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FLORA OF THE USSR

Volume XXVI

COMPOSITAE

Tribes Anthemideae, Senecioneae, and Calenduleae

7 et 'e a at we a i 2s ¥
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FLORA OF THE USSR

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

VOLUME XXVI
COMPOSITAE

Tribes Anthemideae, Senecioneae, and Calenduleae

Volume Editors

B.K. SCHISCHKIN AND E.G. BOBROV

General Scientific Editors
Stanwyn G. Shetler
Ellen Unumb

Translated from Russian

Smithsonian Institution Libraries
Washington, D.C.
2000

SMIN B87-105

Flora SSSR, Tom XXVI
Akademiya Nauk SSSR Publishers,
Moscow-Leningrad, 1961

© 2000 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. -26: V.L. Komarov Botanical
Institute. Academy of Sciences of the USSR.

Reprint. Originally published: Leningrad:
Izdatel’stvo Akademi nauk SSSR, 1934—

Vols. have imprint: Washington, D.C. : Smithsonian
Institution Libraries and the National Science
Foundation.

Includes bibliographies and indexes.

1. Botany—Soviet Union—Classification.
2. Plants—Identification. I. Komarov, V.L.
(Vladimir Leontevich), 1869-1945. II. Botanicheski1
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

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

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

INSTITUTUM BOTANICUM nomine V.L. KOMAROVII ACADEMIAE SCIENTIARUM URSS

FLORA
URSS

(FLORA UNIONIS RERUMPUBLICARUM
SOCIALISTICARUM SOVIETICARUM)

XXVI

EDITIO ACADEMIAE SCIENTIARUM URSS
MOCQUA 1961 LENINGRAD

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

Contributors

K.S. Afanasiev, V.P. Botschantzev, I.T. Vassilczenko,
S.G. Gorschkova, M.M. Iljin, M.E. Kirpicznikov,
O.E. Knorring, L.A. Kuprianova, E.G. Pobedimova, P.P. Poljakov,
A.I. Pojarkova, L.A. Smoljianinova,
An.A. Fedorov, A.I. Tzvetkova, N.N. Tzvelev, B.K. Schischkin

SCIENTIFIC EDITOR’S PREFACE TO VOLUME XXVI

This is the second volume of the six (Vols. XX V—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. In this task, I have had the
expert assistance of Ellen Unumb. 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.

July 2000 STANWYN G. SHETLER
Curator of Botany Emeritus

National Museum of Natural History

Smithsonian Institution

Washington, DC 20560-0166

Assisted by Ellen Unumb
Bethesda, Maryland 20816

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IV*

PREFACE

The twenty-sixth volume includes descriptions of 53 genera and
680 numbered species based on a critical examination of materials of
four tribes of the family Compositae.

In this volume the vast majority of the species belong to the tribes
Anthemideae and Senecioneae.

The work on the Anthemideae in particular required much time
and effort because of the complexity of its classification. As a result,
about 15 new genera were established, making it possible to define
precisely many hitherto poorly delimited genera. In the process nearly
100 new species were described.

Considerable new information has also been added to the
systematics of the genera belonging to the tribe Senecioneae.

About 80 so far unpublished taxa are included in this volume in
Addenda XXV.

The species described in this volume include many economically
important plants. Most important of these are the wormwoods, many
species of which are major pasture plants in the southern zones of the
Soviet Union or have medicinal or industrial importance.

While this volume was in press the latest volumes of the floras of
Azerbaidzhan and Turkmenia were published, which contained material
on these very genera. For technical reasons, not all the illustrations
included in these Republic floras have been cited in the present volume.

Editorial Board

“Page number of the Russian original—General Editor.

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basa Ismotiba

M.E. Kirpicznikov

N.N. Tzvelev

An. A. Fedorov
K.S. Afanasiev

V.P. Botschantzev
L.I. Tzvetkova
E.G. Pobedimova

O.E. Knorring
L.A. Smoljaninova
P.P. Poljakov
L.A. Kuprianova

M.M. Iljin
S.G. Goroschkova

CONTRIBUTORS

Characteristics of the tribes and subtribes and
keys to the genera of the tribe Senecioneae

Keys to the genera of the tribe Anthemideae
and genera Chamaemelum, Santolina,
Otanthus, Argyranthemum, Chrysanthemum,
Leucanthemum, Leucanthemella,
Coleastephus, Pyrethrum, Spathipappus,
Waldheimia, Ugamia, Tricanthemis,
Xylanthemum, Cancriniella, Cancrinia,
Lepidolopsis, Tanacetum, Hemipappus,
Pseudohandelia, Dendranthema, Tridactylina,
Brachanthemum, Ajania, Hippolytia, and
Filifolium

Genus Anthemis

Genus Anacyclus and genus Achillea, except
section Ptarmica

Section Ptarmica of genus Achillea
Genus Handelia

Genera Matricaria, Microcephala,
Tripleurospermum, and Cotula

Genus Lepidolopha
Genus Centipeda
Genera Kaschgaria, Artemisia, Mausolea

Genera Tussilago, Petasites, Nardosmia, and
Homogyne

Genus Arnica

Genus Doronicum

Xil

A.I. Pojarkova Genera Erechtites, Cacalia, Syneilesis, Ligularia
B.K. Schischkin Genus Senecio

I.T. Vassilczenko Genera Calendula, and Gundelia

Addenda XX V—Diagnoses plantarum novarum in tomo XXVI Florae
URSS commemoratarum.

Plates prepared by artists E.S. Gaskevich— I-IV, VIII-XVIII; N.A.
Moiseeva—V-VII, XIX—XXXII; T.N. Voronikhina—XXXIII-XXXVI,;
T.N. Shishlova—XXXVII-XL.

CONTENTS

SCIENTIFIC EDITOR’S PREFACE TO VOLUME XXVI .

PREFACE
CONTRIBUTORS

SYSTEMATIC INDEX OF THE SPECIES OF THE
TWENTY-SIXTH VOLUME OF THE “FLORA OF
THE USSR”

TRIBE 7. Anthemideae Cass.
Subtribe 1. Anthemidinae O. Hoffm.
Genus 1513. Anthemis L.
Genus 1514. Chamaemelum Mill.
Genus 1515. Anacyclus L.
Genus 1516. Achillea L.
Genus 1517. Handelia Heimerl.
Genus 1518. Santolina L.
Genus 1519. Otanthus Hoffmgg. and Link.
Subtribe 2. Chrysantheminae O. Hoffm.

Genus 1520. Argyranthemum Webb. ex Sch. Bip.
Genus 1521. Chrysanthemum L.
Genus 1522. Leucanthemella Tzvel. nom. nov.
Genus 1523. Leucanthemum Mill.
Genus 1524. Coleostephus Cass.
Genus 1525. Matricaria L.
Genus 1526. Microcephala Pobed.
Genus 1527. Tripleurospermum Sch. Bip.
Genus 1528. Pyrethrum
Genus 1529. Spathipappus Tzvel. Gen. Nov.
Genus 1530. Waldheimia Kar. and Kir.
Genus 1531. Ugamia Pavl.

XiV

Genus
Genus
Genus
Genus
Genus
Genus
Genus
Genus
Genus
Genus

Genus
Genus
Genus
Genus
Genus
Genus
Genus
Genus
Genus
Genus
Genus
Genus

1532.
1533.
1534.
153535.
1536:
1537;
1538.
1539.
1540.
1541.

1542.
1543.
1544.
1545.
1546.
1547.
1548.
1549.
1550.
1551.
1552;
15533

Trichanthemis Rgl. and Schmalh.
Xylanthemum Tzvel. Gen. Nov.
Lepidolopha Winkl.

Cancriniella Tzvel. Gen. Nov.
Cancrinia Kar. and Kir.
Lepidolopsis Poljak.

Tanacetum L. Emend. Tzvel.
Hemipappus C. Koch.

Pseudohandelia Tzvel. Gen. Nov.

Dendranthema (DC.) Des Moul.
Emend. Tzvel.

Tridactylina (DC.) Sch. Bip.
Brachanthemum DC.

Ajania Poljak.

Hippolytia Poljak.
Filifolium Kitamura

Cotula L.

Centipeda Lout.

Kaschgaria Poljak.
Artemisia L.

Mausolea Bge.
Turaniphytum Poljak. Gen. Nov.
Neopallasia Poljak.

TRIBE 8. Senecioneae Cass.
Subtribe Senecioninae O. Hoffm.

Genus
Genus
Genus

Genus
Genus

Genus
Genus
Genus
Genus
Genus
Genus

1554.
1533.
1556.

1557.
1558.

1559.
1560.
1561.
1562.
1563.
1564.

Tussilago L.
Petasites Mill.
Nardosmia Cass.

Homogyne Cass.
Arnica L.

Doronicum L.

Erechtites Raf.

Cacalia L.

Syneilesis Maxim.

Senecio L.

Ligularia Cass. Nom. Cons.

TRIBE 9. Calenduleae Cass.

261
274
215
249
280
301
303
344
346
348

370
373
378
391
396
399
400
402
404
600
601
604
606
608
609
610
613
623
624
636
650
651
665
666
7352
814

Genus 1565. Calendula L.

Genus 1565a. Dipterocoma Fisch. and Mey.

TRIBE 10. Arctotideae Cass.
Subtribe Gundelinae (“Gundelieae’”’) Benth.
Genus 1566. Gundelia L.
ADDENDA
INDEX ALPHABETICUS

XV

815
819
819
820
820
822
859

Vv SYSTEMATIC INDEX OF THE SPECIES OF THE TWENTY-

SIXTH VOLUME OF THE “FLORA OF THE USSR”

14371.

14380.

> > Db Db

>>> RRR RRR RRDRDR>DD

Family CLX. Compositae P.F. Gmelin

Subfam. 1. Cardouideae Kitam.

Tribe 7. Anthemideae Cass.

Subtribe 1. Anthemidinae O. Hoffm.

Genus 1513. Anthemis L.

Section 1. Rumata Fed.

. Saportana Alb.

. iberica MB.

. anahytae Woron. ex Sosn.

. carpatica Waldst. and Kit. ex Willd.

ptarmiciformis C. Koch

. anatolica Boiss.
. emiliae Sosn.
. sterilis Stev.

tranzscheliana Fed.
tempskyana Freyn and Sint.

. fruticulosa MB.

. trotzkiana Claus ex Bge.
. calcarea Sosn.

. marschalliana Willd.

. sosnovskyana Fed.

Section 2. Anthemis

. arvensis L.

. ruthenica MB.

. candidissima Willd. ex Spreng.
. grossheimii Sosn.

“Reproduced from the Russian original. Russian page numbers appear in the left-
hand margin of the text—General Editor.

VI

XVili

14390.

14400.

14410.

14420.

20. A. hirtella Winkl.

> > > Sb SD

Section 3. Cota (J. Gay ex Guss.) Rupr.

>> bh BaD RDS Se SS ee eee

. tinctoria L.
. subtinctoria Dobrocz.
. maris-nigri Fed.

saguramica Sosn.
markhotensis Fed.

. monantha Willd.

. cretacea Zefir.

. debilis Fed.

. zephyrovii Dobrocz.
. rigescens Willd.

. jailensis Zefir.

dumetorum Sosn.

. linczevskyi Fed.

dubia Stev.

. woronowii Sosn.

talyschensis Fed.
euxina Boiss.

. macroglossa Somm. and Lev.

schischkiniana Fed.
abagensis Fed.

. zyghia Woron.

melanoloma Trautv.

. altissima L.

. coelopoda Boiss.

. austriaca Jacq.

. wiedemanniana Fisch. and Mey.

Section 4. Maruta (Cass.) Boiss.

. cotula L.

. lithuanica (DC.) Trautv.

. odontostephana Boiss.

. microcephala (Schrenk) B. Fedtsch.
. deserticola Krasch. and M. Pop.

Genus 1514. Chamaemelum Mill.
nobile (L.) All.
Genus 1515.. Anacyclus L.

. cliliatus Trautv.

officinarum Hayne

38

67

68
69

Vil

14430.

14440.

14450.

13.
14.
IS:
16.
My:
18.
199
20.
aks

22:
23e
24.
2).
26.

eT,
28.
20:
30.
al,
a2.
33.
34.

—
SP Se pS PP ae ae

> > Sb D> D> >> S SSS DS DS Db

>> dS SDS Db YD

>> SSS SDS YD DD YD YD

Genus 1516. Achillea L.
Section |. Millefoliatae DC.

. nobilis L.

. neilreichii Kern.
. millefolium L.

. sudetica Opiz.

. borealis Bong.

pannonica Cheele

. setacea Waldst. and Kit.

. asiatica Serg.

. bucharica Winkl.

. distans Waldst. and Kit. ex Willd.
. stricta Schleich.

. latiloba Ldb. ex Norm.

Section 2. Filipendulinae DC.

. filipendulina Lam.

. coarctata Poir.

. ochroleuca Ehrh.

. leptophylla MB.

. glaberrima Klok.

. micrantha Willd.

. biebersteinii Afan.

. cuneatiloba Boiss. and Buhse
. micranthoides Klok.

Section 3. Santolinoideae DC.

. wilhelmsii C. Koch
. kermanica Gand.

. vermicularis Trin.
. tenuifolia Lam.

. schischkinii Sosn.

Section 4. Pitarmica (DC.) Koch

. ligulata Waldst. and Kit.

. biserrata MB.

. griseo-virens Alb.

. ptarmica L.

. acuminata (Ldb.) Sch. Bip.
. ptarmicifolia (Willd.) Rupr. ex Heimer]
. sachokiana Sosn.

. macrocephala Rupr.

X1X

Vill

XX

14460.

14470.

35. A. salicifolia Bess.

36. A. sedelmeyeriana Sosn.

37. A. cartilaginea Ldb.

38. A. septentrionalis (Serg.) Botsch.
39. A. impatiens L.

40. A. ledebouri Heimer!

41. A. alpina L.

42. A. japonica Heimer]

43. A. ptarmiccides Maxim.

44. A. camtschatica Rupr. ex Heimer]
45. A. schuri Sch. Bip.

Genus 1517. Handelia Heimerl
1. H. trichophylla (Schrenk) Heimer]
Genus 1518. Santolina L.

—

S. chamaecyparissus L.
2. S. viridis Willd.

Genus 1519. Otanthus Hoffmgg. and Link
1. O. maritimus (L.) Hoffmgg. and Link
Subtribe 2. Chrysantheminae O. Hoffm.
Genus 1520. Argyranthemum Webb ex Sch. Bip.
*A. frutescens (L.) Sch. Bip.
Genus 1521. Chrysanthemum L.
Section 1. Ismelia (Cass.) DC.
1. Ch. carinatum Schousb.
Section 2. Chrysanthemum

2. Ch. coronarium L.
3. Ch. roxburghii Cass.

Section 3. Xanthophtalmum (Sch. Bip.) Tzvel.
4. Ch. segetum L.
Genus 1522. Leucanthemella Tzvel.

1. L. serotina (L.) Tzvel.
2. L. linearis (Matsum.) Tzvel.

*Cultivated species.—General Editor.

112
113
114
by ie)
116
117
119
120
121
122
Wee,

125

126
127

128

130

hele.

134
15

136

139
139

14480.

[

14490. 3.

—

As See

all a ea =

M.
M.
M.

Genus 1527. Tripleurospermum Sch. Bip.

as es es es es

Pinedo oo

Genus 1523. Leucanthemum Mill.

. rotundifolium (W. and K.) DC.
. vulgare Lam.
. subalpinum (Schur) Tzvel.

Genus 1524. Coleostephus Cass.

. myconis (L.) Cass.

Genus 1525. Matricaria L.
Section 1. Eumatricaria Sch. Bip.

recutita L.
tzvelevii Pobed.

Section 2. Anactidea DC.

matricarioides (Less.) Porter ex Britton ...

aurea (L.) Boiss.
Genus 1526. Microcephala Pobed.

lamellata (Bge.) Pobed.
turcomanica (Winkl.) Pobed.
subglobosa (Krasch.) Pobed.

Subgenus 1. Chionogeton Pobed.

. caucasicum (Willd.) Hayek

. subnivale Pobed.

. grossheimii (An. Fed.) Pobed.

. szowitzii (DC.) Pobed.

. transcaucasicum (Mandden.) Pobed.
. karjaginii (Mand. and Sof.) Pobed.

Subgenus 2. Tripleurospermum

Section 1. Phaeocephala Pobed.

. ambiguum (Ldb.) Fr. and Sav.
tetragonospermum (F. Schm.) Pobed.

phaeocephalum (Rupr.) Pobed.
subpolare Pobed.

Section 2. Chlorocephala Pobed.

. elongatum (F. and M.) Bornm.
. rupestre (Somm. and Lev.) Pobed

XXi

143
143
145

147

148
149

150
[51

153
155
156

161
162
165
166
167
167

168
170
171
172

173
1g

XXil

14510.

14520.

22.

oy
asl ae) |

14.

|

yA

Section 5. Eugastrosulum (Sch. Bip.) Pobed.
T.

me] as) ash ash seh ae) ae as

. colchicum (Mand.) Pobed.
. tchichatchevii (Boiss.) Bornm.

Section 3. Tripleurospermum

. inodorum (L.) Sch. Bip.

. maritimum (L.) Koch

. limosum (Maxim.) Pobed.

. breviradiatum (Ldb.) Pobed.

Section 4. Leiospermum Sch. Bip.

. disciforme (C.A.M.) Sch. Bip.
. decipiens (Fisch. and Mey.) Bornm.
eof.

sevanense (Mand.) Pobed.

parviflorum (Willd.) Pobed.
Genus 1528. Pyrethrum

Section 1. Gymnoclines (Cass.) DC.

. macrophyllum (W. and K.) Willd.
. sorbifolium Boiss.

Section 2. Balsamita (Mill.) DC.

. balsamita (L.) Will.
. balsamitoides (Nabel.) Tzvel.
. majus (Desf.) Tzvel.

Section 3. Parthenium (Briq.) Tzvel.

. peucedanifolium (Sosn.) Manden.
. parthenifolium Willd.

. parthenium (L.) Sm.

. glanduliferum Somm. and Lev.

. grossheimii Sosn.

. sevanense Sosn.

. fruticulosum Biehl.

. kubense Grossh.

Section 4. Leptanthemum Tzvel.

. leptophyllum Stev.

Section 5. Balsamitopsis Tzvel.

. kelleri (Kryl. and Plotn.) Krasch.

174
174

175
177
L7¢
178

181
182
182

183

194
195

196
198
198

202
203
204
205
206
207
209
209

au

212

14530.

14540.

14550.

16

L7.
18.
iD.
20.
21.

22.
23.

24
25
26

af.

Section 6. Cinerariifolia (Heyw.) Tzvel.
. P. cinerariifolium. Trev.
Section 7. Pyrethrellum Tzvel.

P. punctatum (Desr.) Bordz. ex Sosn.
P. marionii Albov

P. roseum (Adam) MB.

P. coccineum (Willd.) Worosch.

P. chamaemelifolium (Somm. and Lev.)
Sosn.

P. silaifolium Stev.

P. daghestanicum (Rupr. ex Boiss.)
Rupr. ex Flerov

. P. aromaticum (Rupr. ex Boiss.) Tzvel.
. P. tricholobum Sosn. ex Manden.

. P. komarovii Sosn.

Section 8. Leucanthemopsis (Giroux) Tzvel.
P. alpinum (L.) Schrank
Section 9. Pyrethrellum Tzvel.

. kotschyi Boiss.
. ordubadense Manden.
. heldreichianum Fenzl ex Tchih.

yD

Section 10. Pyrethrum

. corymbosum (L.) Willd.
. Clusii Fisch. ex Rchb.

. corymbiforme Tzvel.

. poteriifolium Ldb.

oe fe

Section 11. Richteriopsis Tzvel.

. sericeum (Adam) MB.
. oxylepis (Bordz.) Tzvel.
. aucherianum DC.

a oe i >

Section 12. Brachyglossa Tzvel.

. hissaricum Krasch.

. galae M. Pop.

. mikeschinii Tzvel.

. semenovii (Herd.) Winkl. ex O. and
. Fedtsch.

wD Vv vy

XXili

213

215
216
2t7
218
219

220
222.

222
223
224

228

229
230
231

232
234
234
235

237
238
239

241
242
243
243

XXIV

14560.

14570.

42.

==

i.
Genus 1532. Trichanthemis Rgl. and Schmalh.

1.

Section 13. Trichanthemopsis Tzvel.
P. tianschanicum Krasch.

Section 14. Richteria (Kar. and Kir.) Tzvel.

. P. alatavicum (Herd.) O. and B. Fedtsch.
. P. krylovianum Krasch.

. P. abrotanifolium Bdg. ex Ldb.

. P. pulchellum Turcz.

. P. lanuginosum (Sch. Bip. and Herd.)

Tzvel.

. P. pulchrum Ldb.

. P. karelinii Krasch.

. P. songaricum Tzvel.

. P. pyrethroides (Kar. and Kir.) B.

Fedtsch. ex Krasch.
neglectum Tzvel.
transiliense (Herd.) Rgl. and Schmalh.

djilgense (Franch.) Tzvel.
leontopodium (Winkl.) Tzvel.

P:

P. a
. P. arassanicum (Winkl.) O. and B. Fedtsch. ...

iP

|e

Genus 1529. Spathipappus Tzvel.

. 8. griffithii (Clarke) Tzvel.

Genus 1530. Waldheimia Kar. and Kir.
Section 1. Allardia (Decne.) Tzvel.

. tomentosa (Decne.) Rgl.
. Stoliczkae (Clarke) Ostenf.
. transalaica Tzvel.

Section 2. Waldheimia

W. tridactylites Kar. and Kir.
W. glabra (Decne.) Rgl.

Genus 1531. Ugamia Pavl.

U. angrenica (Krasch.) Tzvel.

Section 1. Ligulata Krasch.
T. aulieatensis (B. Fedtsch.) Krasch.

247

249
250
250
Zo
252

293
254
255
256

257
258
259
260
260

262

267
268
269

269
270

2t2

277

XI

14580.

14590.

14600.

Section 2. Trichanthemis

2. T. karataviensis Rgl. and Schmalh.

Section 3. Subbulbosa Tzvel.

3. T. radiata Krasch. and Vved.

Section 4. Leianthus (Novopokr. and Sidor.) Tzvel.

4. T. aurea Krasch.
5. T. paradoxa (Winkl.) Tzvel.

Se ee

oN AKRWN
aaaaaana

>< >< ><

See Sm

as

Q

Section 5. Pyrethroides Tzvel.

. butkovii Koval.

litwinowii (Krasch.) Tzvel.

Genus 1533. Xylanthemum Tzvel.

. pamiricum (Hoffm.) Tzvel.
. fischerae (Aitch. and Hemsl.) Tzvel.
. rupestre (M. Pop. ex Nevski) Tzvel.

Genus 1534. Lepidolopha Winkl.

. komarowii Winkl.

. filifolia Pavl.

. mogoltavica Krasch.

. nuratavica Krasch.

. karatavica Pavl.

. fedtschenkoana Knorr.

Genus 1535. Cancriniella Tzvel.
krascheninnikovii (Rubtz.) Tzvel.
Genus 1536. Cancrinia Kar. and Kir.

Section 1. Polychrysum Tzvel.

. tadshikorum (Kudr.) Tzvel.

Section 2. Tanacetopsis Tzvel.

. goloskokovii (Poljak.) Tzvel.

. subsimilis (Rech. f.) Tzvel.
mucronata (Rgl. and Schmalh.) Tzvel.
. setacea (Rgl. and Schmalh.) Tzvel.

ferganensis (Koval.) Tzvel.
submarginata (Koval.) Tzvel.

. pjataevae (Koval.) Tzvel.

XXV

278

280

281
281

282
283

286
287
287

289
289
290
290
291
291

293

299

301
302
302
304
304
305
306

xii

XXVi

14610.

