... 407
— — var. decurrens Rgl. ex Wink. ..... — borealis Fisch. ............ccccccccceseeeees 498
PR AEs oho as. 490 — brunneopilosa Hand.-Mazz.......... 397
— — var. glabra Ldb. ...............0065 491 — caespitans Ijin. ........... eee 417
— — var. krylovii Serg. .............0+ 491 — calobotrys Diels. ....................000 393
— — var. kuschakeviczii Winkl. ..... 502 = eanai edb... eestor td aierens 413
— — var. lancifolia Serg......... 492, 495 — — var. latifolia Lipsch. ............... 414
— — var. ledebouri Gray. ............... $00 — cana B. angustifolia Ldb............. 413
— — var. leucophylla Turcz. .......... 492 — canescens WinKl. ...............:0:000 418
— — var. schischkinii Serg. ....492, 495 — — var. major Wink. ............ seeeees 418
— alpina var. subacaulis auct. ........ 484 — caprifolia Iljin and Zapriagaev. .... 519
_ — alpina var. viridis Turcz. .....490, 491 — carduicephala (Iljin) Iljin............. 382
— alpina f. angustifolia Kjelim. ...... 500 — centauroides Tausch. ................+.. $20
— — Q. communis Turcz. .............. 493 — ceratocarpa DCME. ............c0cce 382
—=-—= B: subacaulis. Ldb:.2...0cccc2.... 480 — ceterahifolia Lipsch. ...:................ 532
748 Saussurea alpina B. vulgaris Ldb. ......... — chondrilloides Winkl. .................. 512
AOD SEMA apnea Dace 490, 491 — chthonocephala Bormm. .............. 389
— — 6. hypoleuca 5. major seu — coarctata (Herd.) Kitam. ............. 464
genuina Herd. ............0:ccc0000 492 — colorata Wink. ...........:scccceee 510
— — 6. — — a. minima Herd. ....... 480 — compacta Fisch. .............::.:ceceee 454
— — 6. — —c. minor Herd. ......... 492 — congesta TUrcZ. .........:..:cceseesceeees 482
— — 8. maxima LAD. ......ceecccccceees 487 — congesta Turcz. x S. pricei
— &. hypoleuca Lb. ................0000006+ 492 Simps: aet)tsee Sh eto 482
— altissima M. Pop. .............100000000+ 533 — — f. elatior s. Kusnetzoviana
——tamara (EDGE... eee 520 Herd irc. anvils cooled 482, 484
— amara var. glomerata (Poir.) — congesta a. humilior s.
Wrautvics teint Yi eeeeths ait sa. 520 Kiriloviana Herd. ........... 482, 484
— — var. integrata DC. .................. 521 — contracta FreyM. ..........c::cc0ccee 460

— — f. albiflora Lipsch. .................. 521 — controversa DC. ..............::eceeeeee 438