14620.

iif
18.

ble a adc a

C. karatavica Tzvel.

C. pamiralaica (Koval.) Tzvel.

C. urgutensis (M. Pop.) Tzvel.

C. botschantzevii (Koval.) Tzvel.

C. nevskii Tzvel.

C. paropamisica (Krasch.) Tzvel.

C. santoana (Krasch., M. Pop. and Vved.)
Poljak.

Section 3. Matricarioides Tzvel.

. C. discoidea (Ldb.) Poljak.

Section 4. Cancrinia

C. chrysocephala Kar. and Kir.
C. tianschanica (Krasch.) Tzvel.

Genus 1537. Lepidolopsis Poljak.

. L. turkestanica (Rgl. and Schmalh.) Poljak. ..

Genus 1538. Tanacetum L.
Section 1. Omalotes (DC.) Tzvel.
T. bipinnatum (L.) Sch. Bip.
Section 2. Tanacetum

T. vulgare L.

T. boreale Fisch. ex DC.

T. pseudoachillea Winkl.

T. abrotanifolium (L.) Druce

Section 3. Xanthoglossa (DC.) Sch. Bip.

. myriophyllum Willd.

. duderanum (Boiss.) Tzvel.

. heterophyllum Boiss.
tamrutense (Sosn.) Sosn.

. tabrisianum (Boiss.) Sosn. and Takht.
canescens DC.

. tanacetoides (DC.) Tzvel.
crassipes (Stschegl.) Tzvel.
turlanicum (Pavl.) Tzvel.

. scopulorum (Krasch.) Tzvel.
. Saxicolum (Krasch.) Tzvel.

. santolina Winkl.

. karelinii Tzvel.

SSS SSS ee ae ea

306
307
308
309
309
310
ei

373

314
3135

316

324

326
65 |
328
329

331
332
333
334
335
336
337
338
341
342
342
343
344

14630.

14640.

XIV 14650.

19.
20.

a3.
34.

i He Mie ae Be aig ae ms rg me te

ae ie

. kittaryanum (C.A.M.) Tzvel.
. uralense (Krasch.) Tzvel.
. sclerophyllum (Krasch.) Tzvel.

odessanum (Klok.) Tzvel.
millefolium (L.) Tzvel.

paczoskii (Zefir.) Tzvel.

ulutavicum Tzvel.

achilleifolium (MB.) Sch. Bip.
chiliophyllum (F. and M.) Sch. Bip.
longipedunculatum (Sosn.) Tzvel.
oligocephalum (DC.) Sch. Bip.
tenuissimum (Trautv.) Grossh.
uniflorum (F. and M.) Sch. Bip.

. akinfievii (Alex.) Tzvel.

Section 4. Asterotricha Tzvel.

. turcomanicum (Krasch.) Tzvel.
. walteri (Winkl.) Tzvel.

Genus 1539. Hemipappus C. Koch.
canus C. Koch.

Genus 1540. Pseudohandelia Tzvel.

1. P. umbellifera (Boiss.) Tzvel.

11.
12.

SBR ea PRS es

Genus 1541. Dendranthema (DC.) Des Moul.

Soo

VoOoPpooOSE oo ho

Section 1. Dendranthema

. sinuatum (Ldb.) Tzvel.
. indicum (L.) Des Moul.
. morifolium (Ramat.) Tzvel

erubescens (Stapf) Tzvel.
naktongense (Nakai) Tzvel.
zawadskii (Herb.) Tzvel.
mongolicum (Ling) Tzvel.
maximowiczii (Kom.) Tzvel.

. sichotense Tzvel.
. weyrichii (Maxim.) Tzvel.
. littorale (Maekawa) Tzvel.

Section 2. Arctanthemum Tzvel.

. kurilense Tzvel.
. arcticum (L.) Tzvel.
. hultenii (A. and D. Léve) Tzvel.

XXVIi

345
346
347
348
348
349
350
351
332,
354
354
Kets)
356
357

358
359

360

363

370
371
373
374
375
376
378
379
380
383
384

385
386
387

XV

XXVill

14660.

14670.

14680.

ig.

i ci

Saree aS ee
> Pd DSS YS DS BD

fe ee

D.

Section 3. Haplophylla Tzvel.

integrifolium (Richards.) Tzvel.

Genus 1542. Tridactylina (DC.) Sch. Bip.

a

Baws

mrrot

kirilowii (Turcz.) Sch. Bip.
Genus 1543. Brachanthemum DC.

Section 1. Dendrathemopsis Tzvel.

. baranovii (Krasch. and Poljak.) Krasch. ...

Section 2. Brachanthemum

. titovii Krasch.

. kirghisorum Krasch.

. fruticulosum (Ldb.) DC.
. kasakhorum Krasch.

Genus 1544. Ajania Poljak.

. manshurica Poljak.

. pallasiana (Fisch. ex Bess.) Poljak.

. fastigiata (Winkl.) Poljak.

. fruticulosa (Ldb.) Poljak.

. gracilis (Hook. f. and Thoms.) Poljak.
. kokanica (Krasch.) Tzvel. ,
. scharnhorstii (Rgl. and Schmalh.) Tzvel. ...
. trilobata Poljak.

. tibetica (Hook. f. and Thoms.) Tzvel.

Genus 1545. Hippolytia Poljak.

. megacephala (Rupr.) Poljak.

. herderi (Rgl. and Schmalh.) Poljak.
. darvasica (Winkl.) Poljak.

. Schugnanica (Winkl.) Poljak.

Genus 1546. Filifolium Kitamura

. Sibiricum (L.) Kitamura

Genus 1547. Cotula L.

. coronopifolia L.

Genus 1548. Centipeda Lour.

. orbicularis Lour.

388

390

394

395
396
396
397

401
402
405
406
407
408
409
409
410

413
414
415
416

417

421

422

14690.

14700.

14710.

—

Genus 1549. Kaschgaria Poljak.
K. brachanthemoides (Winkl.) Poljak.

2. K. komarovii (Krasch.) Poljak.

16.

aI.

en
Ce ee ees ae oS) See
pa RSP SPRSPPP PPR PH PPP YS

NOK eK
dial s

> > dS DS Sd Dd DS D>

Genus 1550. Artemisia L.
Subgenus 1. Artemisia
Section 1. Artemisia

. vulgaris L.

. opulenta Pamp.

. unalaskensis Rydb.
. tilesii Ldb.

. mongolica Fisch. ex Nakai
. silvatica Maxim.

. ussuriensis Poljak.

. Stolonifera (Maxim.) Kom.
. koidzumii Nakai

. Saitoana Kitamura

. komarovii Poljak.

. integrifolia L.

. subulata Nakai

. rubripes Nakai

argyi Lev. and Van.

. montana Pamp.

. lavandulifolia DC.

. umbrosa Turcz. and DC.

. selengensis Turcz. ex Besser

Section 2. Stellerianum (Rydb.) Poljak.

A. stelleriana Bess.
Section 3. Abrotanum Bess.

. abrotanum L.

. chamaemelifolia Vill.

. pontica L.

. macrantha Ldb.

. adamsii Bess.

. gmelinii Web. ex Stechm.

. santolinifolia Turcz. ex Bess.
. freyniana (Pamp.) Krasch.

. medioxima Krasch. ex Poljak.

. leucophylla Turcz. ex C.B. Clarke

XXIX

424
424

438
439
440
443
443
444
445
446
446
447
448
449
449
450
450
451
452
453
453
454

455

456
458
462
462
463
464
465
467
468

XXX

14720.

14730.

14740.

14750.

eS ee ee eee re ree ae ae ae

pre errr P PP Perr PerP PPP P PRB BDBDBD

. tanacetifolia L.
. maximovicziana Krasch. ex Poljak.
. remotiloba Krasch. ex Poljak.

armeniaca Lam.

. latifolia Ldb.

. laciniata Willd.

. phaeolepis Krasch.

. laciniatiformis Kom.

. macrorhiza Turcz.

. arctica Less.

. punctigera Krasch. ex Poljak.
. norvegica Fries.

. globularis Bess.

. glomerata Ldb.

. furcata MB.

. insulana Krasch.

. Senjavinensis Bess.

. leontopodioides Fisch. ex Bess.
. lagopus Fisch. ex Bess.
. triniana Bess.

. keiskeana Miquel.

. tournefortiana Rchb.

. annua L.

. palustris L.

. aurata Kom.

Section 4. Absinthium DC.

. caucasica Willd.

. Splendens Willd.

. frigida Willd.

. austriaca Jacq.

. caespitosa Ldb.

. cuspidata Krasch.

. schmidtiana Maxim.

. sericea Web.

. aschurbajevii Winkl.

. lagocephala (Bess.) DC.
. rutifolia Steph. ex Spreng.
. persica Boiss.

. hololeuca MB. ex Bess.
. rupestris L.

. obtusiloba Ldb.

. subviscosa Turcz.

468
469
470
471
472
473
474
477
478
479
480
480
481
482
483
484
485
485
486
487
487
488
489
491
492

492
493
494
498
499
500
501
501
503
504
505
506
507
508
509
510

XVII

14760.

14770.

14780.

14790.

a a ee

Section 5. Artanacetum (Rzazade) Poljak.

. A.

- >i P> > bb Ra DR D EDD DDD DDR RE EB ED

. martjonovii Krasch.

. altaiensis Krasch.

. absinthium L.

. sieversiana Willd.

. macrocephala Jacq.

. kulbadica Boiss. and Buhse
. samoiedorum Pampan.

. succulenta Ldb.

. Jacutica Drob.

. anethifolia Web.

incana (L.) Druce

Subgenus 2. Dracunculus Bess.

. dracunculus L.
. pamirica Winkl.
. dracunculiformis Krasch.

daghestanica Krasch. and Por.

. glauca Pall.

. dimoana M. Pop.

. kelleri Krasch.

. trautvetteriana Bess.

. salsoloides Willd.

. halodendron Turcz. ex Bess.
. quinqueloba Trautv.

. arenaria DC.

. albicerata Krasch.

. saposhnikovii Krasch. ex Poljak.
. songarica Schrenk

. lipskyi Poljak.

kuschakewiczi Winkl.

. japonica Thunb.
. littoricola Kitam.

desertorum Spreng.

. macilenta (Maxim.) Krasch.
. capillaris Thunb.

commutata Bess.

. campestris L.

. tomentella Trautv.

. bargusinensis Spreng.
. ledebouriana Bess.

. pannosa Krasch.

XXX1

=} Fe
514
515
517
518
519
520
524
522
522

524

529
534
534
535
535
536
537
537
538
539
540
540
542
542
543
544
544
545
546
549
550
550
551
552
554
555
355
556

XVIill

XXXil

14800.

14810.

14820.

14830.

a
be.
bt3.
114.
11S.
116.

Ly.
118.
119.
120.
ea
122.
123:
124.
125.
126.
127.
128.
129.
130.
131.
132.
133:
134.
135.
136.
137.
138.
139.
140.
141.
142.
143.
144.
145.
146.

147.
148.

> > Sb Db YS Db

polis: De I Ss be pi ih 5 is i yee ee a Bs

. borealis Pall.

. henriettae Krasch.

. limosa Koidz.

. pycnorhiza Ldb.

. scoparia W. and K.
. demissa Krasch.

Subgenus 3. Seriphidium (Bess.) Rouy

. maritima L.

. monogyna Waldst. and Kit.
. szowitziana (Bess.) Grossh.
. fragrans Willd.

. taurica Willd.

. lercheana Web. ex Stechm.

nitrosa Web. ex Stechm.
scopaeformis Ldb.

. halophila Krasch.

. schrenkiana Ldb.

. compacta Fisch. ex DC.

. rhodantha Rupr.

. prolixa Krasch. ex Poljak.

. dubjanskyana Krasch. ex Poljak.
. knorringiana Krasch.

. korovinii Poljak.

. pauciflora Web.

gracilescens Krasch. and Iljin

. terrae-albae Krasch.
. lessingiana Bess.

. aralensis Krasch.

. camelorum Krasch. if
. sublessingiana (Kell.) Krasch. ex Poljak. ...
. transiliensis Poljak.

. tenuisecta Nevski

. kaschgarica Krasch.

. amoena Poljak.

. fedtschenkoana Krasch.
. cina Berg. ex Poljak.
A.

ciniformis Krasch. and M. Pop.

ex Poljak.

A.
A.

glanduligera Krasch. ex Poljak.
ferganensis Krasch. ex Poljak.

527
558
J597
539
560
562

573
574
573
576
578
579
580
581
581
582
585
586
587
587
588
589
589
591
592
594
594
595
596
oF
598
599
600
600
601
603

604
607

XIX

14840.

14850.

14860.

>>> >RRDBDDBD DDD

. A. porrecta Krasch. ex Poljak.

. A. serotina Bge.

. A. valida Krasch. ex Poljak.

. A. gypsacea Krasch., M. Pop. and

Lincz. ex Poljak.

. A. namanganica Poljak.

. A. turanica Krasch.

. A. kemrudica Krasch.

. A. scotina Nevski.

. A. kochiiformis Krasch. and Lincz.

ex Poljak.

. A. sogdiana Bge.

. A. stenocephala Krasch. ex Poljak.

. A. kopetdaghensis Krasch. ex Poljak.
. A. balchanorum Krasch.

turcomanica Gndg.

. prasina Krasch. ex Poljak.
specigera C. Koch.

sieberi Bess.

. lehmanniana Bge.

vachanica Krasch. ex Poljak.

. baldshuanica Krasch. and Zapr.
santolina Schrenk

. lobulifolia Boiss.

. deserti Krasch.

Section 2. Junceum Poljak.

. Juncea Kar. and Kir.
. leucodes Schrenk

Genus 1551. Mausolea Bge.

> >

. M. eriocarpa (Bge.) Poljak.

Genus 1552. Turaniphytum Poljak.

. T. eranthemum (Bge.) Poljak.
. T. kopetdaghense Poljak.

Genus 1553. Neopallasia Poljak.

. N. pectinata (Pall.) Poljak.

. badhysi Krasch. and Lincz. ex Poljak.

XXXill

607
608
610
610

611
612
614
614
615

616
616
617
617
618
619
620
620
621
622
623
624
625
625
626

629
630

631

635
636

637

XXXIV

14870.

14880.

Tribe 8. Senecioneae Cass.
Subtribe Senecioniae O. Hoffm.
Genus 1554. Tussilago L.
1. T. farfara L.
Genus 1555. Petasites Mill.
1. P. spurius (Retz.) Rchb.

2. P. hybridus (L.) Gaertn., Mey and Scherb.

3. P. albus (L.) Gaertn.
Genus 1556. Nardosmia Cass.
Subgenus 1. Nardosmia
Section 1. Nardosmia

. N. saxatilis Turcz.

. N. fominii Bordz.

. N. angulosa Cass.

. N. frigida (L.) Hook.

N. laevigata (Willd.) DC.

AP WN Ee

Section Brevilobae Kuprian.

6. N. palmata (Ait.) Hook.
7. N. japonica S. and Z.

Subgenus 2. Endocellion (Turcz. ex Herd.) Kuprian.

8. N. gmelini Turcz. ex DC.
9. N. glacialis Ldb.

Genus 1557. Homogyne Cass.
1. HA. alpina (L.) Cass.

Genus 1558. Arnica L.

Subgenus 1. Arctica Maguire

. alpina (L.) Olin and Ladau
. iljinii (Maguire) Ijin
. intermedia Turcz.
. frigida C.A.M. ex Iljin

PWN re
> > > >

Subgenus 2. Montana Maguire

5. A. montana L.

641

643
643
644

647
648
648
649
650

651
652

653
654

655

657
658
659
660

663

14890.

14900.

_
> > >

ig fae tm
SSsss

Pee Pe

= AANAAANAA iy SSesyyy

NHN

Subgenus 3. Andropurpurea Maguire

. lessingii Green
. unalaschcensis Less.
. sachalinensis (Rgl.) A. Gray

Genus 1559. Doronicum L.
Section 1. Aronicum Neck.

altaicum Pall.
schischkinii Serg.
bargusinense Serg.
clusii (All.) Tausch.

carpaticum (Griseb. and Schenk) Nym.

Section 2. Pardalianches Tausch

austriacum Jacq.

orientale Hoffm.

macrophyllum Fisch.
pardalianches L.

longifolium Griseb. and Schenk
oblongifolium DC.
turkestanicum Cavill.

Genus 1560. Erechtites Raf.

. valerianifolia (Wolf.) DC.

Genus 1561. Cacalia L.

. hastata L.

. robusta Tolm.

. komaroviana (Pojark.) Pojark.
. praetermissa (Pojark.) Pojark.
. auriculata DC.

. kamtschatica (Maxim.) Kudo

Genus 1562. Syneilesis Maxim.

. aconitifolia Maxim.

Genus 1563. Senecio L.

Section 1. Jacobaea DC.

. erucifolius L.
. grandidentatus Ldb.
. korshinskyi Krasch.

XXXV

665
666
668

672
673
673
674
675

676
677
678
679
680
681
681

683

687
689
691
692
694
695

698

710
pe
712

XXXVI

14910.

14920.

14930.

14940.

18.

ANNNHANAN

ANNNHANN

ANNNNURNURNUNNUNUANURNUANNNANN

. jacobaea L.

. ambraceus Turcz. ex. DC.

. argunensis Turcz.

. saposhnikovii Krasch.

. erraticus Bertol.

. borysthenicus Andrz. ex DC.
. schischkinianus Sof.

. ferganensis Schischk.

Section 2. Incani DC.

. cineraria DC.

. lorentii Hochst.

. lipskyi Lom.

. massagetovii Schischk.
. pseudoarnica Less.

. kubensis Grossh.

. pandurifolius C. Koch.

Section 3. Quadridentati Boiss.
. taraxacifolius (MB.) DC.
Section 4. Crociserides DC.

. subalpinus C. Koch.

. buschianus Sosn.

. pseudoorientalis Schischk.

. kolenatianus C.A.M.

. olgae Rgl. and Schmalh.

. franchetii Winkl.

. paulsenii O. Hoffm. ex Paulsen
. macrophyllus MB.

racemosus (MB.) DC.

. thyrsophorus C. Koch.
. paucifolius Gmel.

schvetzovii Korsch.
umbrosus Waldst. and Kit.

. racemulifer Pavl.
. paludosus L.
. tataricus Less.

Section 5. Oliganthi Boiss.

. pojarkovae Schischk.
. rhombifolius (Willd.) Sch. Bip.
platyphylloides Somm. and Lev.

715
715
716
717
pally
718
719
720

720
724
Tae
122
723
724
724

7125

726
726
Tau
728
728
729
730
730
731
432
133
733
734
735
735
736

739
739
740

XXII

14950.

14960.

38.
39.

40.
41.
42.
43.
44.
45.
46.

47.
48.

49.

50.
Al:

a2.
y3:

54

3.

NANNHANRANAN

S.
S.

S.
S.

Section 11. Karelinoidei O. and B. Fedtsch.
+, SB.

ANRNNHNHANNHAN

. paucilobus DC.
. othonnae MB.

Section 6. Pseudo-oliganthi Sof.

. nemorensis L.

. iljinii Schischk.

. propinquus Schischk.

. lapsanoides DC.

. quinqueligulatus Winkl.
. fuchsii K. Gmel.

. fluviatilis Wallr.

Section 7. Extremiorientales Schischk.

. cannabifolius Less.
. litvinovii Schischk.

Section 8. Carniolici Schischk.

. carniolicus Willd.

Section 9. Montani Schischk.

tianschanicus Rgl. and Schmalh.
renardii Wink.

Section 10. Scaposi (Hoffm.) Schischk.

resedifolius Less.
carpathicus Herb.

karelinioides Winkl.

Section 12. Eriopappus Dumort.

. arcticus Rupr.

Section 13. Tephroseris (Rchb.) DC.

. campester (Retz.) DC.
. czernjaevii Mind.
. jailicola Juz.

amurensis Schischk.

. succisifolius Kom.
. lenensis Schischk.
. cladobotrys Ldb.

subfloccosus Schischk.

. papposus (Rchb.) Less.

XXXVI

741
741

742
743
743
744
744
745
745

747
747

748

749
749

750
750

Fi |

752

753
754
fist]
757
758
758
759
760
760

XXII

XXXVili

14970.

14980.

14990.

15000.

ANNANANNNNANRNANNDANUUANNANNUANANHAN

ANNANNNANNUUNN

. kawakamii Makino
. asiaticus Schischk.
. sukaczevii Schischk.

aquilonaris Schischk.

. igoschinae Schischk.

sichotensis Kom.

. veresczaginii Schischk.
. rivularis (Waldst. and Kit.) DC.

hieraciiformis Kom.
subscaposus Kom.

. porphyranthus Schischk.

flammeus Turcz. ex DC.
besserianus Mind.

. capitatus (Wahlb.) Steud.
. caucasigenus Schischk.

pyroglossus Kar. and Kir.

. karjaginii Sof.
atripurpureus (Ldb.) B. Fedtsch.

tichomirovii Schischk.

. pricei Simps.

turczaninovii DC.

tundricola Tolm.

subfrigidus Kom.
pseudoaurantiacus Kom.
sumneviczii Schischk. and Serg.
reverdattoi Sobolevsk.
jJacuticus Schischk.

Section 14. Obejacae DC.

viscosus L.

calvertii Boiss.

vulgaris L.

dubius Ldb.

sylvaticus L.
krascheninnikovii Schischk.
vernalis Waldst. and Kit.
sosnovskyi Sof.
polycephalus Ldb.
subdentatus Ldb.
noeanus Rupr.

761
762
762
763
763
764
764
765
765
766
766
767
768
768
769
770
770
Va
TZ
TT3
Tt
776
TTT
TB
777
778
17g

173
780
780
781
782
783
783
784
785
786
787

15010.

15020.

15030.

20.

ae
22.
24.
24.
ae

Section 1. Corymbosae (Fr.) Hand.-Mazz.

a
BONE:
a

SS oe &

ait Sail Salll Salll Salll coll soll coll oll

Genus 1564. Ligularia Cass.

Subgenus 1. Ligularia

hodgsonii Hook.
trichocephala Pojark.
calthifolia Maxim.

Section 2. Ligularia

. L. fischeri (Ldb.) Turcz.

. splendens (Lévl. and Vaniot) Nakai
. sachalinensis Nakai

. Sibirica (L.) Cass.

. lydiae Minder.

abakanica Pojark.
sichotensis Pojark.
subsagittata Pojark.

. longipes Pojark.
. arctica Pojark.
. bucovinensis Nakai

Section 3. Stenostegia Pojark.

Subsection 1. Oligocephalae Pojark.

. robusta (Ldb.) DC.

. kareliniana Stschegl.

. narynensis (Winkl.) O. and B. Fedtsch.
. karataviensis (Lipsch.) Pojark.

. Schischkinii Rubtz.

Subsection 2. Racemiferae Pojark.

. jaluensis Kom.

Subsection 3. Paniculatae Pojark.

. thyrsoidea (Ldb.) DC.
. songarica (Fisch.) Ling.

knorringiana Pojark.
thomsonii (Clarke) Pojark.
altissima Pojark.

XXXiX

798
800
800

802
803
805
807
808
810
811
812
816
817
818

820
821
822
824
825

826

827
831
832
834
835

XXIV

xl

15040.

15050.

Section 4. Senecillis (Gaertn.) Kitam.
Subsection 1. Sericochaetae Pojark.

L. macrophylla (Ldb.) DC.
L. heterophylla Rupr.
28. L. talassica Pojark.
L. alpigena Pojark.
L. altaica DC.

Subsection 2. Senecillis (Gaertn.) Pojark.

31. L. glauca (L.) O. Hoffm.
32. L. carpatica (Schott, Nym. Kotschy)
Pojark.

Subsection 3. Stereochaetae Pojark.

33. L. mongolica (Turcz.) DC.
34. L. schmidtii (Maxim.) Makino

Section 5. Glossophyllum Pojark.

35. L. pavlovii (Lipsch.) Cretz.
36. L. renifolia (C.A.M.) DC.
37. L. caucasica (MB.) G. Don.
38. L. correvoniana (Alb.) Pojark.
Tribe. 9. Calenduleae Cass.
Genus 1565. Calendula L.
1. C. persica C.A.M.
2. C. gracilis DC.
3. C. arvensis L.
4. C. karakalensis Vass.
* C. officinalis L. ...

Tribe 10. Arctotideae Cass.

Subtribe Gundelinae (““Gundelieae’”’) Benth.

Genus 1566. Gundelia L.

1. G. tournefortii L.

837
838
840
841
843

847
848

849
850

852
853
855
856

858
859
859
860
860

862

]

Tribe 7. ANTHEMIDEAE Cass. in Bull. Soc. Philom. Paris (1815)
173 and in Dict. Sc. Nat. II, Suppl. (1816) 73, XX (1821) 372, XXIXx
(1823) 176, L (1827) 497, LX (1830) 578; Benth. in Benth. and Hook.
fil. Gen. Pl. II (1873) 203; O. Hoffm. in Pflanzenfam. IV, 5 (1889-—
1894) 267.—Ordo Elichryseae Link, Handb. I (1829) 712 pro min. p.
and ordo Anthemideae Link, ibid. 752, pp.—Trib. Senecionideae
subtrib. Chrysanthemeae and subtrib. Artemisieae Less. Synops. Com.
(1832) 247, 257.—Trib. Helianthées sous-série Anthémidées Baill. Hist.
des Plantes, VIII (1882) 71.—Fam. Anthemidaceae Bessey in Ann.
Miss. Gard. 2 (1915) 164.—Capitula homogamous or heterogamous,
with distinct ray and disk florets, or pseudoligulate ray florets absent.
Receptacle with scales or hairy, less often glabrous. Anthers without
or (rarely) with barely visible basal appendages. Style branches in
bisexual florets usually truncate. Corolla mostly yellow, less often white
or reddish; corolla of outer florets (ray florets) either similar to disk
florets in color (capitula homochromous) or white, pink, purple and
other shades (capitula heterochromous). Leaves almost always alternate,
often incised.

Herbs, semishrubs, and shrubs, usually strongly fragrant. The largest
number of genera of the tribe is found in South Africa (especially in
the Cape Region) and in the Mediterranean Region. Moreover,
Anthemideae is widely distributed throughout Europe and a large part
of Asia; a few genera are represented in America, Australia, and New
Zealand.

KEY TO GENERA OF TRIBE ANTHEMIDEAE*

|. Receptacle covered, with scaly Gracts fo.c.......cc.c5ec-scesescngeoepeneoosss es
+ Receptacle lacking scaly bracts, glabrous or hairy, very rarely bracts
present in outermost florets of capituluM ............ceeeseeeeeteeeeeeee 9.

2. All florets in capitulum tubular; their corollas highly swollen in
lower part and prolonged along both sides of ovary (and achene) as
2 long spurs appressed to and persisting with achene. Whitish-
lanate plants of coastal sands of AbkhaZia ..............cc:cccscceseeees
spe ey Dongen eh. “VN 1519. Otanthus Hoffmgg. and Link
+ Corolla basally lacking long spurs, not persisting with achene .... 3.
Capitula smaller, in terminal clusters on main stem and branches,
in lax or dense corymbose inflorescence (solitary only in very weak
specimens); peduncles always lacking true leaves, but often with

ie

*Compiled by N.N. Tzvelev.

small sulending leaves involucre up to 8 mm, less often up to 10
mm in dia; limb of ligulate florets (if present) broadly ovate to
AlMOSt SEMICITCUIAL co... cyciens apex ca-p op nas Mes openpan< anes pan cao ys ce 4.

Capitula larger, solitary terminal on main stem and leafy branches,

but often numerous on plant; involucre (5)6 —20(25) mm in dia;

limb of ligulate florets (if present) broadly ovate to linear .......... oi
Capitula heterogamous; ray florets pistillate, almost always ligulate,
very rarely tubular (but then also pistillate)....... 1516. Achillea L.*
Capitula homogamous; all florets tubular, bisexual. Soviet Central
PASTA cos sagapsnaeroncouane ie hence tana ee 1517. Handelia Heimerl.
Capitula always without ligulate florets, homogamous, lamina linear
and narrow-linear, pinnatisect or pinnatipartite into many, very short,
obtuse or subobtuse lateral lobes, as if arranged in whorl on leaf
axis. Ornamental plants sometimes escaping to the wild ...........
oda dapbacoas tec eeeign cman neice ae Caseres eae eae ere 1518. Santolina L.
Capitula usually heterogamous, with ligulate florets; very rarely

with tubular florets only, but then leaves of different shape ........ 6.
Achenes, at least in ray florets, highly compressed, dorsally with 2
almost winged. jprominent laveralglWs cane ae ce ee Te

Achenes of all florets with or without 3—5 slightly raised nbs .... 8.
Perennials; achenes lacking apical corona. Carpathians ...........
Ener eet Se 1516. Achillea L. (4. schurii Sch. Bip.)
Annuals; achenes with a short apical corona or auricles .......
adn eneesbsahevawardege Regan’ ty cassatne tease ne Mente eee me 1515. Anacyclus L.
Achenes with 3 prominent ribs, lacking corona; base of corolla tube
with somewhat developed outgrowths enclosing tip of ovary;
receptacle hollow, obtusely conical. Medicinal and ornamental plants
with strong aromatic smell sometimes escaping to the wild .......
Sobslestateslthequentaht ara faiceoe weak Aouad eae ana 1514. Chamaemelum Mill.
Achenes with or without 4-5 prominent ribs; base of corolla tube
lacking downward directed outgrowths; receptacle of different shapes
sao iE Meee castor a cass aired et ate ee ce 1513. Anthemis L.
Capitula smaller (up to 8-10 mm in dia), mostly drooping, lacking
ligulate florets, on relatively short peduncles in corymbs or spikes,
in turn, usually forming paniculate inflorescence; very rarely, in
some arctic and Far Eastern species, few capitula borne in almost
capitate inflorescence (and then pollen grains not spiny) .......... 10.
Capitula, on the average, larger, with or without ligulate florets, in
corymbs, less often in corymbose-panicles, or solitary terminal on
stem and its leafy branches (in doubtful case, pollen grains always
GISEINCELY SPIAYy)) seh -accesceaceansihcsaasMbeanccccotvasteacaes cee ee 14.

*Cf. also couplet 7.

+2.

3

Achenes with somewhat truncate corona, longitudinally lobed al-
most to base; pollen grains spinulose. Russian Central Asia ................
PALA Mid, AOS 4 1537. Lepidolopsis Poljak.
Achenes lacking corona; pollen grains not spinulose ................ 11.
Semishrubs with entire or partly 3(5)-fid leaves, sparsely stellate
hairy; corolla also usually stellate hairy. Eastern Kazakhstan, Tien

Sha iach. vsti scti oa el LORRI RE 1549. Kaschgaria Poljak.*
Plants more or less pubescent or glabrous, but hairs always simple
arte SACRE, AO Rk Nee... 12:

Ray florets fertile, pistillate or bisexual; others bisexual but sterile,
compactly borne on narrow-conical receptacle; achenes strongly
compressed dorsally, in single row, in rosette at base of receptacle.
Annuals with pectinately pinnatisect leaves. Tien Shan, Tarbagatai
PS eee re eee eee ER A A LAR ae Re 1532. Neopallasia Poljak.
Plants with characters otherwise. .).a li Ben ........ 13:
Ray florets pistillate, fertile, with or without highly reduced corolla;
disk florets bisexual, but sterile, staminate (with highly reduced
pistils), with apically hairy corollas; achenes densely hispid; 45—70
cm high semishrubs. Sandy deserts of Soviet Central Asia .....
ATEN RL, Mics W Wa Wvaveritoecvevssne bead 1551. Mausolea Bge.
Ray florets pistillate with highly reduced glabrous corolla, in axils
of carinate involucral bracts; disk florets staminate, with rudimentary
pistil and much longer, 5-toothed, apically hairy corolla; capitula
small (1.0-2.5 mm in dia), often in somewhat globose clusters, in
turn, borne in spicate inflorescences .... 1552. Turaniphytum Poljak.
Disk florets fertile (with normally developed pistil) or sterile, but
in former case achenes either smooth or involucral bracts not carinate
eae ces ERA AI, Ailing. 2A 1550. Artemisia L.**

PRED AVA RS RIEL EN i ER, Reh BE U5:
Perennials with well developed root system .............::cceccceeeeeeees 25:
Receptacle less often hairy; capitula heterogamous, but only with
tubular florets; outer florets pistillate, central bisexual. Altai,
mountainsof Sovie Central tAsiasw...: elect eke...
REO. READ TA ii, 1550. Artemisia L. (A. macrocephala Jacq.)
Redepdible Siabrensiien. fad ebind aectidae cine deeds... 16.
Capitula heterogamous; ray florets pistillate, with or without very
small, indistinctly toothed corolla; disk florets bisexual, often fewer
than ray florets, corolla 4-toothed. Far East ............c:ccecseeseeeees 17;

* Cf. also couplet 38.
** Cf. also couplets 15 and 41.

17.

20.

21.

22,

23%

24.

Capitula homogamous (with only bisexual tubular disk florets) or
heterogamous, with ligulate pistillate ray florets; corolla of disk
florets 5-, less often (in species of Matricaria L.) 4-toothed ..... 18.
Florets (except innermost) and achenes with persistent papillate
processes of receptacle; corolla of disk florets with short obtuse
teeth; achenes of ray florets highly compressed, with thick lateral
ribs and dorsal papillate processes ...........:::00cee8 1547. Cotula L.
Receptacle lacking papillate processes; corolla or disk florets deeply
toothed; achenes of ray florets slightly compressed, 3—5-ribbed,
elaridularshatryosiiyy-: toc: wen iet ces. otal eka, ek 1548. Centipeda Lour.
Achenes with 2—3-highly prominent longitudinal ribs, prismatic ....
ant. JaL. Aa. 1s, Slieon. nt, moon also. oe ont leony, beeenmemeeniaton. ters 19.
Achenes with 5—10 (rarely 4) not so prominent, often inconspicuous
Pernod tirelinnall TAS L ....ssesis sen siisdenneiy cerdehennasesseedona nase eee 20.
Ligulate florets white or absent; achenes more or less coronate,
with 3 thick and obtuse ribs and 2 dark colored punctate glands in
upper part of spine ..............0 1527. Tripleurospermum Sch. Bip.*
Ligulate florets yellow, of various shades; achenes without corona,
with 2—3 almost winged sharp ribs. Omamental plants sometimes
esaaping tocthe Wald... sycccscccueceraayrenes 1521. Chrysanthemum L.
Corolla of ligulate florets yellow or yellowish-white, limb 6-16
mim long; lamina entire, trilobate or 3-fid ...........ceeeeeeeeeeeeeeeeeee Zl
Corolla of ligulate florets white, or capitula with only tubular florets;
leavesmore oriless)pimmatisecty, (6h: 15 dase ote dp ook ls be Ned beeeeetie ee ce 20,,
Achenes with rather long auriculate corona; leaves entire. Crimea
R620. there Deon la redy, wile cercclas, biaets 1524. Coleostephus Cass.
Achenes lacking true corona, but with short obtuse teeth at apex;
leavesitrilabate. or 3=fid.. Baikal Region, 5... ccncsssesg:-ccsckeusteceosuteeeeees
SMALE. cRUGON Basel Eh sary eran hl 1542. Tridactylina (DC.) Sch. Bip.
Achenes with or without inconspicuous corona, less often corona
well-developed, but then corona of tubular florets four-toothed.

Plants with strong aromatic smell .................5 1525. Matricaria L.
Achenes with well developed, usually toothed or lobed corona;
corolla of tubular florets always 5-toothed ............:cccccsccssesseseees 23.

Achenes with highly truncate, more or less lobed corona, densely
covered with whitish scaly hairs; capitula with ligulate, less often
eligulate florets. Soviet Central Asia.......1526. Microcephala Pobed.
Achenes entirely glabrous, with slightly truncate corona; large
biennial or perennial ypkanits:<!3<.001.0). Jevicacs eee eee dee 24.
Capitula solitary terminal on stem and leafy branches, lacking ligulate
florets; leaves with relatively narrow (up to 1.5—2.0 mm wide);

*Cf. also couplet 29.

25.

26.

30.

4 E

32)

2

termmal “lobes Nagakhstanrnt gy. Nk.) nuns oki ealbde.
CRON 33 1536. Cancrinia Kar. and Kir. [C. discoidea (Ldb.) Poljak.]
Capitula usually with ligulate florets in corymbs; leaves with broader

terminal lobes; ornamental plants easily escaping to the wild .............
tA chditiebelbiudeaat 1528. Pyrethrum Zinn. [P. parthenium (L.) Sm.]
Ray florets of capitula ligulate; disk florets tubular ................... 26.
All florets in capitula tubular iil ven ob sh oka iis dhesaese 36.

Achenes with pappus of numerous (25—50) setaceous scales, usually
longer than achenes. Alpine plants of Altai and Soviet Central Asia
PE ard, Ree RO. RL. 1530. Waldheimia Kar. and Kir.
Achenes lacking pappus or pappus coronate, sometimes
ldapitudinally lobed: jj... ete eit. DO Rwk.caile...... 27.
Achenes (and usually also receptacle) densely hairy all over, corona
divided into oblong or ovate lobes right to base. Soviet Central
ESTOS AT Ge 1532. Trichanthemis Rg]. and Schmalh.*
eT enes Ciitire ly SIADOUS eA onic ed bh Wades nee 28.
Achenes with 2-3-highly prominent ribs, prismatic.................... 29.
Achenes with 5—10 often scarcely visible ribs more or less terete ...
CY AR Cl ae ieee Momen, ieecee seed, ds. a0.
Semishrubs; achenes with somewhat winged ribs. Ornamental plants
PRU TUM RUE ROK 1520. Argyranthemum Webb. ex Sch. Bip.
Herbs; achenes with 3 somewhat thick ribs and usually 2 dark
colored punctate glands in upper part of spine uu... ee
ERA UE, oo EE 1527. Tripleurospermum Sch. Bip.**

Achenes lacking corona, with 5 inconspicuous veins ................ Ae
Achenes with distinct corona, if lacking, then with (8)10(12) quite
Prebainentt-110- like Wed 8 soso ceeceseeennsvevevinenasenessestah I 32.

Semishrubs; involucre cupulate or narrow cupulate, 4-7 mm in dia
and 4-6 mm long; ligulate florets 1-10 (entirely absent in some
capitula), with broadly ovate or oblong 1.2—8.0 mm long limb..
SERED evened ced ROM chia reed iB eke Suen 1543. Brachanthemum DC.
Herbs, rarely semishrubs, involucre patellate or broadly cup-shaped,
(6)8—20(40) mm in dia and 4-10(15) mm long; ligulate florets 10—
30 (or more) with oblong or, linear, 8—30(60) mm long limb. ..
SOA eA Ue iiire 1541. Dendranthema (DC.) Des Moul.

Ligulate florets yellow .......... cece 1538. Tanacetum L.***
Ligulate florets white, less often pink or of various other shades ...
RA BOO. due... Sheu adal Suze Saxe 33.

Cf. also couplet 48.

** Cf. also couplet 19.
*** Cf. also couplet 42.

33.

34.

7.

38.

39,

40.

Corona of achenes 1.5—2.4 mm long, auriculate; corolla of ligulate
florets fused with ovary; stems highly woody at base. Mountains of

Soviet CentraltAsiaws a.cd-cael te Rity: 1529. Spathiapappus Tzvel.
Corona of achenes not auriculate, often lacking; corolla of ligulate
florets:not fused with Ovaryeritese 253. b 2 tle eedeetemteeree tee 34.

Achenes with well developed, but sometimes very short corona, 5—
8 (rarely up to 12)-veined; lamina usually divided variously, rarely
entire: weedeat deen 1528. Pyrethrum Zinn.*
Achenes of tubular florets lacking true corona, but often with veins
apically projecting as obtuse teeth; veins (8)10(12), prominent,
forming ribs; leaves usually entire, toothed or slightly lobed, less
often 3—5-fid, with linear, entire lateral lobes ...............:::csecceeee 35.
Ligulate florets sterile; lower leaves falling early. Marsh plants ....
EI GARE Sa AAGR. erie teens 1523. Leucanthemella Tzvel.
Ligulate florets fertile; lower leaves long persisting. Meadow plants
veibisittba Tienda Mptach ithe eek NS oe Ne eat 1522. Leucanthemum Mill.
Achenes lacking corona, but sometimes with obtuse upper margin
slichtly raisediomanciside:wosenn mata Od Reet are Ae Si
Achenes with usually well developed corona, sometimes very short,
but always with sharp and somewhat toothed or lobed margin ... 42.
Plants entirely glabrous; capitula heterogamous, with one row of 1-
10 fertile pistillate ray florets and numerous bisexual, but usually
sterile disk florets, densely clustered because of sticky resinous
secretions; achenes globose-obovoid, highly compressed dorsally.

Siberia; Han Eastusmiatesnes, 2..cevr ees 1546. Filifolium Kitam.
Plants more or less hairy; disk florets fertile, free; achenes more or
LOSS LENGE. .icaccieswnes.ncraianqsdscpieasen: pengiuanmele pelea el ete mene ete. ae 38:
Semishrubs with entire or 3(5)-fid leaves, sparsely stellate hairy;
usually corolla also stellate hairy .......... 1549. Kaschgaria Poljak.**
Pubescence dense; hairs simple and bifurcate; corolla glabrous or
Withomnple hairs hay ic..:boae ..cxuciatg ae. ee ee ee ciel

Achenes arcuately bent, narrow-cylindrical (6-8 times as long as
wide), rather densely covered with short glandular hairs; biennial or
perennial monocarpic plants. Soviet Central Asia ...................
id doadl ded sulted Beat ok Beeb ck aes 1540. Pseudohandelia Tzvel.
Achenes broader, entirely glabrous; perennial polycarpic plants ...
cadet tN MIR sd SO. Ot pees. deal coterie Seite penne 2 40.
Herbs, always with highly reduced nonflowering shoots (forming
rosettes of basal leaves); stems usually with only few reduced leaves;
capitula homogamous (with only bisexual tubular florets). Soviet
Central Asiayans. wath. nelscieet nein: 1545. Hippolytia Poljak.

* Cf. also couplets 24 and 23.
**Cf. also couplet 11.

41.

42.

43.

44.

45.

46.

47.

48.

*

7

Semishrubs or herbs, with densely leafy stems, usually lacking re-
duced nonflowering shoots; capitula heterogamous (ray florets pis-
tillate), rarely some homogamous (then lamina entire) ......... 41.

Capitula in lax corymbose-panicles, few-flowered (less than 20 flo-
rets); corolla of disk florets hirsute along teeth. Caucasus .............
CU RENT, SRA ee 1550. Artemisia L. (A. fasciculata M.B.)
Capitula in dense or lax corymbose inflorescence, rarely solitary,
usually with more than 20 florets; corolla glabrous. Soviet Central

PRL seco ay tcuancoternct tert ener donmeanens tierce naan torent 1544. Ajania Poljak.
Capitula heterogamous; ray florets (sometimes few!) pistillate, of-
ten ligulate; disk florets bisexual ................. 1538. Tanacetum L.*
Capitula homogamous; all florets bisexual ..............ccceeeeeeeeeees 43.

All leaves entire, oblong or ovate, small-toothed; capitula numer-
ous; in dense corymbs; stems 30-120 cm high; aromatic garden
PUANICS OLLI CSCABINGE UO UME WHIT 2...00...1fecct-sasettectaticesceceacessvesecteoeeceessee
2. dee en Rd oe 1528. Pyrethrum Zinn. [P. majus (Desf.) Tzvel.]
LEAVES PinMatiSCChOn PUMMOLNPATINS 6 Eid. ccecesscscsocscacrsetesnses 44.
Involucral bracts fewer and wide, almost equal in length; achenes
smooth, with (8)10(12) very prominent veins forming ribs; dwarf,
almost stemless semishrubs, caespitose, with capitula on long scapes.

Chu-T Mountains 1.0000. ear. Ral 1535. Cancriniella Tzvel.
Outer involucral bracts a third to two-thirds as long as inner ones;
praaies GE itierems Wait S i.) 2605... A A, 45.

Capitula on very short (up to 6 mm long), arcuate peduncles; pa-
ppus of achenes as pappiform corona of 10—20 narrow-linear scales,
longer than achene; dwarf, prostrate, semishrubs. Western Tien

MARRAEA SBE OEE ae L ss eet ca co PORTS RAL CEL 1531. Ugamia Pavl.
Capitula on longer, always erect peduncles; corona of achenes
SCPUCTUT MY Ginretent £2. G CTIA Te, 20a EAE, 46.
Pepenietah MCT NS 112200. TRL... MARES ARS 47.
Semishrubs or semishrubs with somewhat woody stems near base.
Mountains of Soviet Central Asia 3.00. 00.1s 0 itteeseaeeeeede eee 48.
Achenes with entire auriculate corona; capitula in dense terminal
corymbs. Transcaucasia ............cc0 1539. Hemipappus C. Koch

Achenes with corona more or less uniformly bordering their tip,
entire or lobed; capitula in lax corymbose-paniculate inflorescence,
less often in somewhat dense corymbs or solitary. Soviet Central
Faia CUD. AL. ene IE. PRE ERI 1536. Cancrinia Kar. and Kir.**
Ovary and achenes (and often also receptacle) densely hairy .....
ra SR cae Sa Ah 1532. Trichanthemis Rg]. and Schmalh.***

Cf. also couplet 32.

** Cf. also couplet 24.
*** Cf. also couplet 27.

+ :JOvaryiand.achenes glabrous gcxisis.cbtiei.: wapeseehcteesndage iia aet oe 49.
49. Leaves 3-fid or entire; corona of achenes mostly uniformly border-
ing their tip, usually divided to base into oblong scales; semishrubs
Me re en 1534. Lepidolopha Winkl.

+ Leaves (except uppermost) pinnatisect, with 2—5-lateral lobes; co-
rona of achenes entire or lobed, but always obliquely truncate ei-

ther completely or almost to base; semishrub ............:scsssseeseseseseeeeee
sMaiitial adcoole. abborrs, sepaucsty LOO tek 1533. Xylanthemum Tzvel.

Subtribe 1. ANTHEMIDINAE O. Hoffm. in Pflanzenfam. IV, 5
(1889-1894) 268.—Tribe Senecionideae Less. subtrib. VI.
Chrysanthemeae Less. 1. Anthemideae Less. Synops. Comp. (1832)
247 and subtrib. VII. Artemisieae Less. 1. Santolineae Less. ibid 258.—
Receptacle with scales.

GENUS 1513. Anthemis !:?

L. Sp. pl. II (1753) 893; Hoffm. in Pflanzenfam. IV, 5 (1894) 271
p. p-—Maruta Cass. in Dict. Sc. Nat. XXIX (1823) 174.—Cota J. Gay
ex Guss. Fl. Sic. Syn. II (1842-1844) 866.

Capitula many-flowered, with distinctly developed ray (ligulate)
florets or with only disk florets; ray (ligulate) florets 1-rowed, almost
always pistillate, other florets staminate-pistillate (amphisporangiate),
5-toothed, usually with compressed tube, sometimes basally swollen.
Involucre imbricate; bracts sometimes scarious or fimbriate.