737

— — x §. alpina DC. ...... 442 — eopygmaea Hand.-Mazz. ............. 397
— — x §. jadrinzevii Kryl. ............. 439 — eriolepis y. caudata Herd. ........... 466
—-— . S. parviflora (Poir.) DC. ..... 439 — esthonica Baet. ...............006+5 491, 493
— coronata Schrenk. ................ 406, 407 == excelsa IM. sROp srstszeseeesies. 2002-252 533
— coronata var. colorata Herd. ...... 406 — faminziniana Krassn. ................... 510
=—S ChASSHON AWC. css gears. § cecseunee es 504 =— fauricighranch et toes ees, ee 476
— — f. laciniosa Kryl. ..............0.4. 508 — filfolia Rgl. and Schmalh. ........... 711
— — B. muricata Be. ..............0..+. 506 — firma (Kitag.) Kitam. ................... 448
— — fp. papposa Ldb. ............006 507 -=* foliosadb.........eee ae Bees 479
— crepidifolia Turcz. ..............00 527 — — var. altaica Turcz................... 480
—— daurica Adam. ............:ccsseeeeeee 507 — foliosa var. dentosa TurcZ.................
— decurrens Wink]. ...............000000008 AGO i Vian ae ie Bie sacle eR | aes reece 437, 492
— densa Rydb. ...........e eee eeeeeeteees 491 — foliosa var. glabrata Kryl. ........... 480
— denticulata Ldb. ............e eee 456 — foliosa var. tomentosa Krty]......... 480
Be UTC Zia Maeendideseddeeadosdocevaare 492 —— V. pygmaea TUICZ. ...........00 480
— — Ldb. a. genuina Herd. ........... 456 — formosana Hayata. ..........-..:000+ 386
— — B. duiensis Herd. .................... 458 eS hiCIdaQybISCh., «.:.+-.een to 486
— — v. viluiensis Herd. .................. 495 —— frolowiijldbiget See eee 384
— depressa Gren. and Godr. ........... 491 — frolovii var. procera Herd. .......... 387
— derbeckii Kom. ...........::ccccecccceeees 469 — fruticulosa Kar. and Kit. ............. 413
— discolor auct. .........cccccsceeseeeeeeeees 440 — gebleriana C. A. M. .........:.:006- 416
— discolor (Willd.) DC.................... 439 — glacialis Herd. ...............:ce:eeeeeeee 389
== GISCOLOTNDE, BAe sce scese. soins wes 438 — glomerata BOI. ..............0eeceeee 520
— — ajanensis Rl. .........:ccecceeeeees 441 — — var. angustifolia Freyn. .......... 520
— — B. elatior DC. ......e eee 438 — — var. intermedia Freyn............. 520
— — f. s. sibirica (Fisch.) Herd. .... 438 — gmelini Hort. .0.......ceceeseeeeeeeeeeee SZ