Receptacle covered with scales. Achenes cylindrical-obconical,
obpyramidal and angular, sometimes tetraquetrous, more or less
longitudinally striate-ribbed, with corona of small teeth or with long,
auriculate, lateral processes, or almost entirely lacking corona,
sometimes distinctly compressed. Annual or perennial herbs, sometimes
semishrubs.

Lectotype of genus: A. arvensis L.

Until recently it was believed that the genus Anthemis was com-
prised of 130 species [A. Rendle. The Classification of Flowering
Plants, II (1938), London], which are mainly distributed in West Asia
and Europe. But this number may increase significantly if the limits of
species in this genus are understood in the sense of concrete geo-
graphic races (much splitting of polymorphic species such as A. tinctoria

'Treatment by An. A. Fedorov.
*Ancient Greek name used by Linnaeus to designate the species resembling A.
arvensis. The word “Anthemis” is derived from “anthos,” i.e., flower.

9

L. s. 1., A. rigescens Willd. s. 1., A. montana L. s. 1., etc., is possible).
Moreover, new species continue to be described even now also from
other groups of the genus. Therefore, the provisional number of
Anthemis species may be put at close to 150. In the USSR 51 species
of this genus are found.

The majority of species of this genus is concentrated in the region
of West Asia and the Caucasus and partly in the Mediterranean Re-
gion. In Europe, a few species are distributed in the north, only as far
as Scandinavia; only one species is found in Siberia, reaching approxi-
mately as far as Krasnoyarsk. Members of the genus Anthemis are
absent in the Arctic and the tropics. Ecologically, most of the species
are montane, subalpine, partly alpine or rock and semidesert, partly
steppe, rarely forest, as well as desert and weedy plants.

There is no comprehensive monograph of the genus. Therefore,
the publications of Boissier, especially Diagnoses and Flora Orientalis
and a series of papers of D.I. Sosnovsky on Anthemis (cf. literature
cited under different species) are of great importance in understanding
the species found in the Caucasus and Western Asia. Numerous
Anthemis species were described mainly from Asia Minor by the
German Botanist Karl Koch. However, now that the type (authentic)
specimens of Koch’s species preserved in Berlin have been permanently
lost, the species established by him and attempts to understard them
are, in a real sense, the “crux et scandalum botanicorum.”

At present, even with the joint efforts of Soviet botanists to establish
not “aggregate” species but specific geographic races, no identification
of Koch’s species, described carelessly and unintelligibly, with the
species described by other authors—an approach followed, for example,
by Boissier (1. c.)—can help in understanding Koch’s types. Hence, we
are surprised at the boldness shown by D.I. Sosnovsky and A.A.
Grossheim (cf. literature citations), who confidently reported as a matter
of fact the quite unresolved species, Anthemis platyglossa C. Koch,
A. brachyglossa C. Koch, and others in the composition of the “Flora
of the Caucasus.” A botanist using the “Flora of the USSR” can clearly
see from our notes made in the diagnoses of the different species how
confusion has occurred in the names and nomenclatural identities.
Therefore, we have listed certain, especially obscure species of Koch
only in the notes as doubtful species, since hope for strict scientific
clarification of them is essentially lost forever.

Of the two lectotype species proposed for the genus Anthemis, A.
arvensis L. should be preferred over A. maritima L. Although the
lectotype A. arvensis was proposed by A.S. Hitchcock and M.L. Green
only in 1929 and again by E.P. Phillips in 1951, whereas A. maritima
was proposed as a lectotype in 1913 by N.L. Britton and A. Brown,

10

still A. arvensis is more appropriate as the type species of the genus
Anthemis L., because it usually has been included in the long established
type section, which until recently was called Euanthemis (now Anthemis
in accordance with the currently operative nomenclatural rules). The
second lectotype, i.e., A. maritima, in our classification, cannot in any
way be included under this section. Besides, an examination of A.
arvensis convincingly shows that this species actually (even in habit)
includes all the typical characters of the genus. However, as often
happens in similar cases, the species considered by us as the lectotype
of course cannot be considered as the prototype of the genus in the
true genetic sense and is not the oldest (nor the most primitive) species.
In this respect, attention should be paid to other sections of the genus,
partly to section Cota and especially to section Rumata, which not
only include perennial herbs but also semishrubs, and species of a
clearly relict nature. In this sense, species like A. fruticulosa MB., A.
trotzkiana Claus ex Bge. (from section Rumata) and forest species of
the Caucasus Mountains A. zyghia Woron., A. marisenigri Fed., A.
woronowii Sosn. (section Cota), and others should be recognized as
the most typical (oldest) species of the genus Anthemis.

Anthemis species are of interest as ornamental plants. Particularly
valuable in this context are the Caucasian species, for example, A.
marschalliana MB., A. sosnovskyana Fed., A. macroglossa Somm.
and Lev., A. saportana Alb., A. iberica MB., and A. zyghia Woron.

1. | Achenes distinctly compressed; receptacular scales carinate ...... 18.
+ Achenes not compressed, roundish or angular in cross section;
receptacular scales without distinct keel ...........esceeseeeseeeeereeeees 2.

2. | Semishrubs or perennial herbs, with rhizome or many-headed hard
and woody root; stem simple or with few branches, arising from
short caudices, also bearing quite frequently nonflowering shoots
with lax rosettes of leaves; leaves pinnatipartite or pinnatisect,
sometimes pinnatilobate, usually crowded in lower part of stem .

+ Biennials or annuals, with pinnate leaves, lacking basal rosettes ee
CAUGICES LL eas, A Re oe, 2a eee 3.
Achenes angular (usually tetraquetrous); receptacular scales oblong-
lanceGlate 31.2866. M.A OR Be, Ba ee eee 43.
Achenes terete; receptacular scales narrow-lanceolate ................ 47.
Peripheral (ligulate) florets golden-yellow ...........:sscsseseeeseeeeeees 5.
Peripheral (ligulate):florets: white a isac..8..oheeden.. aes 6.
Leaves green, subglabrous or mildly pubescent; involucral bracts
with blackish-brown border ..............0:.008 15. A. sosnovskyana Fed.
Leaves silvery-gray, sericeous; involucral bracts lacking distinct
blackish-brown border ...............:000000 14. A. marschalliana Willd.

OP +

+

oa

-

10.

as

13.

14.

11
Involucral bracts with wide dark brown or blackish fimbriate border

Involucral bracts pale, sometimes with very narrow dark stripe along
margin or with brownish midrib; inner bracts usually broadly scarious
at apex and sometimes fimbriate 0.0.0.0... cscesssseeeeesecseeeeseeseneees 11.
Leaves densely pubescent, scarious-lanate, with approximate lobes
of segments; involucral bracts dark brown along margin; stems
short, usually 10(15) cm high ........... 5. A. ptarmiciformis C. Koch
Plants sparsely pubescent or subglabrous ............:..c:ccccesseceeeeees 8.
Achenes finely punctate; plants caespitose; stems with many
branches, nonflowering shoots bearing rosettes of leaves; capitula
rather large, up to 4.0-4.5 cm in dia; stems usually numerous ......
dav vis-beortaad drraitanwiek 4. A. carpatica Waldst. and Kit. ex Willd.
Achenes not distinctly punctate; plants not caespitose ................. 9.
Leaves glabrous, pinnatilobate or pinnatipartite into somewhat broad
obtuse lobes; corona of achenes short, crenate-lobate ..............:c:0000+
ents A teetateneetetteatas..f..2:bs-.01405..ckdaninles 1. A. saportana Alb.
Leaves pinnatipartite, more or less pubescent; leaf lobes linear...
drancts- tednaath peal. tdaierts His .kanwanateaienl.... 10.
Limb (ligule) of peripheral florets abruptly narrowed toward base;
leaf lobes approximate; involucral bracts obtuse, with roundish apex
sien Reda saan Hie Quast EER a oti ts scena 3. A. anahytae Woron.
Limb (ligule) of peripheral florets basally carinate; leaf lobes
somewhat remote; involucral bracts acuminate .............000:c00
aa Bite aha sss tiie nae cnaastridiinrsn epsntnaned 2. A. iberica MB.
Semishrubs, with many woody but fragile (easily breaking) caudices
and nonflowering shoots with rosettes of leaves in lower part; hairs
usually dense, often white-tomentose; stems sparingly branched or
simple; capitula large, about 2.0-2.5 cm in dia; involucre pale,
white-pubescettindin).ctesnt a oawas.. ince. daiisslacpeenmedgsiecn. 2.
Perennial herbs; hairs not white-tomentose ..............c:ccscceceeseeeeees 14.
Stems simple, short, usually 10-15 cm high; caudices somewhat
spreading at base, thick; leaves densely white-pubescent ...........
\ abt canisetbe caghoaehinn aaninemedian cpa 11. A. fruticulosa MB.
Stems branched; usually 15—30 cm high, sometimes reduced; plants
herbaceous in upper part; pubescence gray or white, but usually not

VEN GOUSE anie vnaascs near cometayitie-tan-ceeubesne 13.
Receptacular scales abruptly acuminate; caudices short, somewhat
thick, very fragile, numerous ........ 12. A. trotzkiana Claus ex Bge.

Receptacular scales 3-toothed at apex; caudices long, slender, few;
stems slender, with long slender branches ..... 13. A. calcarea Sosn.
Lobes of leaf segments very small, about 2 mm long, approximate;
stems very slender, leafless over large part ............:ccccccseeeseseeees 13.

12

18.

19.

22;

pie h

26.

Lobes of leaf segments oblong, sometimes divergent and divaricate,
3-5 mm long; stems strong, erect till flowering ..............::10 16.
Achenes sulcate-ribbed, tuberculate ................ 6. A. anatolica Boiss.
Achenes almost tetaquetrous, tuberculate ............ 7. A. emiliae Sosn.
Stems with reddish longitudinal stripes; leaves pinnate, with few
broad 1OBES. (52... 10. A. tempskyana Freyn and Sint.
Stems without red stripes; leaves bipinnatisect into narrow lobes...
~ USERRA RUE cfs ol cvnnececuven cee ttksGihae tect ne: SN as i Wie
Plants short, 10(15) cm high; disk about 0.7 cm in dia .............cce
STALE RA RRO. SR Seleaeee eee eee 8. A. sterilis Stev.
Plants tall, 30-40 cm high; disk 1.0—1.5 cm in dia «0.0... eeeesceeeeseeeeees
DETTE. SORRY, ROTATE, coe TE, Sie Goren 9. A. tranzscheliana Fed.
Perennial, with thick and strong WOOdY rOOt ...........csssesseesreeeeeees 22,
Anmuals O00 eA Ot 15:
Lobes of leaf segments distinct, somewhat broad, crenate and sharply
dentate; stems and their branches sinuate .................:c:sssssseeeeeeseeee
IAT oD. cenecuscorcivgazes 46. A. wiedemanniana Fisch. and Mey.
Lobes of leaf segments approximate, narrowly linear, with oblong
teeth; stems erect, with straight, divaricate branches ................ 20.
Inflorescence peduncle thick, clavate ........ 44. A. coelopoda Boiss.
Inflorescence peduncle almost not clavate ............:csccssceesseeeeseees 20.
Lobes of leaf segments pinnatipartite, roundish at apex .................
TB, RRL NUREES:, RIS La 45. A. austriaca Jacq.
Lobes of leaf segments pinnatisect, somewhat acute .................00+
Hd tten cel tebeousvesdediavesessconeavss ductadessaadlene aaa etna te ose 43. A. altissima L.
Limbs of peripheral florets (ligule) yellow .............ccsceeseeeseees 23.
Limbs of peripheral florets (ligule) white ..............cssseeesseseeees el
Stems always simple, about 20-30 cm long, less often more, with
single capitulum; leaves crowded in lower part of stem, pubescent
but greenish, with small, approximate lobes ..............cccceseeeeeeees
de ceonovenaddaadeavaaerscesaesde ete aN Reet eae nEE 26. A. monantha Willd.
Stems, as a rule, branched and usually high, generally taller than 30
cm) leafy ‘up*to middleiand above .A42%.. 2202.2) Boise san... 24.
Stems branched above; inflorescence peduncles distinctly clavate
iStape Mei SO ORR MRS Se 24. A. saguramica Sosn.
Stems branched from middle or base, or even simple; inflorescence
peduncle not clavate \s:2:ci2c...0ciscccccuscesdovscevacevraueskerauas eeeeen ee tenes 2S.
Plants 30-50(70) cm high; stem erect or basally arcuate, but never
spreading or bend’downward ... 22. Ra ee ee 26.

Limb of peripheral florets (ligule) short, a half of disk, truncate;
capitula 2omi-1n dia 2g. RU Ree 23. A. maris-nigri Fed.

=

30.

aT

oe:

Plants densely lanate-hairy; leaves large, up to 4 cm long, with
broad lobes; stem densely leafy up to middle ............ cece
Rear Wi ahhh et ML iis ie coke pati oh! BL 25. A. markhotensis Fed.
Plants sparsely and finely pubescent or almost glabrous ............ 28.
Plants grayish; involucre hairy; segments of leaves narrow-linear,
with sharp and pointed teeth; limb of peripheral florets (ligule)
HEU VEHOW tot ck aah 22. A. subtinctoria Dobrocz.
Plants green, almost glabrous; involucre subglabrous; segments of
leaves linear, somewhat broad; stem reddish; limb of peripheral

florets (ligules) bright or deep yellow ............... 21. A. tinctoria L.
Involucral bracts with prominent, bright green midrib; leaves highly
a Mie Ls hg sh le gta Ato e ip Mild la kd he a 30.

Involucral bracts with brownish midrib; leaves oblong, up to 3 cm
long, gray, densely borne in middle of stem; leaf segments narrow-
Tiniedr, GIS Ety STAVISI ::2.ic-seeeoeeisee ise: 29. A. zephyrovii Dobrocz.
Stems usually with single capitulum; leaves small, about 1 cm long,
densely borne and crowded in lower half of stem, greenish ............
week fe A SIRS PELL RL td LEE ar SP at ba 28. A. debilis Fed.
Stems branched, with many capitula; leaves as long as in previous
species, covering stem up to two-thirds, grayish from pubescence ...
recep tetdh! 5 A bd hl Cpe eh teil aed el & 27. A. cretacea Zefir.
Plants with long, creeping stem and branches, subglabrous; stem
branched above; leaves reduced, with toothed segments ............
dealt tran ce AAA PR pe tebe ene ot tM oceed “arabe th 37. A. euxina Boiss.
PPAR WEP CLEC BECIT fc lsc ive ec caticscaneconccecacscoccetaott eee Ren toes wi 32.
Leaves densely velutinous-tomentose, silvery, with broad and
approximate segments, broadly obovate; involucral bracts with

brownish-black border .............c:cceeseseeeseeseeeees 41. A. zyghia Woron.
Leaves glabrous or pubescent, but never velutinous .................. a3.
Involucral bracts with dark brown or black border ................066 34.
Involucral bracts pale, whitish, or greenish ...............::ceseeseeeees a

Leaf segments broad, reduced, approximate, obtuse, truncate-toothed;
plants bright green, somewhat pubescent; capitula not more than 3—
ROUT IE UE re ae coe iatrescr te artesenereceesie tt 40. A. abagensis Fed.
Leaf segments narrow, oblong, sometimes leaves thrice-pinnate,
with very small lobes; capitula large, 4—5 cm in dia .................. a.
Leaves finely divided, sometimes thrice-pinnate; lobes small .......
erect ps Lt 4) ae pee ts Pet 39. A. schischkiniana Fed.
Leaves divided into narrow or somewhat broad segments lacking
small pinnate lobes, truncate-toothed .............:::ccseeesssesseeeeseeeeeeeees 36.

14

36.

42.

Leaf segments oblong-linear; involucral bracts somewhat obtuse,
broadly scarious-fimbriate and blackish-brown along margin ..........
ee ee ee ne 38. A. macroglossa Somm. and Lev.
Leaf segments and lobes narrow-linear, acuminate, grayish,
involucral bracts sharply oblanceolate or pointed deltoid, with
distinct and narrow blackish-brown border ..........::::esseceeceeeees
ad Rega a0 cent Noa Haak edna tery at ae 42. A. melanoloma Trautv.

Seth pli nasty itachi baags cuba Ban Suasddes« dictly tue ON, cat eagle te ai, «2 38.
Leaves green, subglabrous or sparsely pubescent ...........::c::00000 42.
Plants densely tomentose; leaf surface entirely covered with
tomentum, gray; internodes of young branches white-pubescent ...
ee ee ee 36. A. talyschensis Fed.
Plants grayish but not densely tomentose; leaf surface always clearly
Visible, <.:te1..-susgbiv pions uta eriaa neat iecames haat Qerurmnateneah tebege 39.
Capitula large, up to 4-5 cm in dia; disk up to 1.5—2.0 cm in dia
Ce ee Es eee OR ee io cs 31. A. jailensis Zafir.
Capitula up to 3-4 cm in dia; disk up to 1 cm in dia ................ 40.
Stems numerous, usually highly branched; inner involucral bracts
With wustypiimbtia atapexocr. 6 iste... cca 34. A. dubia Stev.
Stems few or solitary, slightly branched, rarely simple; involucral
bracts with or without deep brown fimbria at apex ..............000 41.
Leaves minute, 1.0—-1.5 mm long, reduced, with approximate lobes,
orbicular, crenate-pinnate ............-.:eeeeeeees 35. A. woronowii Sosn.
Leaves large, up to 3 cm long, oblong, with divergent segments and
lobes; segments unequal, lobes acuminate ................escesseseseeeeeeseenees
Lamas es, cal, aanab i iaueuea teacaleanan tailed, eden 32. A. dumetorum Sosn.
Leaf segments very narrow, long, lanceolate-linear, acute, along
margins incised and sharply toothed; leaves vertical; capitula 2.5—
8.0) cain Gia weiss, .cteali.. ceamuerye beaut tae 33. A. linczevskyi Fed.
Leaf segments linear-oblong, somewhat broad; capitula large, about
4:0-4.5 jor dingy ava Sieioss , wench coettcteeec hebeeins 30. A. rigescens Willd.
Plants small, 10-15 cm high, silvery-sericeous with slender stems
and their branches divaricate from base .........ccccssceseeseeseeeeseees 45.
Plants 20-30(40) cm high, appressed and grayish-pubescent or
villousyistems branched: butverect ters: vor deseseecacrog eee 44.
Plants lanate-tomentose-VillOusS............:.:c00+ 17. A. ruthenica MB.
Plants appressed-pubescent, grayish...............0:00 16. A. arvensis L.
Corolla tube of disk florets swollen. belOW............ccccccsseceeseseereeees
eiastonibah barecbphag: «Shes AMY. Os 2. eons. bttebs ea 18. A. candidissima Willd.
Corolla, tubeiof other shapes, i:.wssecancseejunes-saheek fenton eee 46.
Corollastube straightiiagehs diidee beds eclevee.) 19. A. grossheimii Sosn.

15

+ Corolla tube with constriction in middle, resembling sand clock in

SHape Wes.c:: ee RRA OE Pe a, 20. A. hirtella Winkl.

47. Leaves twice pinnate, lobes numerous, small, linear; plants usually
BS 2h hth 1. £0) hh asceet EMOR Halg hb DIR, ARS. 48.

+ Leaves pinnate, with small number of segments and lobes; plants
Siiall-S~1O¢IS Pom Mish Kh RR RAMAR ID. BELL aa,

48. Lobes of leaf segments reduced, narrow-linear......47. A. cotula L.
--)SAGbEeS GF leaf sepments Ubion py florins Mate NOE RE RN BN.
ABO OUP EST ARI NTS) 620002 BOS 48. A. lithuanica (DC.) Trautv.

49. Leaves usually linear, often entire, lacking segments or with few
laterae TOBES* ene. 208. 51. A. deserticola Krasch. and M. Pop.

BP OL Nibeaves pinnate 2. 102228. AR BOY BR RD SPs DARE? 50.
50. Plants pubescent; lobes of leaves reduced, narrow-linear ...............
BOR PRY AO ORI BAe DAI, SRAM 49. A. odontostephana Boiss.

+ Plants almost entirely glabrous; lobes of leaves long, narrow-linear
WATE ARR AS 50. A. microcephala (Schrenk) B. Fedtsch.

Section 1. Rumata Fed. sect. nova in Addenda, XXV, 865.—
Euanthemis DC. Prodr. VI (1837) 7 pp.—Sect. Chamaemelum Cass.
in Dict. Sc. Nat. XXIX (1823) 279 pp.—Sect. Euanthemis ser. Perennes
Boiss. Fl. or. III (1875) 278.—Ser. Sphacelatae Sosn. in Vestn. Tifl.
Bot. Sada, Ser. 2, 3 (1927) 173.—Achenes angular in cross section,
obconical or obpyramidal, not compressed. Receptacle concave or
hemispherical. Receptacular scales more or less carinate and usually
acute. Involucre bracts with blackish border or pale, usually with wide
scarious margin. Perennial herbs or semishrubs, with somewhat hard
rhizome or woody root and simple or slightly branched stems, each
bearing single capitulum. Leaves lobate or pinnate, crowded in lower
part of stem forming loose rosette.

Type of section: A. saportana Alb.

In section Rumata, we have included the montane perennial
(rhizomatous) and partly semishrub species of the genus Anthemis,
characterized by broad scariously bordered and fimbriate involucral
bracts, frequent caudices and nonflowering shoots bearing leaf rosettes.
In the true (genetic) sense, the species of this section are the most
ancient, but the initial nomenclatural type of the genus, Anthemis
arvensis L., is included in type section Anthemis, which is
phylogenetically definitely more advanced than section Rumata.

Series 1. Saportanae Fed.—Involucral bracts with dark brown or
blackish, broad, scarious-fimbriate border. Leaves pinnately lobed,
pinnately parted or pinnately divided into numerous segments and lobes.
Capitula large, up to 3—4 cm in dia; limb of peripheral florets (ligule)
white. Plants exclusively alpine.

16

1. A. saportana Alb. in Bull. Herb. Boiss. III (1895) 92; ej. Prodr.
Fl. Colch. 135; Sosn. in Vestn. Tifl. Bot. Sada, Ser. 2, 3, 173; Grossh.
Fl. Kavk. IV, 120; Sosn. in Fl. Gruzii, VIII, 314; Kolak. Fl. Abkhaz.
IW, 226.

Perennial. Whole plant covered with inconspicuous, sometimes
somewhat tomentose or appressed, very small hairs, sometimes almost
glabrous, short, usually about 10—15(20) cm high; stock (caudex) highly
branched, somewhat woody, brown, covered with scaly remnants of
dried leaves, besides nonflowering shoots terminating into leaf rosette;
stems basally slightly arcuate, leaf only below, with single capitulum
and few longitudinal stripes, not clavate below capitulum; rosulate
leaves and subtending leaves pinnately lobed or almost palmately lobed,
or pinnately parted, sometimes with few lobes, ovate-cuneate, narrowed
into petiole; segments (or lobes) broad lanceolate, obtuse, sometimes
with 2—3 terminal teeth; cauline leaves short-petiolate, cuneate-oblong,
divided into few broadly linear or linear-deltoid, obtuse or acute lobes
with minute cusps, sometimes bidentate, approximate or slightly
divergent; upper leaves narrower, less incised or almost entire. Capitula
rather large, about 4 cm in dia. Bracts of sparsely white-pubescent
involucre with blackish border; outer bracts deltoid-lanceolate; inner
bracts ovate-lanceolate, with wide scarious margin, slightly incised,
not acuminate. Receptacular scales linear-lobate, blackish-tipped,
divided, acute. Ligules white, oblong-elliptical or elliptical, as long as
disk. Achenes longitudinally sulcate, basally extended; corona a fifth
as long as achene, crenate. Flowering June to August. (Plate I, Fig. 1).

In alpine zone, on rocks, predominantly limestone screes, as well
as in meadows, near thawing snow.—Caucasus: Western Transcaucasia
(particularly in mountains of the Bzyb River basin and in region of the
Caucasian Reserve). Endemic. Described from Abkhazia (“along high
rocks of the Adzituko Range”). Type in Leningrad; isotype in Geneva.

Note. The distribution of A. saportana is confined to the western
part of the Main Caucasus Range, mostly to the limestone ranges
(Abkhazia, region of the Caucasian Reserve). In the east, the range of
this species does not even reach Elbrus, and beyond that another race—
A. iberica MB.—predominates. In turn, in the Lesser Caucasus, the
latter species is replaced by a separate race—A. anahytae Woron. In
Asia Minor, beginning from Ararat, other species of the series
Saportanae are found. A. carpatica, found in the mountains of Central,
Eastern and Southern Europe beginning at the Eastern Carpathians, is
the far western species of this series.

17

2. A. iberica MB. FI. taur.-cauc. II (1808) 328; ej. ibid. III, 580;
DC. Prodr. VI, 7; Spreng. Syst. veget. III, 593; Ldb. Fl. Ross. II, 2,
522, excl. syn. nonnull.; Boiss. Fl. or. III, 289; Sosn. in Vestn. Tifl.
Bot. Sada, Ser. 2, 3 (1927) 173.—A. iberica var. elata and var.
tomentosa Sosn. ex Grossh. Fl. Kavk. IV (1934) 121; Sosn. in FI.
Gruz. VIII, 315.—A. iberica var. minor Boiss. 1. c. quoad pl. e Cauc.-
orient.—A. montana Georgi, Beschr. d. Russ. Reichs. III, 1254, nom.—
A. caucasica C.A.M. Verzeichn. (1830) 75, non Adam (1805), nec
Hernem. et al.—A. nivalis Adam ex Tausch. in Ldb. op. cit. 523, pro
syn.—A. alpina var. B. Willd. Sp. pl. III (1804) 2178, non A. alpina
L.—Pyrethrum ptarmicifolium Willd. ex Ldb. op. cit. 523, pro syn.—
P. nivale Ad. ex Ldb. |. c. pro syn.; Boiss. 1. c. pro syn.

Perennial, with multicauline habit, 10—15(20) cm high, with woody
root producing numerous, branched, reduced brown caudices, covered
with remnants of dried petioles; nonflowering shoots bearing leaf
rosettes. Stems ascending from arcuate base, erect above, 15—20(30)
cm high, unbranched, unicapitulate, longitudinally striate, glabrous
above, mildly pubescent below. Leaves (up to middle cauline leaves)
petiolate, mildly appressed-hairy, pinnate, with linear, three-parted,
less often 5-parted, sometimes even entire and rather narrow segments;
lobes also narrow, acute and short or long, linear. Capitula large, about
4-5 cm in dia, with white ray (ligulate) florets; ligule oblong-lanceolate,
3-toothed. Involucral bracts scarious along margin and apex, lanate-
tomentose, acuminate with brownish-black border. Receptacular scales
lanceolate, scarious at apex, somewhat cuspidate. Achenes apically
along margin with denticles of indistinct corona. Flowering June to
August. (Plate I, Fig. 2).

In alpine zone on stony places.—Caucasus: Ciscaucasia, Dagestan,
Eastern Transcaucasia (north); Western Transcaucasia (eastern part).
Endemic. Described from Georgia. Type in Leningrad.

3. A. anahytae Woron. ex Sosn. in Vestn. Tifl. Bot. Sada, Ser. 2,
3 (1927), absque diagn. lat., nec ross., sed tantum cum observ. brevibus
de florum forma Descr. nostra in Addenda, XXV, 865; Grossh. FI.
Kavk. IV, 120, diagn. brev. ross. in clave; An. Fed. in Sov. Bot., XIII,
4, 18, nom.

Perennial. Subglabrous or sparsely pubescent plants; rhizome
woody, brown, producing reduced caudices and nonflowering shoots
bearing leaf rosettes. Stems rather short, 10—15(20) cm high, ascending
or erect, mostly leafy at base. Lower cauline leaves and leaves of
nonflowering shoots obovate or lobate, more or less long-petiolate
(other leaves sessile), pinnate; leaf segments oblong-lanceolate or linear,
entire or lobed, the latter entire or 3-toothed at apex; rachis somewhat

20

18

broad, lacking teeth; upper cauline leaves reduced, finely lobed or
entire. Capitula rather large, about 4—5 cm in dia. Outer bracts of more
or less hairy involucre deltoid; inner bracts oblong or oblong-obovate;
all bracts broadly scarious and dark brown or blackish on margin, with
gradually increasing membranous roundish apex from outer to inner
bracts. Ligules white, oblong-elliptical, indistinctly 3-toothed at apex,
narrowed toward lower side. Achenes as in A. iberica. Flowering June
to August.

In alpine zone on rocks and screes.—Caucasus: Southern
Transcaucasia. General distribution: Balkans-Asia Minor (east).
Woronow collected as the holotype a specimen from Azerbaidzhan
(Murov-Dag Range, Gyamysh Mountain). This specimen and also the
isotype are preserved in Leningrad.

Note. The specific epithet is derived from the Armenian feminine
name Anait. Unfortunately, it is impossible now to determine whom
J.N. Woronow had in mind when naming this species. The spelling of
the specific epithet varies on the labels written in Woronow’s hand:
instead of the transcription “Anahytae” published by Sosnowsky (I. c.)
and Grossheim (1. c.), Woronow wrote “Anahitae.” However, the first
version, which had already put into print, must be recognized as the
valid variant.

Specimens are preserved in Leningrad from Surbkhach Mountain
in Asia Minor, some of which were identified earlier as A.
ptarmiciformis C. Koch, that should be included in A. anahytae. There
is also no doubt that all specimens from Alagyoz in Armenia should
also be included under this species. Thus, A. anahytae, growing in the
mountains of the Lesser Caucasus, is distributed beyond the limits of
this region and, therefore, is not endemic to the Caucasus.

4. A. carpatica Waldst. and Kit. ex Willd. Sp. Pl. III (1800)
2179; Simonkai, Enum. FI. Transs. (1886) 316; Boiss. Fl. or. II, 290,
pro max. p.; M. Popov, Rast. i Fl. Karpat (1949) 247; Katina in Vizn.
Rosl. URSR (1950) 536; Dostal, Kli¢ k iplné kvéetene CSR (1954)
846.—A. montana Bielz apud Schur in Verh. Seibenb. Ver. Naturw. I
(1850) 104, nom. L.—A. kitaibelii DC. Prodr. VI (1837) 7, non
Spreng.—A. petraea Tenore, Fl. Napol. I (1811-1815) 1; ibid. II (1820)
246.—A. stiriaca Vest in Syll. Ratisb. I (1824) 12.—Ic.: Tenore, op.
cit. II, t. 82; Dostal, op. cit. p. 846, f. 23600 (mediocr.).—Exs.: L.
Vagner, Fl. der Marmaros, sine numero (e loco classico, ex mont. Pop-
Ivan): F. Schultz, Herbar. normale, nov. ser. Cent. 23, No. 2205 (e
loco classico); Fl. Exs. Reipubl. Bohem. Sloven. No. 1274 (e loco
classico) Fl. Exs. Austro-Hungar. No. 1002 (e loco class. Anthemidis
stiriacae, ex mont. “Jurgbaueralpe”); Domin and Krajina, Fl. Cechoslov,

23

19

exs. No. 400; Dorfler, Herb. normale, No. 4114 (sub Anthemide petraea,
e loco classico, ex mont. “La Majella’); Fiori and Béguinot, FI. Ital.
Exs. ser. II, No. 1577 (sub Anthemide petraea, e loco classico speciei).

Perennial. Glabrous or weakly appressed-hairy caespitose plants,
about 10-15 cm high, bearing numerous caudices densely covered with ©
brown scaly remnants of old leaves; non-flowering shoots bearing leaf
rosettes, or usually numerous unicapitulate stems and not leafless all
along except lower densely leafy part. Lower cauline and rosulate
leaves rather long-petiolate; lamina cuneate-ovate, pinnately cut into
few broadly linear or narrow, entire, 3-fid or 3-lobed, somewhat acute
segments; cauline leaves sessile, gradually reduced upward; uppermost
leaves entire, linear. Capitula large, about 4 cm in dia; limb of peripheral
(ligulate) florets elliptical-oblong, wide, apically 3-toothed, with middle
tooth much smaller than 2 lateral. Involucral bracts glabrous or weakly
hairy, with scarious, black-bordered margin; outer bracts deltoid, acute;
inner apically truncate, incised along membranous margin and slightly
fimbriate. Receptacle flat-hemispherical; scales oblong-lanceolate,
membranous, as long as disk florets. Achenes somewhat long, extended
below, punctate, with narrow border in place of inconspicuous corona.
Flowering June to August.

On rocky places in alpine zone. European part: Upper Dniester
(Eastern Carpathians). General distribution: Central Europe, Balkans-
Asia Minor (western part), Mediterranean Region (western). Described
from Eastern Carpathians (Pope Ivan Mountain). Type in Berlin, several
topotypes in Leningrad.

Examination of the topotypes of A. carpatica, A. petraea and A.
stiriaca, in the Leningrad Herbarium (cf. list of exsiccatae), confirms
that the same species was described under these names which, in
accordance with the priority rule, should be called A. carpatica.
Probably, the classical locality (locus classicus) of this species is Pope
Ivan Mountain in the Marmarosha Range of the Soviet Carpathians.
This point was not reported by Willdenow (l. c.), but was indicated by
Kitaibel, which Domin and Krajina indicate in the note on the label to
the exsiccatae issued by them (I. c.).

5. A. ptarmiciformis C. Koch in Linnaea, XXIII (1850) 319;
Boiss. Fl. or. III, 289; Sosn. in Vest. Tifl. Bot. Sada, Ser. 2, 3, 175;
Grossh. Fl. Kavk. IV, 120.—A. iberica var. burgeana Trautv. in Tr.
Peterb. Bot. Sada, II (1873) 546.—A. iberica var. minor Medved.
Rast. Kavk. I, 1 (1919) 260, quoad pl. em. Ararat, non A. iberica
MB.—A. iberica auct. non MB.; Bge. PI. Abich. 11 (1859).—A. argaea
Boiss. and Bal. Diagn. pl. or. II, 6 (1842) 97.

20

21 Plate I.
1—Anthemis saportana Alb., habit, involucre, inner involucral bract, ray floret,
and disk floret, 2—A. iberica MB., habit, involucre, involucral bract, ray floret,
and disk floret. |

24

21

Perennial. Root many-branched, producing short caudices above.
Plants sericeous. Stems short, 5-10 cm high, densely leafy at base,
sparsely above, leafless at apex, pubescent, simple, unicapitulate. Lower
leaves small, oblong, petiolate, pinnate, with linear-oblong, entire or
simple segments, sericeous; uppermost leaves almost entire. Capitula
about 3 cm in dia. Involucral bracts hairy, scarious along margin,
oblong, imbricate, marginal brownish fringe never broad and blackish,
usually sharply demarcated, sometimes so narrow that entire involucre
appears pale brown. Ligules broadly oblong, slightly longer than disk,
3-toothed. Receptacular scales as long as disk florets, cuneate, with
brown border at apex; receptacle slightly concave. Flowering June to
August.

On rocks and screes in alpine zone.—General distribution: Balkans-
Asia Minor (east), Armenia and Kurdistan (north). Described from the
Pontic Range (on protogenic rocks, at about 3,000 m). Type lost.

Note. Found near the border of the USSR on Mt. Ararat. Possibly
growing in the Lesser Caucasus.

According to C. Koch, who described this species, it is close to A.
carpathica but differs in pubescence and from A. montana L., which
belongs to this group, it differs by a broad brownish border of the
involucral bracts.

The identity of the Caucasian plants (from Mt. Ararat) with the
species described by Koch under this name was first proposed by D.I.
Sosnowsky (I. c., 1927); however, to date one cannot have full
confidence in the correctness of this identification, inasmuch as the
authentic specimens of Koch’s species, preserved in Berlin, were
destroyed and not a single specimen of Anthemis from Koch’s
collections exists in the herbaria of the USSR.

Boissier (1. c.) considers A. argaea Boiss. (described from Argei
Mountain or the Ardzhish-Dag in Turkey) as a synonym of this species.
We have seen the latter species with Boissier’s own label. The specimen
of A. argaea examined by us is very similar to Caucasian specimens
identified by Sosnowsky as A. ptarmiciformis. This indirectly confirms
the correctness of Sosnovsky’s identification and leads us to think that
A. ptarmiciformis C. Koch actually grows on Mt. Ararat and possibly
also in neighboring areas of the territory of the USSR.

Series 2. Anatolicae Fed.—Involucral bracts pale, whitish, with
more or less membranous margin and apex. Leaves pinnately cut into
approximate or remote, linear, usually narrow and small lobes of
segments. Capitula usually small; limb of peripheral florets white. Rocky
plants from middle and lower ranges.

Pas

22

6. A. anatolica Boiss. Diagn. pl. or. II, 11 (1849) 10; ej. Fl. or.
III, 294; Grossh. Fl. Kavk. IV, 119.

Perennial. Appressed gray-pubescent, somewhat sericeous plants,
about 10-15(20) cm high. Rhizomes slender, producing strong,
ascending short caudices, bearing leaf rosettes or simple, or weakly
branched stems in lower part. Stems slender, erect, arcuately bent below,
becoming erect toward maturity, leafy in lower half, leafless above,
very finely and inconspicuously longitudinally striate, usually less than
1 mm thick. Lower leaves petiolate, ovate, twice pinnately cut into
small, oblong-linear, obtuse lobes; cauline leaves highly reduced.
Capitula rather small, about 2 cm in dia, one on each stem, with long
and thin peduncles. Involucral bracts weakly hairy, small with very
narrow brownish border, pale, whitish; outer bracts triangular, somewhat
acute; inner oblong, obtuse. Peripheral (ligulate) florets white; limb
oblong, emarginate, scarcely as long as disk. Receptacle conical.
Achenes whitish, sulcate-ribbed, auriculate; outer achenes smaller,
plump, tetraquetrous, tuberculate. Flowering July to August.

In middle mountain zone, in dry stony places.—General
distribution: Balkans-Asia Minor. Described from Asia Minor (“In
mountains and foothills . .. at the foot of Kadm, to the east of the city
of Denizli and between Denizli and Geira’). Type in Geneva.
Discovered and collected long ago in the former Kars Province near
the present border of Turkey with the USSR. Possibly found in the
Lesser Caucasus.

7. A. emiliae Sosn. in Izv. Kavk. Muz. X (1916) 1; Grossh. FI.
Kavk. IV, 120; Sosn. in Fl. Gruzii, VIII, 316.

Perennial. Plants covered with appressed grayish hairs. Root many
headed, very thick, brownish, producing short caudices, bearing sterile
rosettes of leaves and unicapitulate, erect or ascending, simple or
branched stems up to 25 cm high, leafy only in lower part and up to
middle. Lower and basal leaves long-petiolate, obovate, twice pinnately
cut into obovate segments, narrowed as if into petioles and divided
into obtuse small lobes; middle leaves simple pinnate; upper leaves
scaly. Capitula small, up to 1.5 mm in dia, with long peduncles. Outer
bracts of sparsely hairy involucre triangular; inner bracts oblong-linear,
narrow-scarious and blackish along margin, slightly carinate and
brownish along keel. Ligules white, 5-veined, as long as or longer than
disk. Receptacle conical; receptacular scales lanceolate, acute, carinate.
Achenes obpyramidal-tetraquetrous, slightly curved, longitudinally
sulcate, finely tuberculate; corona oblique, serrate. Flowering June to
August.

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23

On rocks in middle mountain zone.—Caucasus: Eastern
Transcaucasia (west). Endemic. Described from Georgia (Tana River
Valley near Ateni). Type in Leningrad.

8. A. sterilis Stev. in Bull. Soc. Nat. Mosc. XXIX, 4 (1856) 379;
Boiss. Fl. or. III, 293; Trautv. in Tr. Peterb. Bot. Sada, VIII, 393;
Schmalh. FI. II, 60; Fedtsch. and Fler. Fl. Evrop. Ross., 968; Stankov-
Taliev, Opred. 381; Rubzsov in Tr. Nikit. Bot. Sada, XXIX, 51.—A.
saxatilis auct. non DC.: MB. FI. taur.-cauc. III (1819) 581, 648.—A.
fruticulosa MB. FI. taur.-cauc. II (1808) 329, quoad pl. tauricam. non
MB. in Tabl. prov. Casp. (1798).—A. fruticulosa auct. non MB.::
Spreng. Syst. Veget. III (1826) 593, quoad pl. Taur.; Ldb. Fl. Ross. II,
521, quoad pl. Taur.; Fedtsch. and Fler. 1. c. quoad nom.—A. montana
auct. non L.: Ldb. FI. Ross. II (1844-1846) 523.

Biennial or perennial. Sericeous, glaucous-gray plant. Root many-
headed, with very short, scarcely visible caudices and non-flowering
shoots bearing crowded rosettes of leaves. Stems slender, hard on
fruiting, straight, short, about 10-15 cm high, simple, unicapitulate,
leafy only below, leafless above almost throughout, arcuately bent,
rather hard and erect, spreading at fruit maturity. Leaves crowded at
base, with thin petiole and lamina, twice or thrice pinnately cut into
few, narrow-linear or oblong-linear segments terminating into corny
cusps. Capitula small, 1.5—2.0 cm in dia, with long peduncles. Involucre
pubescent, pale; outer bracts deltoid, somewhat acute; inner oblong,
with wide scarious margin and apex. Receptacle hemispherical.
Peripheral (ligulate) florets with oblong-elliptical limb as long as disk,
and usually 2-toothed at apex. Achenes somewhat oblong-sulcate,
slightly angular in cross section, with auriculate process on one side;
outer achenes slightly curved. Flowering May to July.

In sterile stony and sandy places near seacoast and along river
banks.—European part: Crimea (southern coast). Endemic. Described
from Crimea (“between the villages of Kapsikhor and Uskut on the
southern coast, in Juniperus excelsa forests, on dry soil,” as well as
“in stony places near Kutlak’’). Type in Helsinki; isotype and paratype
in Leningrad.

Note. This species is often considered to be the one described by
Marschall-Bieberstein as A. saxatilis, but actually Bieberstein
overlooked it at first in Flora tauricocaucasica and lumped it together
with his A. fruticulosa, reporting the latter species not only from the
Caucasus (Kurt-Bulak), as was originally done correctly but also from
Crimea (Kutlak). In volume III (p. 581) of Flora taurico-caucasica,
Bieberstein recognized his mistake and stated: “...habui olim pro
varietate macilenta A. fruticulosae, sed nunc, suadente amiciss.

2

~

24

Steveno, specie separo et cum A. saxatili Decand. conjungo”. In other
words, on the advice of his friend Steven, Bieberstein segregated from
A. fruticulosa its “deparperate” (macilenta) variety growing in Crimea
and included it in the De Candollean species A. saxatilis DC. In doing
this, Bieberstein committed a new, less serious mistake, not realizing
that the “deparuerate” plants represent a totally new species, but ulti-
mately understood, however, as in the very beginning, that the Crimean
plant is not equivalent to the Caucasian one (the actual type of A.
fruticulosa MB., preserved with just such a label and collected from
Kurt-Bulak). It must be remembered that in fact Bieberstein did not
describe any species of his own under the name A. saxatilis. There-
fore, Steven quite consciously described his own A. sterilis and based
it on a new type (see above), citing (as paratypes) the collections of
Bieberstein from Kutlak (i.e., A. saxatilis). At the same time he did not
forget to add that Bieberstein in Flora taurico-caucasica partly had
this species in mind under the name A. fruticulosa, which originally
was quite correctly attributed only to the Caucasian plant. After
Bieberstein, only Ledebour (1. c.) mixed up these two species, having
identified A. saxatilis (sensu Bieberstein) with the entirely different
species A. montana L. without noting that the plants from Crimea do
not belong to the purely Caucasian species A. fruticulosa. After Steven’s
description of A. sterilis and annotations to the diagnosis, it would
seem that the confusion and errors should have disappeared. All this
was correctly shown already by Schmalhausen, but recently, Stankov
(cf., Stankov and Taliev, loc. cit.) again reports both A. sterilis and A.
fruticulosa from Crimea, despite the fact, of course, that nobody has
ever found anything similar to true A. fruticulosa in Crimea.

The new specimens of A. sterilis from the classic locality in Crimea
(village of Morskoe, i.e., Kapsikhor) were collected by N.N. Tzvelev
in 1960.

9. A. tranzscheliana Fed. in Addenda, XXV, 866.

Perennial. Entire plant finely lanate. Root hard, woody, with an
oblong branched caudex, covered with remnants of dead petioles,
producing stems and sterile leaf rosette. Stems highly elongate, 30-45
cm high, simple, unicapitulate, leafy at base, lower part, and up to
middle, leafless above, arcuate at base, straight and slightly sinuate
above, inconspicuously striate longitudinally, not clavate below
capitulum. Lower cauline leaves and those of nonflowering shoots
broadly ovate, divided into linear, sparse, entire or apically 2—3-fid,
divaricately pinnately cut hairy segments; rachis not toothed, narrow.
Capitula medium-sized about 3—4 cm in dia. Outer bracts of more or
less pubescent involucre triangular, pale brown; inner bracts oblong-

28

25

lanceolate, stramineous, with dorsally dark brown midrib, scarious at
apex and sometimes with black border. Ligules white, broadly ovate,
entire, divergent. Receptacle hemispherical; scales dense, stramineous,
membranous, acute. Achenes (immature) brown, subglabrous; lacking
corona or with very narrow apical border. Flowering May to June.

Shady rocks of middle mountain zone, in crevices.—European
part: Crimea (southern coast). Endemic. Described from Crimea
(Karadag Mountain). Type in Leningrad.

Note. This species is close to the Crimean race of A. sterilis Stev.,
but differs from it in being very tall with large capitula and thin,
divided leaves with distant lobes. Large specimens (particularly the
type) are not at all similar to A. sterilis, but smaller plants resemble
this species. However, the disk even of these specimens is always very
large, with a diameter up to 1.5 cm, which is never the case in A.
sterilis—a small and, as mentioned by Steven, “depauperate” plant.
The race described here is endemic especially to Karadag and was
collected only from Karadag (by V.A. Transchel in 1926 and by Al.
Fedorov, An. Fedorov, and M.E. Kirpicznikov in 1952).

10. A. tempskyana Freyn and Sint. in Bull. Herb. Boiss. III, 7
(1895) 347; Sosn. in Vestn. Tifl. Bot. Sada, Ser. 2, 3, 175; Grossh. FI.
Kavk. IV, 120.

Perennial. Semishrubs. Stems woody at base, numerous, about 30—
40 cm long, straight, unicapitulate, terete, with red longitudinal stripes,
leafy below and not above middle; whole plant white-sericeous. Leaves
of nonflowering shoots and lower cauline leaves rather long-petiolate;
lamina pinnate, ovate, divided into few, rather broadly lanceolate,
somewhat acute or obtuse lobes; cauline leaves short-petiolate, upper
leaves sessile, gradually reduced upward, oblong; lobes oblong, small,
obtuse, usually broad. Capitula rather large, 2.5-3.0 cm in dia.
Involucral bracts pubescent, pale, with black border; inner bracts ovate-
lanceolate, outer lanceolate, scarious at apex; all bracts somewhat acute.
Ligules white, oblong-elliptical, apically crenate, two times as long as
disk. Receptacle concave, scales cuneate-oblong, 3-toothed at apex.
Achenes (immature) with very short corona. Flowering June to August.