— — Q. genuina s. europaea Herd. ..... — gnophalodes (Royle) Ostenf. ...... 390
issih ai le tte atacand 439 — gnaphalodes (Royle) Sch. Bip. ... 390
— dissecta LAD. ...........c::ccccceeeeeeeee 523 — graminea Dunn. ................::000+ 397
— dorogostaiskii Palibin. ................. 394 — graminifolia Wall. .........0....0..... 397
— dshungarica Iljin f. canescens — grandifolia Maxim. ...................... 464
MUI. Sececacccewnceerene eee 404, 405 — grandifolia var. caudata (Herd.) Kom.
— — f. viridis Njin. ................. 405406 BER user te bemeie, & ee S. 466
— dubia Freyn. ............. ce sceeeeeeeeeeee 460 — — Q. genuina seu maximovici-
— duiensis Fr. Schm. ...................02- 458 Ziana Herd......)190 edie 464
=—=4 CL ALA AUCH ete. cass ssedaock eaeaeestestvees 472 — — f. tenuior seu Raddeana Herd. ...
==jelatasledb: 222k... dei eens ADD RI ca cece ebarcece a sace. nouns terete des Wes 464
— — f. albiflora Lipsch................... 422 —— 6. coarctata seu wilfordiana Herd.
=—elegans db: .. tise. 2. gett: 423) ARB teas tn et eee Bos 464
749 — — ssp. turkestanica Lipsch. ........ 423 — — Vv. asperifolia Herd. ................ 464
— — var. amoena (Kar. and Kir.) — hypargyrea Lipsch. and Vved. .... 440
Serp S22 ious. Gs hesvar cess 423 — — x §. pseudoangustifolia Lipsch....
— — var. latifolia Kar. and Kir. ..... A ecco nt steel, cece ete 292. 2 440
— — var. nivea Lipsch. ........... 423, 424 — Intermedia Freyn. ............. 456, 460
— — var. tenuis (Ldb.) Serg. ....... 423 — involucrata Kar. and Kir. ............ 383
— — f. albiflora Lipsch................... 425 — involucrata (Kar and Kir.) B.
— elongata AUct. ..........cesceeeceeeeeeeeees 445 Fedtschy, <ccic:s.osctsth-s see 383
— elongata DC. 0... eee 444 — ispajensis IIjin. ................:eeeeee 409
— elongata DC. a. genuina Herd. ... 444 — jacea (Klotzsch) Clarke. .............. 514
— — B. polypodiifolia DC. ............. 444 = jadrinzevii Kay]. .....0..04:..020...0. 429
— — B. recurvata Maxim. .............. 445 — — var. intermedia Serg. .............. 430
— — 6. clehniana Maxim............... 445 — JAPONICA AUC. ........ceceeceeeeeseeseeeees 523
— — 6. pectinata Herd. .................. 445 — — (Thunb.) DC. .......... eee 522