In open deciduous montane forests.—General distribution: Balkans-
Asia Minor (east). Described from Asia Minor (Giimiisane in Turkey,
in maple forests of Darsosdag Mountain, and also Agridag Range).
Type in Vienna; isotype in Leningrad.

Note. Quite possibly, it occurs within the limits of southwestern
Transcaucasia (on spurs of the Pontic Range and along Chorokh River),
as well as in Armenia. After Sintenis, who collected this species from
Asia Minor, specimens corresponding completely to the type specimen

29

26

were collected by D.I. Litvinow in the former Kars Region in 1914
(mountains above the village of Bazat and the vicinity of the village
of Khan-Dara).

Series 3. Fruticulosae Fed.—Involucral bracts whitish, scarious
along margin and apex. Leaves pinnately cut into narrow lobes. Capitula
small or medium; limb (ligules) of peripheral florets white. Semishrubs,
growing on rocks of middle zones, and partly plains.

11. A. fruticulosa MB. Tabl. Prov. Casp. (1798) 119; ej. Beschr.
Casp. 201; ej. Fl. taur.-cauc. II, 329; quoad pl. cauc.; idem. |. c. III,
p. 580; Spreng. Syst. veget. III, 593, quoad. pl. Cauc.; DC. Prodr. VI,
5; C. Koch in Linnaea, XXIII, 320 quoad nom.; Ldb. FI. Ross. II, 521,
pl. cauc.; Trautv. in Tr. Peterb. Bot. Sada, VIII, 449, p. p. Boiss. Fl.
or. III, 288; Schmalh. Fl. II, 61; Fedtsch. and Fler. Fl. Evrop. Ross.,
968, quoad nom.; Grossh. Fl. Kavk. IV, 120; Sosn. in Fl. Gruzii, VIII,
314; Stankov and Taliev. Opred. 381, quoad nom.

Cultivated. Plants appressed white-tomentose. Root woody, many-
headed, bearing numerous hard but very short, branched and then long
caudices, with brown remnants of dry leaves, passing over to erect,
rather thin, but hard, about 10-15 cm long, basally leafy, unicapitulate
stems; caudices partly covered only with sterile rosettes of leaves.
Basal and rosette leaves usually rather long-petiolate, oblong, divided
into linear or ovate-oblong, very small, obtuse or acute lobes; upper-
most leaves linear, finely lobed. Capitula medium, 2.0-2.5 cm in dia.
Outer bracts of compressed and gray-hairy involucre deltoid-oblong,
somewhat acute, membranous at apex; inner bracts obtuse, with very
broad, scarious, obtuse but incised border. Ligules oblong-elliptical,
white, equaling disk. Receptacular scales linear-oblong, carinate,
truncate. Achenes small, tetraquetrous, with 8 grooves, and very short
indistinctly lobed corona. Flowering May to July (Plate II, Fig. 2).

Dry rocks in middle mountain zone.—Caucasus: Ciscaucasia
(Central Caucasus), Dagestan, Eastern Transcaucasia. Endemic.
Described from the eastern part of the Main Range (near Kurt-Bulak).
Type in Leningrad.

Note. Contrary to some reports (for example, most recently by
Stankov, loc. cit.), it is not found in Crimea, which, incidentally, has
long been known (for details cf. the Note to A. sterilis Stev.).

12. A. trotzkiana Claus ex Bge. Delect. sem. Dorpat. (1847) 3, in
obs. Claus in Mater. k Blizh. Pozn. Prozyabaemosti Ross. Imper. 8, 2,
299; Trautv. in Tr. Peterb. Bot. Sada, VIII, 393; Boiss. Fl. or. III, 287;
Schmalh. Fl. II, 62; O. and B. Fedtsch. Perech. Rast. Turk. IV, 179;

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27

Taliev, Opred., 571; Krascheninn, in Fl. Yugo-Vost. Evrop. Ch. SSSR,
VI, 334.—A. cretacea Zing, in sched., non Zefir.—Pyrethrum
achilleifolium auct. non MB.; Rgl. in Tr. Peterb. Bot. Sada, I (1871-
1872) 254.—Ic.: Krascheninn, op. cit. Fig. 674 (bona).—Exs.: GRF
No. 3260a (topotypus!) and 3260b.

Cultivated plants. Root thick, woody, producing numerous short
caudices, hard, but fragile, dark brown, covered with remnants of dry
leaves; some caudices terminating into sterile leaf rosettes. Stems 10—
15(30) cm high, arcuate at base, straight above, simple or slightly
branched; branches leafless, unicapitulate. Young leaves white-
tomentose, sometimes subglabrous later, slightly fleshy, twice pinnately
cut; segments linear, small, entire or deeply divided, with cartilaginous
cusps; terminal segment pinnately cut. Capitula small, about 2 cm in
dia. Outer involucral bracts ovate, acute, with white-pubescent margin;
inner bracts with hyaline, truncate or entire processes. Receptacle short
conical and apically acute; scales oblong, abruptly acuminate, carinate,
slightly shorter than disk florets. Ligules very broad, yellow. Achenes
obpyramidal, tetraquetrous; corona very small, toothed. Flowering July
to September.

On chalky cliffs and limestone deposits.—European part: Lower
Volga; Soviet Central Asia: Aralo-Caspian Region. Endemic. Described
from “chalky mountains near Khvalinsk’”. Type in Leningrad.

13. A. calcarea Sosn. in Vestn. Tifl. Bot. Sada, XX VII (1913) 10;
Grossh. Fl. Kavk. IV, 120.—Ic.: Sosn. op. cit., Plate 1 (phot., ad
specim. typ.).

Cultivated perennials. Plants 10—20(30) cm high, with spreading
or erect shoots, entirely appressed gray-pubescent. Nonflowering shoots
and few simple or slightly branched, straight or somewhat curved
branches at base arising from woody rhizome. Lower leaves petiolate,
4—6 cm long, oblong, pinnately cut into oblong-lobate segments, with
pinnately parted lobes, with small cartilaginous cusps; cauline leaves
reduced and sessile. Uppermost leaves linear-lobate or filiform. Capitula
rather small, up to 2 cm in dia, terminal on stem or its branches, long-
pedunculate. Involucre white-tomentose; outer bracts somewhat oblong,
acute, slightly membranous at margin; inner bracts oblong-ovate, with
wide membranous and serrate apical process. Ligules white, oblong, as
well as involucre, but exceeding disk. Receptacle convex; scales oblong,
carinate, with 3 apical teeth. Achenes sulcate-ribbed, with small corona.
Flowering June to August.

On limestone rocks, in mountains.—General distribution: Balkans-
Asia Minor (east).

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Described from Olor in Olta District in Turkey (limestone rocks).
Type in Leningrad. Possibly found in the Lesser Caucasus.

Note. The specimen from Olor, the third among those cited in the
diagnosis, should be considered the type, because it is illustrated in
the good photograph appended to the description.

Series 4. Marschallianae Fed.—Involucral bracts pale or brown,
or with blackish border, broadly membranous at tip. Leaves pinnately
cut; lobes narrow, falcately curved. Capitula large; limb (ligules) of
peripheral florets golden-yellow. Plants of high-mountain zone.

14. A. marschalliana Willd. Sp. pl. III, 3 (1804) 2187; MB. FI.
taur.-cauc. II, 332; Spreng. Syst. Veget. III, 595; DC. Prodr. VI, 5;
Ldb. Fl. Ross. II, 520; Schmalh. FI. IJ, 62 p. p.; Sosn. in Vestn. Tifl.
Bot. Sada, Ser. 2, 3, 168; Grossh. Fl. Kavk. IV, 115; Sosn. in FI.
Gruzii, VIII, 312.—A. marschalliana var. typica Trautv. in Tr. SPb.,
Bot. Sada, IV, 1 (1876) 148.—A. biebersteiniana C. Koch in Linnaea,
XXIII (1850) 320; Boiss. Fl. or. III, 286; Schmalh. Fl. II, 62, excl.
var.—A. biebersteiniana o. marschalliana Boiss. op. cit. p. 287; Somm.
and Lev. Enum. pl. Cauc. (1900) 231, 232.—Chrysanthemum
biebersteinianum Adam in Weber and Mohr. Beitr. Naturk. I (1805)
70.—Chamaemelum biebersteinianum C. A. Mey. Verzeichn. (1831)
75.—Exs.: Fl. Cauc. Exs. No. 149; GRF No. 624.

Perennial. Root thick, woody, producing numerous short or long,
thick, brown caudices, densely and imbricately covered with scaly
remnants of dead leaves; some caudices and their branches not
producing stem but sterile rosettes of leaves, resulting in somewhat
caespitose plant. Stems numerous, ascending, unicapitulate, about 10-
15(20) cm high, leafy below. Leaves gray-sericeous; basal leaves
oblong, pinnately cut into linear, approximate, acute segments; cauline
leaves sessile, gradually reduced upward, with fewer lobes. Capitula
about 3 cm in dia. Involucral bracts tomentose outside; inner bracts
with broad and hairy scarious, brownish, but not black margin.
Receptacle concave; scales carinate, apically truncate, almost deltoid.
Ligules golden, as long as disk. Achenes whitish, angular, almost
tetraquetrous, prolonged below, with indistinct or very short corona,
slightly toothed. Flowering June to August. (Plate II, Fig. 1).

On rocky places in alpine zone.—Caucasus: Dagestan, Ciscaucasia
(Main Range), Western Transcaucasia. Endemic. Described from
Central Caucasus (Kaishaur). Type in Berlin; isotype in Leningrad.

Note. Kaishaur should be considered the collecting locality of the
type. About the collecting locality of the type Willdenow (1. c.) said
only: “... Habitat ad Caucasum.” Adam (I. c.), who independently

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29

described this species under a different name, stated that the plant
“occupies crevices in the rocks of Ossetia” (“Occupat rupium fissaras
Ossetiae’). At the same time, on the isotype label written by Adam
and bearing the name Chrysanthemum biebersteinianum mihi it is
specified thus: “... grows on the peak of Kaishaur Mountain” (“...
habitat in summitate montis Kaischaur’). Considering that Willdenow
cited this name as a synonym and referred to the report by Count
Mussin-Puschkin, for whom Adam collected plants at the time of their
joint expedition to the Caucasus, one can be sure that the specimen
sent by Mussin-Puschkin to Willdenow was also collected near
Kaishaur.

i5. A. sosnovskyana Fed. nom. novum.—A. rudolphiana Adam
in Weber and Mohr. Beitr. Naturk. I (1805) 72, nom. illegit. (vide
obs.); MB. FI. taur.-cauc. II, 333; Spreng. Syst. veget. III, 596; C.
Koch in Linnaea, XXIII, 321; Sosn. in Vestn. Tifl. Bot. Sada, Ser. 2,
3, 169.—A. rudolphiana var. pumila Sosn. ex Grossh. Fl. Kavk. IV
(1934) 115; Sosn. in Fl. Gruzii, VIII, 313.—A. marschalliana 6.
rudolphiana C. A. Mey, ex DC. Prodr. VI, (1837) 5; Ldb. Fl. Ross. II,
520; Trautv. in Tr. SPb. Bot. Sada, IV, 2, 376.—A. marschalliana var.
subglabrescens DC. op. cit. 5.—A. biebersteiniana B. rudolphiana C.
A. Mey. ex Boiss. Fl. or. III (1875) 287; Schmalh. Fl. II, 62, excl.
nom. auct.—A. biebersteiniana y. pectinata Boiss. Fl. or. III (1875)
287.—Pyrethrum orientale Willd. Sp. pl. III, 3 (1804) 2159, nom.
prius sed ob homonymiam non restituendum (vide obs.).—P.
xanthomelanum Krasch. ex Grossh. Opred. Rast. Kavk. (1949) 459,
nom. nud.—Chrysanthemum orientale Adam in sched. ad. specim. typ.
and in Willd. |. c. pro. syn.—Exs.: Fl. Cauc. Exs. No. 149 (sub
Anthemide biebersteiniana).

Perennial. Plants often caespitose, consisting of large number of
intertwined, woody, brown, short or long caudices, arising from root,
covered with setiform or scaly remnants of dead leaves and producing
stems, or sterile leaf rosette. Stems usually few, unicapitulate, at base
straight or arcuately ascending, leafy only below, about 10—15(30) cm
long. Leaves often almost glabrous, green, pinnately cut into relatively
short acute lobes; cauline leaves gradually reduced above. Capitula up
to 3 cm in dia; peripheral (ligulate) florets with wide, short-elliptical,
apically distinctly 3-toothed, bright golden-yellow limb. Involucral
bracts membranous on margin and at apex, with blackish-brown broad
border; outer bracts deltoid, inner oblong and distinctly longer.
Receptacular scales carinate. Achenes angular, usually lacking corona.
Flowering June to August.

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On rocks in alpine zone.—Caucasus: Dagestan, Ciscaucasia, Eastern
and Western Transcaucasia. General distribution: Armenia and Kurdistan.
Described from Georgia. Type in Berlin; isotype in Leningrad.

Note 1. Even though Willdenow (1. c.) and Adam (I. c.), who
redescribed this species from the same area but under a different name
and with a different taxonomic status (sub Anthemide, non sub
Pyrethro), mentioned only that the plant “grows in Georgia” (“in
Georgia,” “in Iberia’), it is quite probable, nevertheless, that the exact
collection locality of the type is Kaishaur, where the authentic specimen
of the previous species was collected, and where Anthemis rudolphiana
was subsequently collected by Marschall-Bieberstein (“in Caucasi alpe
Kaischaur’). The specimen from Adam’s herbarium (isotype) does not
mention the place of collection, and there are no other indications.
Therefore, it is quite possible that it was indeed collected near Kaishaur
(preserved in the herbarium together with the type specimen of
Anthemidis marschallianae [sic] and it was considered unnecessary to
make separate mention of the collecting site).

Note 2. Restoration of the original Willdenowian specific epithet
“orientalis” is not possible because the binomial name Anthemis
orientalis (L.) Degen, based on the earlier basionym Anacyclus
orientalis L., was already in use. But since the name “rudolphiana”
was superfluous at the time of its publication, then in accordance with
the International Code of Nomenclature, the name of the author (Adam)
should be dropped henceforth after the species name Anthemis
rudolphiana, or a new name should be given to this species. We prefer
to call it by the new name A. sosnovskyana, in honor of Prof. D.1.
Sosnowsky, who contributed much toward an understanding of the
systematics of the genus Anthemis. The recent (Grossheim, 1949, 1. c.)
name, Pyrethrum xanthomelanum Krasch. ex Grossh., which was used
erroneously to designate a certain specimen of A. sosnovskyana, can-
not be used as basionym for a new combination, because the name P.
xanthomelanum is invalid, being only a nomen nudum.

Section 2. Anthemis—Sect. Chamaemelum Cass. in Dict. Sc. Nat.
XXIX (1823) 279 p. p.—Sect. Euanthemis ser. Annuae Boiss. FI. or. II
(1875) 276 p. p.—Achenes obtuse or truncate at apex, with or without
auriculate corona, not compressed. Receptacular scales oblong or
lanceolate. Annual or biennial herbs, with highly divided pinnate leaves
and usually branched stems. Capitula often lacking ligulate florets.

Type of section: Lectotype of genus.

Series 1. Arvenses Fed.—Inner involucral bracts obtuse, with
membranous margin. Stems 20-30(40) cm high branched. Pubescence
appressed or somewhat squarrose, velutinous.

31

33 Plate II.
1—Anthemis marshalliana Willd., habit, involucre, inner involucral bract, ray
(ligulate) floret, and disk floret; 2—A. fruticulosa MB., habit, involucre, inner
involucral bract, ray (ligulate), and disk floret.

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32

16. A. arvensis L. Sp. pl. II (1753) 894; DC. Prodr. VI, 6; MB.
Fl. taur.-cauc. III, 582; Ldb. Fl. Ross. II, 521; Boiss. Fl. or. II, 301;
Schmalh. Fl. II, 61, excl. var.; Fedtsch. and Fler. Fl. 968, excl. var.;
Grossh. Fl. Kavk. IV, 119; Fl. Yugo-Vost. VI, 334; B. Fedtsch. Rast.
Turk. 735.—A. multicaulis Boiss. and Heldr. in Boiss. 1. c. pro syn.—
Ic.: Fedtsch. and Fler. Fl. 968; Fl. Yugo-Vost. VI, 460, 459, 15
(achene).—Exs.: Pl. Finn. Exs. No. 976; Herb. Fl. Ingriae, No. 311;
Fl. Polon. Exs. No. 641.

Annual. Appressed-pubescent, slightly ash-gray plants. Stems
numerous, arising from single root, branched, 15—30(40) cm long.
Leaves oblong, pinnately cut into narrow, linear-cuneate, acute
segments. Peduncles of inflorescences oblong, somewhat clavate.
Involucre hispid; bracts ovate-oblong, outer acute, others obtuse,
broadly scarious on margin and apex. Receptacle oblong-conical; scales
lanceolate, carinate, abruptly cuspidate, exceeding disk florets. Capitula
2-3 cm in dia; ligules of peripheral, exclusively pistillate, florets
elliptical; tube of disk florets swollen at base. Achenes somewhat
tetraquetrous, longitudinally sulcate, smooth; outer achenes somewhat
plump, inner with short acute border at apex in place of truncate corona.
Flowering May to September.

Long dry herb-covered places, fields—European part: Karelia-
Lapland (south), Ladoga-IImen, Baltic Region, Upper Volga, Upper
Dnieper, Volga-Don, Middle Dnieper, Upper Dniester, Bessarabia,
Black Sea Region, Lower Don, Crimea; Caucasus: Ciscaucasia, Western
Transcaucasia. General distribution: Scandinavia, Atlantic Europe,
Central Europe, Mediterranean Region, Balkans-Asia Minor. Described
from “Europe, especially from fields of Sweden” (“Europae, praesertim
Sueciae agris”). Type in London.

17. A. ruthenica MB. FI. taur.-cauc. II (1808) 330 and 465; DC.
Prodr. VI, 11; Ldb. Fl. Ross. I, 522; C. Koch in Linnaea, XXIII, 315;
Boiss. Fl. or. III, 304; Grossh. Fl. Kavk. IV, 119; Krascheninn, in FI.
Yugo-Vost. Evrop. Ch. SSSR, VI, 335.—A. maritima Pall. ex MB.
l. c. pro syn. A. maritima L. non A. arvensis B. ruthenica Schmalh. FI.
II (1887) 61; Fedtsch. and Fler. Fl. 968.—A. pontica Ldb. op. cit. p.
5.—A. pontica auct. non Willd.: d’Urville. Enumerat. (1822) 114.—A.
pontica d’Urville in Ldb. |. c. pro syn.—A. neilreichii Ortmann in
Flora (1852) 672.—Ic.: Javorka-Soo, Néven kiz. t. 146, f. 884
(mediocr.).—Exs.: GRF No. 733; Lang. and Szovitz, Herb. Ruthen.
Cent. I, No. 8; Schultz. Herb. normale, No. 504 (sub A. neilreichii);
Dorfler, Herb. normale, No. 5275.

Annual. Tomentose-lanate, many-stemmed plants, highly branched
from base. Stems short, 15-20 cm high, with straight, divergent,

cH

33

divaricate branching. Leaves oblong, divided into linear or oblong-
lobate segments with deltoid acute lobes. Peduncles of capitula long,
but not clavate below capitulum; capitula about 2 cm in dia. Involucre
hispid; bracts pale; outer bracts acute, inner oblong and obtuse, with
wide scarious margin. Receptacle conical; scales lanceolate, acute,
membranous except midrib, equaling disk. Ligulate florets only
pistillate; ligules white, elliptical, tube of disk florets swollen at base.
Achenes smooth, obconical; outer achenes deeply sulcate-angular, inner
sulcate, all truncate, very narrowly and usually indistinctly bordered at
tip, i.e., almost lacking corona. Flowering May to July.

In steppe and partly forest-steppe zone.—European part: Upper
Volga, Upper Dnieper, Bessarabia, Middle Dnieper, Volga-Don, Trans-
Volga Region, Black Sea Region, Crimea; Caucasus: Ciscaucasia,
Dagestan, Eastern and Western Transcaucasia. Described from Crimea
and Ukraine. Type could not be located.

Series 2. Candidissimae Fed.—All involucral bracts somewhat
acute. Stems short, (5)10-15 cm high, divaricate from base, simple or
branched. Pubescence dense silvery-tomentose and somewhat squarrose
or appressed.

18. A. candidissima Willd. ex Spreng. Syst. nat. III (1826) 593;
DC. Prodr. VI, 9; Boiss. Fl. or. III, 305; Ldb. Fl. Ross. II, 523; Grossh.
Fl. Kavk. IV, 119; Sosn. in Fl. Gruzii, VIII, 317; B. Fedtsch. Rast.
Turk. 735(?).—A. candidissima var. epapposa and intermedia Trautv.
in Tr. Peterb. Bot. Sada, IV (1876) 337.—A. pterygantha C. Koch in
Linnaea, XVII (1843) 44; Ldb. Fl. Ross. II, 522.—A. peregrina auct.
non L.; MB. FI. taur.-cauc. II (1808) 331.—A. chamamilloides Stev.
ex MB. Il. c. pro syn.; C. Koch in Linnaea, XXIII, 316.—Ic.: Takht.
and Fedor. Fl. Erevana, Atlas, Plate 107 (mediocr.).

Annual. Entirely silvery-tomentose plants, with roots simple or
branching from collar. Stems shorts, 5—10(15) cm high with erect
branches. Leaves short-ovate, 2—3-pinnatisect into narrow-linear, acute
segments; lobes ternate or palmately divided, approximate. Peduncles
of inflorescence slender, long, but not clavate; capitula small, 1.5—2.0
cm in dia. Involucre lanate, bracts lanceolate, acute. Receptacle conical;
scales narrow linear-lanceolate, carinate, acuminate, as long as disk
florets. Corolla tube swollen at base. Achenes small, roundish in cross
section or indistinctly tetraquetrous, slightly tuberculate along ribs on
outside; all achenes truncate at apex, lacking corona or with acute and
very narrow apical border, sometimes nonuniformly developed.
Flowering May to June. (Plate IV, Fig. 2).

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In semidesert region.—Caucasus: Eastern Transcaucasia, South-
ern Transcaucasia; Soviet Central Asia: Kara-Kum, Kyzyl-Kum. De-
scribed from Caucasus (Georgia). Type in Berlin; isotype in Leningrad.

Note. On the basis of Willdenow’s herbarium, Sprengel (1. c.)
described under the name A. candidissima undoubtedly a duplicate of
the same plant which was found also in the herbarium of Marschall-
Bieberstein (collections of Adam) and whose specimens (isotypes) were
sent to Willdenow, probably, by Mussin-Puschkin. However, Willdenow
failed to establish a new species based on them in “Species plantarum,”
which was subsequently done by Sprengel. Marschall-Bieberstein
himself considered his plants identical with the western (southern
European) species A. peregrina, which is why Sprengel cited this name
in his diagnosis as a synonym. Thus, there is sufficient reason to
consider the specimen from Georgia (Ex Caucaso Iberico), preserved
in Bieberstein’s collection, as an isotype of A. candidissima.

We have cited the synonym A. pterygantha C. Koch relying on
Boissier’s authority, who identified the Koch’s plants with A.
candidissima. Unfortunately, many species described by Koch remained
poorly explained, until almost all the authentic specimens perished.
Koch’s original diagnoses have long provoked justified criticisms for
their poor and sometimes defective description (the exact collecting
site and other details are not mentioned).

19. A. grossheimii Sosn. in Vestn. Tifl. Bot. Sada, Ser. 2, 3 (1927)
176; Grossh. Fl. Kavk. IV, 119; Sosn. in Takht. and Fed. Fl. Erevana,
307.—A. candidissima var. longepapposa Trautv. in Tr. Peterb. Bot.
Sada, IV (1876) 337.

Annual. Plants appressed-hairy, grayish-green or grayish, with
numerous, short, up to 8 cm high, branched, erect stems, arising from
root collar. Leaves ovate-oblong, thrice pinnately cut into short, fine,
filiform, divaricate lobes, with hyaline cusp. Capitular peduncles long,
later somewhat thickened. Involucre appressed-hairy; bracts pale
brownish along margin, membranous, green outside; all bracts
lanceolate, with acute and emarginate apex. Receptacle conical; scales
oblong-lanceolate, carinate, apically truncate, serrate, with cusp, as
long as florets. Peripheral florets white, fertile, with elliptical ligules
as long as disk; tubes of all florets not swollen, pale-ocher-yellow.
Outer achenes small, terete, prominently ribbed and tuberculate, plump,
lacking corona, achenes of disk florets narrow-obpyramidal, prominently
ribbed, less tuberculate, with serrate-incised auricles, thrice as long as
achenes. Flowering May to June.

In semideserts.—Caucasus: Southern Transcaucasia. Endemic.
Described from Armenia (Yerevan, on stony slopes). Type in Tbilisi.

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35

Note. According to D.I. Sosnowsky (I. c.), this species is close to
A. haussknechtiana Boiss. and Reut., but differs in having pale yellow
and not swollen corolla tubes.

20. A. hirtella Winkl. in Tr. SPb. Bot. Sada, XI, 2 (1892) 278; O.
and B. Fedtsch. Perech. Rast. Turk., IV, 179; Zakirov in Tr. Uzbeksk.
Univ. Nov. Ser. No. 89, 147.

Annual. Plants entirely grayish-tomentose, 10-15 cm high, with
branched stems; branches leafy up to middle, but always bearing solitary
capitula and leafless peduncles of inflorescence. Leaves 2—3 cm long,
twice pinnately divided; lobes ovate-lanceolate, somewhat acute.
Peduncles of inflorescence clavate below capitulum; capitula somewhat
large, hemispherical, about 12 mm in dia. Receptacle conical, punctate;
receptacular scales lobate, membranous, but split toward apex, with
acute cusp. Involucral bracts numerous; outer bracts leafy, broadly
membranous along margin; lanate-hairy outside, broadly lanceolate,
acute; inner bracts slightly narrowed, gradually becoming membranous
(inward); innermost bracts similar to receptacular scales. Peripheral
florets pistillate, ligulate; ligules somewhat broad, oblong-roundish,
longer than involucre, white, with 3 subobtuse teeth; corolla tube
swollen, lanate outside; disk florets yellow, tubular; tube constricted in
middle (resembling sand clock in shape), with 5 apical teeth. Anthers
very short-sagittate; style branches short, almost erect, truncate,
clustered. Achenes turbinate, with longitudinal striations, lacking
corona. Flowering April.

In desert region.—Soviet Central Asia: Pamiro-Alai. Endemic.
Described from Tadzhikistan (eastern Bukhara, Karatau Range, at source
of Chiglin River). Type in Leningrad.

Note. It is close to A. hemistephana Boiss., but its achenes lack
a corona, whereas the latter species has a unilateral corona.

Actually, Winkler’s article (Decas septima ...) was published in
1892, but 1890 is mentioned on the title page of the reprints, which
can be considered as the time of publication of A. hirtella.

Section 3. Cota (J. Gay ex Guss.) Rupr. Fl. Ingr. (1860) 589;
Boiss. Fl. or. II], 278.—Sect. Euanthemis DC. Prodr. VI (1837) 7
p.p.—Cota J. Gay in Guss. Fl. Sic. Syn. II (1842-1844) 866, pro
gen.—Achenes tetraquetrous, with somewhat sharp corners, usually
highly compressed. Receptacular scales carinate. Stems usually
sparingly branched; capitula often large. Perennial or biennial plants
with pinnate leaves; leaf lobes somewhat broad.

Type of section: A. tinctoria L.

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36

Series 1. Tinctoriae Fed.—Involucral bracts pale; peripheral (ligu-
late) florets golden-yellow. Stem branched or simple. Leaves grayish
from pubescence or green and subglabrous. Perennials.

21. A. tinctoria L. Sp. pl. II (1753) 896; Ldb. Fl. Ross. II, 524,
excl. var.; Boiss. Fl. or. III, 280 p. p. excl. var. and syn.; Schmalh. FI.
II, 61 p. p., excl. vat.; Fedtsch. and Fler. 967 p. p.; Sosn. in Fl. Gruzii,
VIII, 306, quoad nom.; Grossh. Fl. Kavk. IV, 116, excl. syn. quoad
nom.; Kryl. Fl. Zap. Sib. XI, 2720, quoad nom.—Ic.: Syreistsch. FI.
III, 260 (mediocr.).—Exs.: GRF No. 3700; Fl. Finl. Exs. No. 975; Fl.
Polon. Exs. No. 352a and b; Herb. Fl. Ingr. No. 312; Pabo and
Tscholowskii, Herb. Mohil. XIX, 2.

Perennial. Plants usually with many stems. Stems virgate, especially
at fruiting, strong, 30-40(60) cm high, reddish, often cymosely and
divaricately branched. Leaves appressed-hairy, greenish (not gray),
oblong, pinnately divided into oblong, usually folded and compressed
segments, with deltoid-lanceolate, acute, pectinate lobes, terminating
into white cusp; leaf axes serrate. Capitula rather large, up to 3 cm in
dia, with long peduncles. Involucre subglabrous, bracts pale; outer
bracts lanceolate, more or less acute; inner oblong-linear, with obtuse,
scarious apex. Ligules yellow, shorter than disk. Receptacle
hemispherical; scales broadly linear, carinate, acuminate almost as long
as disk florets. Achenes rectangular-compressed, with many grooves
on both sides and very short, almost entire, corona at tip. Flowering
June to September.

In dry meadows, fields and weedy places.—European part: Karelia-
Lapland (south), Ladoga-Ilmen, Baltic Region, Upper Volga, Dvina-
Pechora (south), Volga-Kama. General distribution: Scandinavia,
Central Europe, Atlantic Europe, Mediterranean Region (north).
Described from “dry sunny meadows of Sweden and Germany.” Type
in London.

Note. The northern (especially Swedish) plants (topotypes) should
be considered as typical specimens of A. tinctoria, which are
characterized by highly branched stems with many capitula. In this
northern race, the ligules are intensely golden-yellow (not pale), the
leaves green, and the stems distinctly reddish. Another race, A.
subtinctoria Dobrocz., grows in the south, beginning from the Ukraine
and southeast, as well as in Crimea and the Caucasus, in addition to
a whole series of endemic races with small distribution ranges. A.
subtinctoria differs quite sharply from typical A. tinctoria by having
grayish pubescence, narrow leaf segments, and pale ligules.

A hybrid, A. tinctoria x A. cotula, was found in the former Venev
District in the former Tula Province, which was identified as A. bollei

4

—

37

Sch.-Bip. We have not included this form among the species of
Anthemis, as hybridization apparently is a common feature among its
species, but the degree of constancy of the hybrids is unknown. As
regards the name A. bollei, it is hardly suitable in the present case. The
Venev hybrid probably appeared spontaneously and entirely
independently of the hybrid form described from Western Europe under
this name. Concerning A. bollei from Venev, see Majevsky “Fl. Sred.
Russ.” [Flora of Central Russia], Ed. 5 (1917) 301, edited by D.
Litvinow, where this form was first reported on the basis of the
published notes and herbarium of V.V. Rozen who collected it in 1912
in the locality mentioned above.

The characteristics of the Venev hybrid, according to Litvinow
(1. c.), are as follows: “Outer florets yellowish-white. Stem erect, quite
branched, sparsely pubescent (like leaves). Leaves 2-pinnatipartite;
lobes pectinate, linear or linear-lanceolate, with entire or dentate margin
or with two or three notches; lobes of first order close together. Capitula
long-pedunculate. Involucral bracts obtuse, somewhat oblong.
Receptacular scales lanceolate-linear, with prickly cusp. Annual, 35-—
45.

According to A.A. Grossheim (Fl. Kavk., p. 121), the hybrid form,
also called A. bollei by him, was found in the northern Caucasus.

22. A. subtinctoria Dobrocz. in Ukr. Bot. Zhurn. XVIII, 2 (1961)
67, Fig. 2.—A. caucasica Henckel, Adumbrat. plant. nonnull. Horti
Halensis Acad. Select. (1806) 9, non Adam (1805).—A. kochiana Sosn.
ex Grossh. Fl. Kavk. IV (1934) 116, pro syn.—A. chrysantha Sosn. ex
Grossh. 1. c. pro syn. and in Opred. Rast. Kavk. (1949) 452, non
Gay.—A. tinctoria var. chrysantha Trautv. in Tr. Bot. Sada, IV (1876)
377.—A. tinctoria B. macro-glossa C. Koch in Linnaea, XXIII (1850)
318.—A. tinctoria B. floribus pallidis C. Koch op. cit. XVII (1843)
44.—A. tinctoria auct. non L.: Ldb. Fl. Alt. I1V, 122; B. Fedtsch. Rast.
Turk. 735; Kryl. Fl. Zap. Sib. XI, 2720; M. Popov, FI. Sr. Sib. II, 725;
Katina in Vizn. Rosl. URSR, 536; Kraschennin, in Fl. Yugo-Vost.
Evrop. Ch. SSSR, VI, 334.—A. pallescens auct. non Heldr.: Trautv.
in Tr. Bot. Sada, VIII, 391.—Exs.: Callier, Iter. taur. (1900) Nos.
631-632 (sub A. finct.).

Perennial. Plants grayish from more or less dense and somewhat
appressed hairs. Stems 30—50(70) cm high, branched usually in lower
part, less often simple, often not spreading. Leaves oblong-ovate or
oblong, divided into narrow linear segments, with few divergently sharp-
toothed cuspidate lobes. Capitula medium, up to 2—3 cm in dia, less
often larger; ligules yellow, sometimes pale yellow, oblong-lanceolate,
narrowed at apex and with 3 teeth, equal to disk or larger. Involucre

42

38

hairy; outer bracts oblong-deltoid, acute; inner bracts oblong-lanceolate,
somewhat obtuse, lacking fimbriate apex; all bracts pale, lacking green
or brownish vein. Receptacle hemispherical; scales brownish, oblong,
abruptly narrowed into darker cusp. Flowering June to August.

European part: Black Sea Region, Lower Don, Volga-Don, Trans-
Volga Region, Crimea; Caucasus: All regions; Siberia: Upper Tobol,
Irtysh, Angara-Sayans (rarely); Soviet Central Asia: Lake Balkhash
Region. Endemic. Described from the Black Sea Region of Ukraine
(Stalino Region [now Donetsk Region—Ed.], Novoazovsk Region,
“Khomutovskaya Steppe” Reserve). Type in Kiev.

23. A. maris-nigri Fed. Addenda, XXV, 866.

Perennial. Plants sparsely covered with short or longer hairs, but
green or slightly grayish. Stems arising from somewhat woody root,
arcuate below, erect above, with spreading branches, about 40—60(70)
cm high, strong, longitudinally striped, leafy up to middle and above,
leafless near tip. Basal leaves; caducous; cauline leaves obovate,
somewhat wide, pinnate, divided into somewhat obtuse and rather wide
oblong-linear, sharply toothed segments, teeth uneven, pointed forward,
with very small cusps. Capitula rather small, about 2 cm in dia; ligules
yellow, short, obtuse, indistinctly 3-toothed at apex, almost a half as
long as disk. Involucre lanate; outer bracts deltoid, sharply acute; inner
bracts oblong-lanceolate, somewhat obtuse, all bracts blackish on
midrib, brownish-ribbed at apex. Receptacle hemispherical; scales
brown, oblong, abruptly narrowed into an obtuse cusp. Achenes
(immature) stramineous, indistinctly tetraquetrous, with indistinct teeth
at apex, glabrous. Flowering June to September.

In montane forest zone.—Caucasus: Western Transcaucasia
(Kolkhida). Endemic. Described from cliffs between Nakhe and
Chukhuk rivers in Sochi District. Type in Leningrad.

24. A. saguramica Sosn. in Vestn. Tifl. Bot. Sada, Ser. 2, 3 (1927)
150; Sosn. in Sosn. and Grossh. Opred. Rast. Tifl. (1920) 284, nom.
nud.; Grossh. Fl. Kavk. IV, 116; Sosn. in Fl. Gruzii, VIII, 305.

Perennial. 40-50(70) cm high plants, sometimes with nonflowering
shoots at base. Stems erect, branched in upper half, spreading, angular,
striate at base, subglabrous or grayish from scattered curly hairs. Lower
cauline leaves early falling, petiolate, spatulate, pinnately cut; segments
oblong, along margin sharply toothed or bipinnate; middle leaves sessile,
ovate, pinnate; segments gradually reduced toward leaf base, lowermost
segments amplexicaul, somewhat broadly spatulate, divided to middle
into short, oblong, sharply acute, toothed lobes; all leaves greenish,
glaucous beneath, gradually reduced upward; some upper leaves scaly,

43

39

pinnately toothed. Capitula on very long, subglabrous peduncle,
medium, about 2.0—2.5 cm in dia. Involucral bracts linear-lanceolate,
acute, white-pubescent, narrow membranous at apex. Liguies golden,
oblong, as long as disk, with small teeth at apex. Receptacle
hemispherical; scales narrow and sharply acute, brownish, indistinctly
carinate outside. Flowering June to September.

In middle mountain zone.—Caucasus: Eastern Transcaucasia
(west). Endemic. Described from Georgia (Sagurami). Type in Tbilisi;
autotypes and topotypes in Leningrad.

25. A. markhotensis Fed. in Addenda, XXV, 876.

Perennial. Ali plants sericeous-lanate from dense hairs. Stems
branched and leafy in middle, about 40-50(60) cm high, leafless at
base and below capitula, longitudinally striped, reddish. Leaves dense,
rather large, about 3—4 cm long, obovate, pinnate; segments somewhat
wide, oblong-obovate; lobes truncate obconical-dentate. Capitula rather
large, about 2—2.5 cm in dia; ligules yellow, short, roundish at apex,
entire or sometimes indistinctly toothed, oblong-ovate, distinctly shorter :
than disk. Involucre lanate; outer bracts deltoid, acuminate, inner bracts
oblong, somewhat obtuse; all bracts pale. Receptacle spherical; scales
brown; outer scales gradually narrowed into cusp, inner abruptly
narrowed into emerginate cusp or bilobate tip. Achenes stramineous,
glabrous, slightly ribbed, unevenly and indistinctly toothed at apex.
Flowering June to September.

On cliffs of coastal limestone rocks and partly on herb slopes in
calcareous soil.—Caucasus: Western Transcaucasia (western part).
Endemic. Described from Markhot Range. Type and paratypes in
Leningrad.

26. A. monantha Willd. Sp. pl. III, 3 (1804) 2187; Zefirov in
Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI, 370-372, XVIII,
253, 254 p. p.; N. Rubzov in Tr. Nikitsk. Bot. Sada, XXIX, 50, in
obs.—A. coarctata auct. not Sibth. and Sm.: Trautv. in Tr. Peterb.
Bot. Sada, IX (1884) 391.—A. tinctoria var. B. MB. FI. taur.-cauc. II
(1808) 332.—A. tinctoria var. monantha DC. Prodr. VI (1837) 11;
Ldb. FI. Ross. II, 524.—A. tinctoria var. uniflora MB. in sched. (1810).

Perennial. Stems 1-2, simple, about 25-30(40) cm high,
unicapitulate, sparsely pubescent like leaves, rather slender, erect,
somewhat arcuate at base, generally leafy only below, less often up to
middle, leafless above. Leaves small, somewhat wide, up to 1.5—2.0
cm long, pinnate; segments linear, divergent, finely and sharply incised-
toothed, greenish (not gray). Capitula medium, about 2.0-2.5 cm in
dia, with narrow-oblong or lanceolate, bright yellow ligules, lacking

ot

40

apical teeth. Involucre slightly hairy; outer bracts acute-deltoid, lack-
ing greenish veins; inner bracts oblong, somewhat obtuse. Receptacular
scales oblong, brownish, terminating into long cusp. Flowering June to
August.

In mountains, on herb slopes.—European part: Crimea. Endemic.
Described from Crimea. Type in Berlin.

Note. B.M. Zefirov, who for the first time established this species
for the Crimean flora after a long period of oblivion, understood it in
the broad sense, including the yellow chamomile of the affinity of
Anthemis tinctoria s. |., which very often has tall, branched stems, and
grayish leaves with narrow lobes and is the most widely distributed
race in the south of the European USSR, especially in Crimea and the
Caucasus. However, D.N. Dobroczaeva (personal communication) quite
correctly segregates this race as a separate species, A. subtinctoria
(cf.), assigning the name A. monantha Willd. only for the Crimean
montane yellow-flowered chamomile, which always has simple
unicapitulate stems and greenish (not gray) leaves with thin pubes-
cence not imparting the typical grayish appearance to the plant as

‘found in A. subtinctoria.

27. A. cretacea Zefir. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVI (1954) 371; N. Rubzov in Tr. Nikitsk. Bot. Sada, XXIX,
50.

Perennial. Stems numerous, ascending, partly prostrate, highly
branched from base, 10-20 cm high, finely tomentose, leaves almost
glabrous above, lanate-tomentose beneath, ovate, 10-15 mm long,
pinnately cut into 2-5 narrow-lanceolate segments on each side, further
divided into oblong-lanceolate, acute lobes. Capitula solitary terminal
on branches, 1.2—1.8 cm in dia. Outer bracts of arachnoid involucre
lanceolate, green, with pale border on margin; inner bracts obtuse,
with fimbriate apices, brownish. Ligules bright yellow, slightly shorter
than disk. Receptacle almost hemispherical; scales shortly acute.
Achenes compressed-tetraquetrous, slightly curved, with unequally
toothed and very short border at tip. Flowering June to August.

On chalk slopes, limestone deposits, in middle and lower mountain
zone.—European part: Crimea; Caucasus: Western Transcaucasia
(west). Endemic. Described from Crimea (Mangub-Kale, chalky scree).
Type in Leningrad.

Note. At first glance, the holotype of this species gives the im-
pression of a small specimen of the common A. ftinctoria (or A.
subtinctoria) that grows in poor soil and is also grazed by cattle.
However, we collected an Anthemis in 1952 near Gelendzhik in the
Caucasus from limestone rocks that is very similar to A. cretacea, but

45

41

with simple, and not branched, unicapitulate stems. These specimens
can all be referred to A. cretacea evidently, and at the same time, it
can be suggested that the branching of the type specimen and the large
number of small capitula on the lateral shoots is the result of cattle
grazing, causing the regeneration of branches. Normally, A. cretacea
is a plant with unbranched stems, each bearing a single capitulum.

28. A. debilis Fed. in Addenda, XXV, 867.

Perennial. Plants grayish from short and rather dense hairs. Stems
arising from hard brownish root, ascending and weak, sparingly
branched or simple, rather slender, about 15—20(40) cm high, leafy up
to middle, leafless above. Leaves only cauline, dense, reduced, small,
about 7—10(15) mm long, broadly ovate, pinnate, with linear, acute,
truncate-toothed approximate segments in turn divided into acute lobes
with small cusps. Capitula medium or small, about 2.5 cm in dia;
ligules golden gray, sometimes divergent, oblong-lanceolate, with
indistinct apical teeth. Involucre lanate; all bracts pale; outer bracts
deltoid, greenish on midrib; inner bracts oblong-lanceolate, somewhat
obtuse, more densely tomentose at tip. Flowering May to June.

On rocky cliffs and other similar sterile places —Caucasus: Western
Transcaucasia (south, occurrence possible). General distribution:
Balkans-Asia Minor (east). Described from Asia Minor (Artvin District,
Godrokhevi Ravine); type and paratypes in Leningrad.

Note. One might think that this species had already been described
earlier by C. Koch as A. brachyglossa C. Koch. However, Koch’s
diagnosis is so poor that there can be no certainty in such assumption.
Besides, the type specimen, on the basis of which Koch described his
plant, perished irretrievably like the other authentic specimens preserved
in Berlin after Koch’s death. The other assumption about the possible
identity of A. debilis m. and A. brachyglossa C. Koch is based only on
Koch’s report that his species is found in the Chorokh District [“... im
Tschorukgebiete”. C. Koch in Linnaea, XXIII (1850) 318]. In order to
simplify the further attempts at elucidation of A. brachyglossa C. Koch,
a translation of the original diagnosis from Koch’s publication is given
here: “Perennial (plant), gray due to dense appressed pubescence. Stems
branched at base, ascending, 1/2 foot or little taller, with almost simple
branches, unicapitulate, leafless above. Leaves ovate-oblong, 5-7 mm
long, pinnatipartite with oblong lobes, incised-dentate, with very sharp
teeth. Capitula hemispherical. Involucral bracts imbricate, tomentose.
Ligules golden, highly reduced, later recurved downward. Scales
(receptacular) oblong, abruptly narrowed into cusp, equaling disk
florets. Achenes distinctly bordered laterally, with 3 stripes on dorsal
and ventral sides, glabrous, with very short corona.”

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42

29. A. zephyrovii Dobrocz. in Ukr. Bot. Zhurn. XVIII, 2 (1961)
70, Fig. 1—A. monatha var. orientalis Zefir. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XVIII (1957) 253-254, absque diagn. lat.

Perennial. Root strong, woody, thick, bearing short, erect, spreading
stems. Stem leafless below, but with remnants of dry leaf petioles,
densely leafy in middle, rather thick, up to 2 mm or more in dia,
slightly reddish, and like leaves rather densely grayish-pubescent, simple
or weakly branched, with solitary terminal capitula on each branch or
stem. Leaves rather large, up to 3-4 cm long, grayish-pubescent, broadly
ovate, pinnately cut into linear segments, sharply incised-serrate along
margin and remote. Capitula medium or rather small, with narrow yellow
ligules. Involucre lanate; outer bracts acute-deltoid; inner bracts oblong,
somewhat acute, lacking apical fringe and greenish midrib. Flowering
June to August.

On stony slopes, in foothills, sometimes in oak forests —European
part: Crimea (Kerch Peninsula). Described from the above mentioned
region (near the village of Marfovka). Type in Leningrad.

Note. B.N. Zefirov (1. c.), who first discovered this species but
described it as a variety (of A. monantha), noted its characteristic
feature: “Stem . . . often branching already from middle; leaves oblong
in their general outline, noticeably exceeding very short internodes.”
The range of this species covers only the eastern part of Crimea (Kerch
District, Feodosia), spreading along the southern coast not farther west
than Karadag and Sudak.

Series 2. Rigescentes Fed.—Involucral bracts greenish or whitish,
lacking dark broader; peripheral (ligulate) florets white. Stems slightly
branched or simple. Leaves green or grayish, with sparse or dense
pubescence, or glabrous. Perennials.

30. A. rigescens Willd. Hort. Berol. I (1816) 75; Boiss. FI. or. III,
28 p. p.; Fedtsch. Rast. Turk. 735, quoad nom.; Sosn. in Vestn. Tifl.
Bot. Sada, Ser. 2, 3, 158, diagn. lat. compl.; Fl. Gruzii, VIII, 308.—
A. rigescens vat. latifolia Sosn. ex Grossh. Fl. Kavk. IV (1934) 118.—
A. rigescens var. genuina C. Koch in Linnaea XXIII (1850) 317.—A.
caucasica Hornem. Hort. Hafn. II (1813) 832, non Henk. (1806), nec
Adam (1805).—A. triumfettii auct. non All.: Ldb. Fi. Ross. II (1844—
1846) 525, p. max. p.—A. tinctoria var. triumfettii Trautv. in Tr. Bot.
Sada, IV (1876) 337, non A. triumfettii All.—A. tinctoria var. rigescens
Schmalh. Fl. II (1897) 62 p. p.—Ic.: Willd. loc. cit. t. 62 (bona).