— — scabiosaefolia Turcz. ............. 444 — — var. alata (Rgl.) Kom. ........... 523

738

— — var. ovata (Rgl.) Kom. ........... 23") — macrocephala LESS. ...........0:..0006+ 520
— — var. pterocaulis Franch. ......... 523 — macrophylla Be. ........1....00cc00000 478
— — var. pinnatifida (Rgl.) Kom..... 523 — manshurica Kom. ..............:c00008 471
— — var. subintegra (Rgl.) Kom. .... 523 — manshuriensis Kom. ................-..+ 472
— — B. alata Rgl. ......e eee $25 — marginata BOrsZeZ. ........-.00000 520
— — Vv. latifolia (Maxim.) Kom. ..... 525 — masarica Lipsky. ..............cc:cceee 513
— karaartscha Saposhn. ........,.......... S11 — maximoviczii Herd. ..................... 448
— karategini Lipsky. ...2.......0....:00055 710 — mikeschinii Iljin. ...........000.... 408
— karelini Stschegl. ............0:c.00008 383 — microphylla C. A. M. ........10.00006+ 413
= KAFOL EVEN. 20s. eee 462 — miyagii Miyabe and Kudo. ......... 455
— kaschgarica Rup. .............cc0 509 — MONGOlica AUCt. .........cccccceceeeeenees 468
— kitamurana Miyabe and Tatewaki..... — morozeviczii B. Fedtsch. .............. 516
Cas ORS NOL MING 432 — multiflora DC. -...c.cccccecece 410, 412
— klementzii Lipsch. ..........00.0.00.... 498 — nakaiana Kom. ............:020+ 431, 434
— komarovii Soczava. .........:.c0:0c000 432 — neopulchella Lipsch. ..................... 325);
— konuroba Saposhn. ..................6+ 407 — neoserrata Nakai. .....0...0...eceee 475
— korotkyi Lipsch. ..............cccccceeee 482 — — x S. umbrosa Kom................. 476
— krylovii Schischk. and Serg. ....... 396 —ninae Iljin. .....00 404, 407
— kurilensis Tatewaki. ..............00..... 452 — nivea Kom. and I. Schischk. ....... 431
— — var. arachnoidea Tatewaki. .... 452 nda hdbe i. A eee 515
— kuschakeviczii Winkl. ................. $02 — — x §S. oxyodonta Hult. ............. 518
— — var. caulescens O. Fedtsch. .... 502 — — x §S. pseudo-tilesii Lipsch. ..... 518
=—laciniataldbse a ee $29 — — var. exaltata (Maxim.) Trautv. ex
— laciniata var. lasiocephala Turcz. .... Fedisch: 4232) See 487
Sate TET A ROR TEM oo 526 — — var. subsinuata B. Fedtsch..... 516
— laciniata Ldb. var. latifolia — nuda Ldb. f. uniflora Lipsch. ...... 518
bipscht( (Se Pee ees 532 — nuda a. genuina Rgl. ...............--- 515
— laciniata var. pinnatifida — — f. subsinuata Rg. ................. 515
Schischky 3.2) ee ae 527 — — jy. tilesii Rg. oo... ee 484
— laciniata Ldb. var. pygmae Lipsch. .. — nupuripoensis Miyabe and
Deck A TEE Be 532 Miyake tse ae nerd 0)
— — f. ramosa Lipsch..................... 532 — obvallata Edg. ............cceeeeeeee 384
'— — Ldb. f. simplex Lipsch............ 530 — odontolepis Sch. Bip. .................. 428
— lanicaulis Hand.-Mazz. ................ 397. — odontophylla Freyn. ...................- 456
=arionovil’ Winkle 425 = ovata Benth a. sues 503
— larionovii var. nivea Lipsch. ....... 425 — oxyodonta Hult... 488
— latifolia Ldb. «0.00.0... ceeeeseeeees 478 — pamirica Winkl..............:0cc0c 389
— — f. exalata Kryl. ..... 478 — PAPPOSA TUNCZ. .0......eccceceeeseeteeeees 507
— — f. pubescens Lipsch................ 478 — x paradoxa Lipsch. .................0... 506
— ledebouri auct. ..........ccccccccccseeee 484 — parviflora auct. .............. SRR oes 474
= Ona ORE | Pee 515 = parviflora (Por) DEE. ee 472
— — B. subsinuata Rgl. .................. 515 — — x §. stubendorffii Herd. ......... 474
— — v. kruhseana Herd.................. 488 — — ssp. purpurata (Fisch. ex Herd.)
750 Saussurea Ledebouri o. Tilesii Rg. ....... erpscha teers ea et 473
tetas TR, RS PE 484 — — var. incisa Lipsch. ..................474
— — &. exaltata C. A. M. and — pectinata Bee. ........ Pest reccerey 429
Maxim see) ee ae 487 — pectinata B. amurensis Maxim. .........
— lenensis M. Pop. ............::::0c000000 ay RRR a cI aie entering 428, 446
— leucophylla Schrenk. ................... 399 — petiolata Kom. ..........::ccccceeeceeeees 465
—=+— f. caulescens: Lipsch. 103% 400 — pohlei Gandoget. ...............e 490
— — f. subacaulis Lipsch................ 400 — poljakovii Glehn. «0.0.0.0. 483
— liatroides Fisch. .........::ccccccceeeees 393 — polyphylla Schrenk. .............:.0+ 398
==lioui Ling Yong. ...252. 2008 383 — polypodiifolia auct. ............0000. 445

— lomatolepis Lipsch...............0..0-. 412 SANTOS Se VME ARE hs 444

139

pete UTZ ase ee 528, 529 — — y. heteromorpha Turcz. ........:.......
— — RB. angustifolia Turecz. ............. 529 Nee aa phones 396, 397, 398
— poophylla Diels. ................000 397 — — y. polyphylla Turcz. seu Turcza
—— POpovil’ Wipschs suv cee wk oe. 513 ninowiana Herd. ...........000005 398
— porcellanea Lipsch. .................004. 431 — recurvata (Maxim.) Lipsch. ........ 445
— porcil Degen. .............ccsccesseeeees 474 — — var. Glehniana (Maxim.)
Be picchSInCS 480 Lipsch. Le Adis ccaveaes Mires Ree AO,
als 2 = leden Herd! es See ee 453
— propingua Ijin. ............:ccccceseeseees 438 : nue
Winkl — — subsp. yezoensis (Maxim.)
—PEOSEEL TE yh ee ee 334 Kitam! hecoseonnenl sae. 454
— — var. erecta Winkl. ......00..00.0.. 534 — riederi var. yezoensis Maxim. ..... 454
— — var. exscapa Winkl................. 534 == ripidaiedb. ee ee ae 403
— pseudoangustifolia Lipsch. .......... 501 robusta ledby 200 ee ay o. 533
— pseudoblanda Lipsch. ................: 408 — robusta \db. var. discolor Rgl. and —
— psedofoliosa Lipsch. ............+..0+++. 478 Schmalhie, 2.22068 Voy) ne 534
— pseudosalsa Lipsch. ................04.. 504 PROC IMMMEMPSEN. 2... soees 406
— pseudosquarrosa M. Pop. and == OSCR KOM... Uae See 461
LipSCh. ..........0seessesessessessessseees 437 a> MUNCINGLA-AUCK.....c2-20c2nc eee ete 532
— pseudo-tilesii Lipsch. ................... 487 — runcinata DC. «......cccccccccsesessceccsees 528
— — var. alpicola (Kom.) Lipsch. ... 488 — — x §, salsa (Pall). Spreng......... 529
= pulchella RiSGh nh ee 522 — — yar. albiflora Herd. ................. 529
— pulchella B. alata auct. ............... 525 — runcinata B. dentata Ldb. ........... 527
— — var. alata Rgl. ........ceceeceeees 523. — russowii Winkl............ 400, 402, 403
— — var. dentata Kom. Sees ocee one 525 — sachalinensis Fr. Schm. ............... 455
— — var. lineariloba Nakai. ........... 523 — sajanensis Gudoschnikov. .....:..... 433
— pulchella Fisch. popul. paucice- — salemannii Winkl................. 420, 421
phalae Lipsch. ieteeteeseeertaceeeenes 525 — salicifolia auct. .........cccccccccceeeeees 423
— pulchella Fisch. popul. plurice- HS salicifolia(L.)sDG 1. Aa 410
phalae Lipsch. .................000 525 — salicifolia var. angustifolia DC. ... 412
—— a. subintegra Rgl. ... 522, 523, 525 — — var. elegans Trautv. ............... 423
— — 8. latifolia Maxim. ......0.......... 525 — salicifolia var. intermedia Ldb. ... 412
——y. pinnatifida Rg]. ... 522, 523, 524 — — var. macrocephala Turcz. ....... 410
— — 6. ovata Rgl. ......... 522.523, 525 — salicifolia var. macrocephala need
Ae. 1 . ift . Wi kl. pinecone, RI ROGY, 1 Yo SOURS arn sta A oliali 9 ECS CEC SCRE RES a eee aa
a Dee ee ot ay — salicifolia var. major Ldb. ........... 412
— pumilio Fisch. ...........c.ccceccceeeeeeees -480 Sener Fig
‘ — salicifolia var. ramosissima Ldb. ......
— purpurata FiSCh. ............c0ccccceeees Dame. eee eee OE he Ca oe ee 423
— pycnocephala auct. ............c0:00000 400 — — var. scabra Rgl. and Schmalh.
LAD. ose eects BBS BEO 6!) MIBK, silts act etc. cAI Lets at 423
— — a. genuina Herd. ................... 393 — — var. squarrosa BBE. ...........+: 414
— — a. papillosa Kry]l. ............00. 393 — — var. stenocephala Trautv. ...... 423
— — B. paleata Kryl. .......cece. 396 — — var. transbaicalensis Ldb....... 411
— — 8. sordida (Kar. and Kir.) — — var. viridifolia Winkl.............. 421
Herd pedt aie biG) ie 400 — = ¥- Encisa LAD. ose 414
— PYQMAEA AUCH. ..eeeeeeecseeeeeeee: sce 397 — — brevifolia TUtez. «1... see 412
7 PYSRIACAUS DIN een eee eee 397 — — € regione transbaicalensi Ldb. ....
— pygmaea vat. juldusica Winkl. ...399 «eet ttttettteteettsetertsseesettseereesseeeeees 411
751 ——a. communis Turcz. ............... 397 — salsa (Pall.) Spreng. .........1.........- 504
Pee, SPPICA RIGA rene ee 307 — salsa (Pall.) Spreng. x S. laciniata
"8. teucophylla Lab. 2... 399 edbiyaat a Diuheea RO Be
MS — — var. aspera inten | eta
B. polyphylla Schrenk. ........... 398 an Fipinnanpdialeleeid
— — B. schanginii DC.\ ................... 397 :
Ue Schmalhi224.). Bee ee 508
— — f. sibirica Herd. .................0... 397

— — var. fastigiata Winkl. ............. 504

740

— — var. laciniosa (Kryl.) Serg. ....508 | — sumnneviczii Serg. ........--....:e 414
— salsa. var. pamirica Lipsch. .......... 506 752 Saussurea sumneviczil var.
— salsa var. pinnatifida Wink_........ 508 communis Serg. ..........:ecee 415
— salsa f. albiflora Lipsch. ...... 505, 509 © — — var. mutabilis Serg. ................ 415
— saxosa Lipsch. ............ mdidos lem oeetes 497 — — var. sergievskajae Lipsch. ...... 415
— scabiosaefolia TurcZ. ..........:0000 444 — susuyensis Miyabe and Miyake. ... 458
— SCADA LESS. .......ccccscsesseesseesseseceaes 521 — tadshikorum Iljin and Gontsch. .... 514
— schanginiana (Wyd1l.) Fisch. ........ 397 — takhtadjanii Lipsch. .................... 425
— schanginiana (Wydl.) Fisch. — talassica LIipsCh. ...........:ccsccceeeees 504
var. heteromorpha Turcz. ...... 399 — tomentosa Kom. ............:eeeeeee 430
— schanginiana var. heteromorpha — tenuis LAD. .00.....eceeceeeesteeeeeeeeees 423
(Turez.) Lipsch. ...........::cccce 398 == tilesii-auct 24 Bee AE 487
— — var. polyphylla (Schrenk) Lipsch. filestt- Abies. LERARA 484
pe ee td sot en on 398 — tilesii var. exaltata Maxim. ......... 487
— — (Wydl.) Fisch. f. dentifolia Lipsch. — — var. kurilensis Kudo. .............. 488
a slusdaadeseeostes car abeaaeen &. Abaat teres 397 — — f. alpicola Kom. ..:.............+.-.. 487
— schischkinii Kom. ..............0000008 469 — — f. glabrata Kom. ..............006 487
— semenovii Herd. .............::seeceeeee 635 — — f. subacaulis Kom. ................. 484
— serawschanica B. Fedtsch. ................ — triangulata auct. ...........cccccccccerees 471
Lf er EAE 5 es 400, 402, 403 — triangulata Trautv. and Mey......-. 469
== serrata’ DORs Sentai: 472 — triangulata subsp. manshurica
= = auch tae PP 474, 475, 495 (Kom.) Kitam. ...........0000..000008. 471
— serrata var. contracta Ldb............ 473 — — a. genuina Herd. ...............06. 469
— — var. patens Ldb. ................0.. 473 — — B. elatior Herd........................ 469
— serrata vat. pygmaea Tutcz........ 492 — tschuktschorum Lipsch. ............... 518
— — c. amurensis Herd. ................. 475 — turczaninovii Lipsch. ................... 496
— — d. purpurata Herd. .................. 473 — turgaiensis B. Fedtsch. ................ 508
— serratuloides Turcz. ...............:0000 416 — umbrosa Kom. ............cceceeeeeereees 462
— shiretokoensis Sugaw. ................. 459 — x uralensis Lipsch. ..............00.0 442
— sinuata Kom................. Sey See 468 — ussuriensis Maxim. ..............000 446
— soczavae Lipsch..............:ccecee 486 — — var. firmifolia Lipsch. ....447, 448
— sordida kar and Kir. .................... 400 — — qa. genuina Maxim.................. 447
— sordida var. oligocephala Winkl. — — B. incisa Maxim. .................. 447
PEE ace eeene Petraes Axcahsabevet bess 402 — — jy. pinnatifida Maxim. ............. 447
— sordida var. russowii (Winkl.) — ussuriensis Maxim. o.
Lipschi cea 400, 403 odontolepis Herd. .................-- 428
— sorocephala auct. ..........ccccccsceeeee 389 — VITESCENS FL€YM. ...........ceceesceeteees 460
— — Hook. f. and Thome. ............... 390 — viscida Hult... 486, 500
= = Schrenkwt. 2 cde. 390 — vwvedenskyi Lipsch. ................0....- 420
— sovietica Kom. .....cccccceceeee 434 — zeaensis Frey. .........0.... 92. cer: 460
— splendida Kom. .............c..ccccc0e 443 Sciadocousinia Tschern., sect. .......... 186
— squarrosa Fisch. ..............c:cc00s000 414 Serratula alpina L. ...1..cccccecccecseeeees 490
— squarrosa TUICZ. ...........cc:cccceseees 436 == “alataiDes ie seh ee eee 673
— stenophylla FreyM. ...........c0:c00000 456 == GMARASS cov ctacesesassiveas eaters 520
— stubendorfii Herd. ................ 491, 495 — ambigua DC. ........ccccccesccesseseeseees ‘579
— — x S. parviflora (Poir.) DC. ..... 496 — angustifolia Willd. ...........-...:000 498
— subacaulis (Ldb.) Serg. ............... 480 — blanda M. B. ............:0::0000 659, 668
— subhastata Kom............. ator ~- 466 — chaetocarpa LAD. ............c0.cc000 680
— subsinuata auct. ........:c.cccccseesenees 488 — cyanoides DC. .........ccesceeseeeseeteees 546
pee SS Lb in PR Se tla | 515 eB BOP ANNA ruta cute a 548, 563
— subtriangulata Kom. .................... 465 Se PEMA. PARES. osapevet vs vas vessnvacttee: 554
— sukaczevii Lipsch. ............... 491, 492 — davurica Adams. ............00000cccce08 507
— — var. pumila Lipsch. ................ 492 — decurrens LAD. ...........:c1ccecereeeee 526

— sulcata Ijiti cil. .ccscecciseecceseseeess 388 === AEPTESSAN SUC Vine careers eee 705

— discolor Wild. ............cccccesscseseees 439
— dissecta Steph. ...........cccccccsccereees $29
— elegans Stev. ......ccccccccccsscesseesseeees 601
— japonica Thumb. ............c:cecccee 522
— liatroides Adams. ..............000000+ 393
— microcephala Trautv. ....... Pernt 145
NUL LON Nee 22.25 Sececeses Sec secceteciess 579
Sera De. cbse eetae cues Se aiics Seecdasoees 472, 474
— parviflora POAT. ...........c0.cceseeeeeees 472
— polyclonos M. B. ............0006+ 558, 561
— pulchella Fisch. and Mey. ........... 581
SBI Stee fete er lester sosevee ta mean cess 523
= ALG (0) Lah Ue eer re pee ECE 410
FMC DCC case esac tsk ence ascessanbonsesaesse 579
Wit eee ce ascectoses cds tagetalatcdaaute 580
Se SOCING Pall Lo. .csccccoseseMesdanse tee: 579
— SAlina Pall. ........ccccecccesceeseseeeseees 504
SAA PEN cece icin seceee cone eee ee eee 504
— salsuginosa Si€vets. ...........0.00000 526
= squalida Steph. ..............ccccccceeees 526
——SLELIEFE SLOP erecassecsnesasc kee seceteeees 444
— stoechadifolia (M.B.) DC. ........... 580
SHEL CSIP ACAD a cose sears soot edesceescesces eeeseee 484
ea PETICIO (CASS. tose, ccsstedetvestacensine tees 579
SEKTAlMICACMDE, 2 ce cccssseeeet oe eesncoesses 90
Serratuloideae Bge., Sect................+. 276
913 OLSS ESC CUS a aren eos Uae ese sere 278
Serratulopsis Tschern., sect. ............. 248
Schmalhausenia Wink. ................... 360
— eriophora Wink. ..............c0000006 361
— nidulans (Rgl.) Petrak.................. 361
Slebera HOppe. .........:c:.cccccecsssceeeseeees 67
Stebera Gaye: ccccslec tkeeccssssssevieoes 57
—= (HEED ( CEI s Ararend lececaonesiad ose eeu 68
— pungens (Lam.) DC. ...................... 68
— — BP. nana DC, o0....ccceccceccsececeeeee 68
ISIEDERIaESPTEN Be eee e esses saeseccaeceseenees 67
Sphaerocephalae Bge. sect. ............. 152
Sphaerocephalus Be. .........c0c00c000c000 40
See INIZESiee merece wen ery a Ler eel 2
== dauricus O. KtZe. v.......cstccsceosce-s 45
Spicatae Winkl., S€Ct. ............cc0c0e 168
Squarrosae BOiSS., SCCt. ..........60cccccccceeees
Pedi ee tne 146, 152, 160, 269
Stechmannia Boiss., sect. ................. 647
SECCHMANNIGND] oo ae ecrcca. cos 647
Stenocephala Benth. .............0.:0000060 647

Stenocephalae Benth. and Hook., sect.
ieee sie eee ME ed at SOs Fe, 578

753

741

Stenocephalae Bge., Sect. ................. 146
Stenoloma Juz., sect. ................00 a0)
Subacaules Boiss., Sect. ..........00...0000 567
Subappendiculatae Tschern., subsect. ....
So eaces SRA ncrs Une Aen nse EM 319
Suffrutices Iljin, sect. ................ 594, 721
Tenellae Bge., S€Cct. .........:...0ccc00000s0060 333
HenmasEndle secthee ee 47
Theodorea Cass. «0.1.2... 361, 520
= pil cnellalCaSsweteres seen 523
Uncinatae Winkl., sect. .........0000.......e000
SEE Cee 177, 180, 182, 238
Xeranthelia Chrtek, subgen. ............... 60
Xeranthemeae DC, .......0..ccecccseceeeeeeeees 57
Xeranthemum L. ......................... 57, 59
Xeranthemum, subgen. ...................... 60
Xeranthemum, SECCt. ............cecceeceeee 60, 62
— annettae Kalenicz. ................... 62, 64
= PATI TLULUL IT [ecstecseeenee ee eeee 60, 62, 64
— — f. gracilius Fisch. and Mey. .... 62
— — f: inapertum L..........:......... 64, 66
— — y. orientale Ly ..........60ccccscccccees 70
— cylindraceum Sm. ....... 60, 62, 66, 67
EACHECLUIM DINOS lessens sseescte nesses 64
— x foetidum Moench. ......... 64, 66, 67
— X inapertum auct. .........0..cceeeeeeeeee 66
— inapertum (L.) Mill. .......... 62, 64, 67
— incanum foetens, flore minore
INI OTI SS eee ose ee eee Keke eee oe 66
— longipapposum Fisch. and Mey. .......
nce eek PEA Rus Me CIe 62, 65
— orientale Mill. ............000cccccccceeseeeeee 70
== pungens WAI 2... .cscc<:2.eessccese00eses: 68
— radiatum Lam. ...........:000..cccc0000000- 62
— — var. a. caule stricto etc. Ldb. .... 62
— — var. a. macranthum C. Koch. .... 62
— — var. b. caule subsquarroso etc.
{BiG [oR Ae ene orn ee Ey ae 63
— — var. c. micranthum C. Koch. ... 63
— squarrosum Boiss. ............ 62, 63, 64
— — Q. pictum BOISS. .............:cc000 63
— — B. unicolor Boiss. ..............:0-++ 63
Xerarctium JuzZ., SCct. ...............0000088 180
Xeroloma Cass. ..........s000000000+ - 59, 60, 66
Xeroloma (Cass.) DC., sect. ............... 66
Xiphacanthae Winkl., sect. ....... 186, 277
Xiphiolepides Bge., sect............ 276, 278

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