Perennial. Sparsely appressed-hairy or almost glabrous, green
plants. Stems short, 40—50(70) cm high, simple or cymosely branched,
erect. Leaves broadly ovate-oblong, pinnately cut into oblong-lanceolate

47

43

segments, divided into deltoid pale-acuminate and usually also toothed-
pectinate lobes, with a broad, toothed, rachis. Capitula long-pedunculate,
usually large, up to 3 cm in dia. Involucral bracts pubescent or sub-
glabrous; outer bracts deltoid-lanceolate, somewhat acute; inner bracts
oblong-linear, obtuse, with scarious apex, pale or slightly blackish,
somewhat fimbriate along margin. Ligules white, as long as disk or
longer. Receptacle hemispherical; scales oblong, carinate, terminating
into cusp as long as disk florets. Achenes tetraquetrous-compressed,
narrow-winged, with numerous stripes on both sides and short corona.
Flowering July to August.

In subalpine zone, meadows.—Caucasus: Ciscaucasia, Dagestan,
Eastern, Western and Southern Transcaucasia. General distribution:
Balkans-Asia Minor, Armenia-Kurdistan. Described from a specimen
raised in the Berlin Botanical Garden. Place of seed collection not
mentioned in description. Type in Berlin.

Note 1. Specimens received at one time from Chamisso from the
Berlin Botanical Garden were found in the Leningrad herbarium. Most
probably, these specimens were raised even by Willdenow from the
seeds undoubtedly collected from the Caucasus. The specimens totally
resemble the description and diagram given by Willdenow in Hortus
Berolinensis (1. c.). Thus, these specimens are none other than the
isotypes of A. rigescens described by Willdenow. Attention must be
drawn to the fact that the pubescence of these plants is very sparse, the
leaves are green, the lobes of the leaves fairly broad, and the involucre
not as pale as in A. dumetorum Sosn., and also not as black-bordered
as is typical for A. macroglossa Somm. and Lev. and some other spe-
cies.

Note 2. A. sachokiana Sosn. ex Grossh. [Grossheim FI. Kavk. IV
(1934) 117], described very briefly only in Russian and without
designation of the type, is also listed in the Flora of the Caucasus. Its
characteristics are repeated in even more abridged form by Grossheim
in Opred. Rast. Kavk. [Key to the Plants of Caucasus] (1949, 452).
The plant, that is the holotype of A. sachokiana was collected, as
indicated on the label, in the Shemakha District, “between the Chevranly
pasture and Kurudag Mountain, July 28, 1928, M. Sakhokia” (A.A.
Grossheim’s herbarium, which is preserved in Tbilisi). Examination of
this specimen allows us to suggest that, possibly, it is a cattle-grazed
specimen of A. dumetorum, which, after grazing, produced a mass of
new leaves and a single shortened stem with a capitulum (a second
flowering in autumn) which frequently is observed in damaged herba-
ceous plants.

4

co

14

31. A. jailensis Zefir. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVIII (1957) 251; N. Rubzov in Tr. Nikitsk. Bot. Sada, XXIX,
FAX

Perennial. Stems numerous, erect or ascending, simple or weakly
branched, finely lanate, 20-50 cm high. Leaves subglabrous above,
lanate beneath, oblong, pinnately cut into 3-5 mm long segments, with
lobes terminating into whitish cusp. Capitula solitary terminal on stems
and their branches, large, 3-5 cm in dia. Involucral bracts sparsely
pubescent, oblong-ovate, yellowish-green, with deeper greenish-gray
vein, more or less brownish and fimbriate above along margin. Ligules
white, 1.5 cm long. Receptacular scales oblong, with obtuse but drawn
out, dark brown tip. Achenes compressed-tetraquetrous, curved, with
finely and indistinctly serrate or entire, slightly oblique corona.
Flowering June to August.

In meadows, on mountain peaks and in forest glades.—European
part: Crimea. Endemic. Described from Crimea (Babugan-Yaila, moun-
tain-steppe). Type in Leningrad.

32. A. dumetorum Sosn. in Vestn. Tifl. Bot. Sada, Ser. 2, 3 (1927)
160.—A. dumentorum var. subtripinnata Sosn. ex Grossh. Fl. Kavk.
IV (1934) 118; Sosn. in Fl. Gruzii, VIII, 309.—A. triumfettii auct. non
All.: Ldb. Fl. Ross. II (1844-1846) 525 p. p.—A. collina var. intermedia
Sosn. in Sosn. and Grossh. Opred. Rast. Tifl. (1920) 285, nom. nud.—
Exs.: Pl. or. oxs. No. 274.

Perennial. Plants arachnoid, covered with appressed, but not very
dense hairs, some grayish green. Stems numerous, about 30—40(60) cm
high, erect, angular in cross section, not densely leafy, branched from
middle and above, with erect unicapitulate branches, or stems simple,
unicapitulate. Leaves green above, subglabrous, grayish beneath from
appressed hairs, sessile, oblong-ovate, pinnately cut; segments oblong
or lanceolate, or oblong-linear, in turn divided into short, lanceolate,
serrate lobes and with sharply acute cusps, rachis narrow, with few
teeth as on leaf lobes. Capitula long-pedunculate, 2.5-3.0 cm in dia.
Bracts of sparsely hairy, involucre pale; outer bracts deltoid and obtuse,
inner oblong, somewhat obtuse, with brownish, scarious, fimbriately
truncate apex. Ligules oblong-spatulate, with almost entire, somewhat
truncate apex. Receptacle hemispherical; scales oblong, narrow, carinate
on Outer side, terminating into very narrow cusp as long as or longer
than florets. Flowering June to September (Plate III, Fig. 2).

In middle mountain zone.—Caucasus: All regions except Talysh
Mountains. Described from Georgia (Tbilisi). Type in Tbilisi; numerous
autotypes in Leningrad. Cf. Note 2 to A. rigescens Willd.

49

Plate III.
1—Anthemis zyghia Woron., habit, involucre, inner involucral bract, peripheral
(ligulate) floret, and disk floret; 2—A. dumetorum Sosn., habit, involucre,
involucral bract, peripheral (ligulate) floret, disk floret.

45

51

46

33. A. linczevskyi Fed. in Addenda, XXV, 868.—A. rigescens
auct. non L.: B. Fedtsch. Rast. Turk. (1915) 735.

Perennial. Plant greenish, with woody root, sparsely covered with
somewhat appressed hairs. Stems long, erect, slightly arcuate at base,
erect above, longitudinally striate, unicapitulate, about 60 cm high,
leafy from base to middle, leafless above, strong and virgate at fruiting.
Basal leaves caducous; cauline leaves oblong, antrorse, pinnately
divided into narrow-linear truncate-toothed segments; teeth and incisions
pointing forward, unequal, sharply acute. Rachis narrow, more or less
grooved. Capitula medium, 2.5—3.0 cm in dia; ligules white, scarcely
shorter than disk, oblong-elliptical, narrowed toward both ends, with
an apical notch or indistinctly 2—3 toothed. Involucre pubescent; all
bracts pale; outer bracts deltoid-lanceolate; inner oblong-lanceolate,
somewhat obtuse, brownish and weakly fimbriate at apex. Receptacle
hemispherical; scales stramineous, membranous or sharply acute.
Achenes lilac, with rather broad corona. Flowering June to September.

On dry mountain slopes and among shrubs.—Soviet Central Asia:
mountainous Turkmenia. Endemic. Described from Karakala District
(loldere Ravine, on the slopes of Khasardag Mountain). Type and
paratypes in Leningrad.

34. A. dubia Stev. in Bull. Soc. Nat. Mosc. XXIX, 4 (1856) 380;
Trautv. in Tr. Peterb. Bot. Sada, VIII, 392; Schmalh. FI. II, 61, tant.
nom.; Zefirov in Izv. Krymsk. Otd. Geogr. Obshch. SSSR, 3, 39-44,
in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI, 369-370, XVIII
(1957) 251-253; N. Rubzov in Tr. Nikitsk. Bot. Sada, XXIX, SO.

Perennial. Roots strong, fusiform, thick. Stems usually numerous,
simple or highly branched, ascending, arcuately or geniculately bent,
about 25—30(40) cm long, unicapitulate or with unicapitulate branches,
rather thick and, like leaves, grayish from rather dense pubescence,
leafy up to 1/2—2/3 of its height. Leaves obovate, up to 3 cm long,
pinnately divided into broadly linear, subobtuse, not strongly divergent,
somewhat distant segments, in turn divided into slightly toothed acute
lobes (cusps very small). Capitula medium, about 3.0—3.5 cm in dia;
ligules white, somewhat acute, lacking apical teeth, slightly shorter
than disk. Involucre lanate; outer bracts sharply deltoid, inner some-
what obtuse, oblong, with brownish membranous fimbria at tips, all
pale, but with greenish midrib, particularly prominent in upper half of
bract. Receptacular scales with slightly obtuse acumen. Flowering May
to August.

On lawns, in oak forests, on sands and patchy turf slopes.—
European part: Crimea (western part). Endemic. Described from

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47

Crimea (near Simferopol and Alma, on marly hills). Type possibly in
Helsinki. Isotypes apparently do not exist.

Note. This species could not have a more appropriate name, A.
dubia, i.e., “dubious chamomile.” In reality, big doubts remain as to
which of the species of Anthemis actually occurring in Crimea the
name proposed by Steven (Il. c.) should apply. Nevertheless, in
introducing this species now as a member of the Flora of the USSR,
one can base it on the intuitive, probably correct conclusion of B.M.
Zefirov (1. c.) that A. dubia, a quite widespread plant in Crimea, is
taxonomically close not to A. arvensis, as considered evidently by
Steven himself, but to A. rigescens and A. dumetorum. In any case, a
chamomile is found in Crimea, which, on the basis of its small (but
characteristic) distinctions, resembles the latter two species but, as a
matter of fact, has a branched stem and habit somewhat resembling A.
arvensis. If this plant can not be lumped with A. dubia Stev., then it
has to be considered a separate (and very good besides) new species.

Unfortunately, in the present case, Steven (I. c.) provided an
unsatisfactory diagnosis of his A. dubia, and it is impossible, without
having the type in hand, to get a clear idea of this species by reading
the diagnosis. I consider it useful to provide a translation of the origi-
nal diagnosis and notes made by Steven: “Leaves pubescent,
pinnatipartite; lobes small, lanceolate, absolutely entire and slightly
cuspidate; receptacle hemispherical; scales shorter than disk, lanceolate,
with cusp, slightly shorter than scale and somewhat obtuse; achenes
compressed-tetraquetrous, with membranous corona. Perennial. In habit
(similar) to A. arvensis, but many stemmed for the most part, height
up to 1 foot; with few unicapitulate branches, leafless over two-thirds
(the length). Florets (ray) white, but of same size as in A. arvensis;
disk (florets) yellow. Involucral bracts pale, lanate, with creamish
midrib. Ligule equal (in length) to diameter of disk. Cusp of scales
(receptacular) not acute, as in other (Crimean) species, but cylindrically
rolled. Achenes almost the same as those in A. rigescens, which has
conical receptacle. Because of presence of unique scales (receptacular),
it cannot be included under any other (Crimean) species.”

35. A. woronowii Sosn. in Vestn. Tifl. Bot. Sada, Ser. 2, 3 (1927)
162; Grossh. Fl. Kavk. IV, 117; Sosn. in Fl. Gruzii, VIII, 310.—A.
collina Sosn. in sched. ad specim. typ.—A. rigescens var. collina
Woron. in Sched. ad. Herb. Fl. Ross. IV (1908) 48.—A. triumfettii
auct. non All.: Ldb. Fl. Ross. III (1844-1846) 523 p. p.—A. tinctoria
var. rigescens Schmalh. FI. II (1897) 62 p. p. non A. rigescens Willd.—
Exs.: GRF No. 1734 (sub A. rigescente var. collina).

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48

Perennial. Plants appressed-hairy, grayish (especially stems). Stems
erect, about 40—50(60) cm high; somewhat dichotomously branched
above middle; branches unicapitulate. Leaves subglabrous above, but
with small glandular hairs, appressed-hairy beneath, sessile, obovate,
pectinately pinnately cut into short, oblong, approximate segments,
with very short-lanceolate, entire lobes, terminating into corny cusps;
rachis narrow, lacking teeth. Capitulum medium on rather long, leafless
thin peduncles. Involucral bracts pale, dorsally flattened; outer bracts
deltoid-lanceolate and acute, inner oblong, obtuse, narrow-scarious on
margin, slightly brownish at apex. Ligules lanceolate-oblong, as long
as disk, white. Receptacle hemispherical; scales oblong-lanceolate,
abruptly narrowed into cusp, carinate dorsally, as long as disk florets
or longer. Flowering June to August.

In lower forest zone, on herb-covered places.—Caucasus: Western
Transcaucasia. Endemic. Described from Abkhazia (near Tsebelda).
Type in Leningrad.

36. A. talyschensis Fed. in Addenda, XXV, 869.—A. triumfettii
auct. non All.: Ldb. Fl. Ross. II (1844-1846) 525, quoad pl. talysch.

Perennial. Root woody, many-headed, covered at apex with dry
remnants of leaf petioles; whole plant grayish-lanate. Stems ascending
from curved base or erect, about 20-30 cm high, usually branched;
branches unicapitulate, and some nonflowering, leafy up to lower 1/2,
leafless above, about 20—30 cm high, with white-lanate short internodes.
Leaves small, broadly ovate; all leaves sessile, divided into oblong-
linear, toothed, approximate segments. Capitula solitary, medium, about
3-4 cm in dia; ligules white, oblong, narrowed toward both ends,
inconspicuously 3-toothed at apex, as long as disk or slightly longer.
Involucre pubescent; outer bracts deltoid, inner oblong-lanceolate; all
bracts pale, scarious along margin and tip. Receptacle hemispherical;
scales pale-brown, membranous, oblong, abruptly narrowed into acute
brownish tip. Achenes obconical, brown with prominent ribs and
indistinctly toothed corona. Flowering May to August, fruiting August.

In middle mountain zone.—Caucasus: Talysh. General distribution:
Iran (Elburz Range). Described from Talysh Mountains (between the
villages of Tuli and Dzhoni). Type and paratypes in Leningrad.

37. A. euxina Boiss. FI. or. III (1875) 282; Trautv. in Tr. Peterb.
Bot. Sada, VIII, 392; Sosn. in Vestn. Tifl. Bot. Sada, Ser. 2, 3, 163;
Grossh. Fl. Kavk. IV, 117; Sosn. in Fl. Gruzii, VIII, 310.—A. rigescens
var. ochroleuca Somm. and Lev. Enum. pl. Cauc. (1900) 230.—A.
tinctoria f. floribus pallide luteis var. albidis Alb. Prodr. Fl. Colch.
(1895) 134.—Exs.: Pl. or. exs. No. 378.

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49

Triennial or perennial. Weakly pubescent plants with white curly
hairs or subglabrous, slightly fleshy, and with many stems. Stems long,
40-50(60) cm, divaricate, ascending, in lower part somewhat woody,
prostrate—usually creeping, weakly branched above. Leaves usually
small, oblong, pinnately cut into 3—4 lobes (on each side); lobes short
oblong-lanceolate, toothed, adjacent and almost fused at base; teeth
sharply acute; subtending leaves small, subentire. Capitula medium,
2.0—2.5 cm in dia, long-pedunculate. Involucral bracts appressed-white-
hairy, pale; outer bracts short-lanceolate, subacute, serrate, inner oblong-
linear, obtuse; all bracts greenish, longitudinally striate on outside.
Receptacle concave; scales carinate, abruptly narrowed into cusp slightly
shorter than disk. Ligules white, oblong-linear, usually entire at apex.
Achenes small, compressed-triquetrous, with many stripes, terminating
with short-truncate corona. Flowering June to August.

On coastal sands.—Caucasus: Western Transcaucasia (Black Sea
Coast). General distribution: Balkans-Asia Minor. Described from
Abkhazia (between Poti and St. Nicolas Bay). Type in Leningrad.

Series 3. Macroglossae Fed.—Involucral bracts with dark brown
or blackish, usually wide border; ray (ligulate) florets white. Stems
simple or sparingly branched. Leaves velutinous-tomentose, grayish,
scabrous-hairy or subglabrous and green. Perennial plants.

38. A. macroglossa Somm. and Lev. in Nuov. Giorn. Bot. Ital. II,
2(1895) 85; ej. Enum. pl. Cauc. 230.—A. platyglossa auct. non C.
Koch: Sosn. in Vestn. Tifl. Bot. Sada, Ser. 2, 3 (1927) 1927, p. max.
p.; Grossh. Fl. Kavk. VI. 117; Sosn. in Fl. Gruzii, VIII, 307, p. max.
p.

Perennial. Bright green, appressed-hairy or grayish plants. Stems
erect, unicapitulate, about 40-50 cm high, somewhat thick, hollow.
Leaves punctate-glandular, grayish-pubescent beneath, sessile, broadly
ovate or almost orbicular, pinnately cut; segments lanceolate-elliptical,
divided into linear, entire or pinnately toothed lobes with whitish cusps;
rachis narrow, with isolated teeth on foliar lobes. Capitula rather long-
pedunculate, large, up to 5—6 cm in dia. Involucral bracts carinate with
brownish border; outer bracts deltoid, acute arachnoid-hairy outside,
inner linear-oblong, subglabrous, attenuate into brown and scarious,
fimbriate apex. Ligules elliptical-lanceolate, wide, somewhat acute,
white. Receptacle hemispherical; scales linear, carinate, emarginate,
abruptly narrowed into somewhat obtuse and extended tip, as long as
disk. Achenes (immature) of disk florets triquetrous, compressed; outer
achenes distinctly winged, with short, almost entire, corona. Flowering
June to August.

5

Nn

50

In subalpine meadows.—Caucasus: Ciscaucasia (central), West-
ern Transcaucasia (east). Endemic. Described from Central Caucasia
(Svanetia, Latpari Range). Type in Florence.

Note. D.I. Sosnowsky (1927, 1. c.) included A. macroglossa Somm.
and Lev. with A. platyglossa C. Koch, described earlier from the Pontic
Range in Turkey. Besides, Sosnowsky used the description by Sommier
and Levier (I. c.) which pertains to A. macroglossa, but added a few
words from the description given by C. Koch for A. platyglossa.
However, there are insufficient grounds for lumping these species. A.
macroglossa is unquestionably endemic to the Main Caucasus Range
and is distributed essentially only in its central part. It is not really
known which race of this group actually grows in the Pontic Range. In
any case it is absolutely impossible to guess which species Koch ac-
cepted as A. platyglossa. The diagnosis of this species, as was always
the case with Koch, is totally unintelligible; the type is not designated
(it only is stated: “Auf pontischen Hochgebirge, 8—10.000' hoch. auf.
Urgestein”), and now the specimen also has disappeared (perished in
Berlin together with other authentic specimens remaining after Koch’s
death).

However, to facilitate possible future attempts to clarify the spe-
cies described by Koch, at least in some indirect manner, we are pro-
viding a translation of the original diagnosis given by Koch for A.
platyglossa [C. Koch in Linnaea, XXIII (1850) 319]: “Multi-cauline
(plant), weakly sericeous from appressed hairs. Stems entirely un-
branched, ascending, foot or more long, densely leafy below, leafless
above, fistular. Leaves orbicular-oblong, bipinnate, lobes elliptical,
sharply acute, 1-2-toothed. Involucre imbricate; bracts lanate, ovate-
lanceolate. Capitula large, together with ligules 1.5—2 inch in dia; ligules
broadly linear-oblong, pale yellowish, fertile. Achenes (immature) stri-
ate, with distinct unilateral corona. Scales (receptacular) oblong,
abruptly narrowed into spiny cusp, equaling disk florets.”

Characteristic is Koch’s remark that the stems of A. platyclossa
are fistular (fistulosi). Is some sort of deformity being described by
this term, for example, a case of fasciation of the common species A.
rigescens Willd.? The most characteristic feature, typical for A.
macroglossa Somm. and Lev., by which these species were identified
(blackish border of involucral bracts) was not mentioned at all. Exactly
such particularly large inflorescences can be found in fasciated plants.

39. A. schischkiniana Fed. in Addenda, XXV, 870.—A. tinctoria
var. bipinnatisecta Trautv. in Tr. Bot. Sada, IV (1876) 319.—Exs:.: PI.
orient. exs. No. 245 (sub Anthemide platyglossa), sp. typ.

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51

Perennial. Plants sparsely whitish-pubescent, but green. Root
woody. Stems few or solitary, unicapitulate, leafy from base to middle,
leafless above, 50-60 cm high, erect from arcuate base, finely striate.
Basal leaves absent (possibly caducous); cauline leaves broadly ovate,
sessile, twice or thrice pinnately cut into segments with oblong-linear,
acute, lobes with small cusps. Capitula medium, about 4 cm in dia.
Involucre subglabrous; outer bracts oblong-deltoid, inner oblong-
lanceolate, with brown scarious margin. Receptacle hemispherical;
scales membranous, acute, much longer than disk. Ligules, white,
oblong-broad lanceolate, narrowed toward both ends, indistinctly 2-3-
toothed or emarginate, somewhat yellowish when dry. Flowering July.

In subalpine zone.—Caucasus: Western Transcaucasia (south).
Endemic. Described from dried specimens (1. c.) from western Georgia
(Bakhmaro). Type in Leningrad.

40. A. abagensis Fed. in Addenda, XXV, 869.

Perennial. Plants subglabrous or sparsely covered with whitish
scabrous hairs, but bright green. Root brown, woody. Stems erect,
unicapitulate, leafy up to middle, leafless above, with thin stripes, 40-
50 cm high, arcuately bent at base. Basal leaves caducous; remaining
cauline leaves sessile, broadly obovate, pinnately divided into broad
elliptical pectinate and obtusely toothed segments, teeth terminating
into very small corny cusps. Capitula medium, about 4-4.5 cm in dia.
Outer bracts of almost glabrous or slightly hairy involucre oblong-
deltoid, narrowly dark brown along margin; inner bracts oblong,
yellowish green outside, broadly dark-brown or blackish along margin
and apex, scarious and fimbriate. Receptacle hemispherical; scales
membranous, pale, acute. Ligules white (somewhat yellowish on drying),
oblong, slightly shorter than disk. Flowering June to August.

In subalpine meadows.—Caucasus: Western Transcaucasia.
Endemic. Described from Caucasian Reserve (Abago Mountain). Type
in Leningrad, paratype in Reserve’s herbarium.

Note. Undoubtedly related to A. zyghia Woron. but at first glance
differs by its bright-green, almost glabrous leaves or weak but scabrous
(not soft and velutinous-tomentose and gray) pubescence.

41. A. zyghia Woron. in Izv. Kavk. Muz. XI (1917) 19; Sosn. in
Vest. Tifl. Bot. Sada, Ser. 2, 3, 156 (diagn. lat. compl. sub illegitima
et mutata transcriptione “zygia”); Grossh. Fl. Kavk. IV, 117; Sosn. in
Fl. Gruzii, VIII, 306.—A. rigescens var. albo-tomentosa C. Koch in
Linnaea, XXXIII (1850) 317.—Ic.: Kolak. Fl. Abkh. IV, Plate XXII
(mediocr.).—Exs.: GRF No. 3245.

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52

Perennial. Plants with many stems, entirely soft pubescent, ash-
gray. Stems 30-40 cm high, simple. Leaves broadly ovate, pinnately
cut, with lanceolate-ovate segments, divided into lanceolate or linear-
lanceolate, acute, sometimes toothed-truncate, slightly decurrent, and
pectinate-pinnate lobes; rachis with few teeth or incisions. Capitula
solitary, on long peduncle, clavate below capitulum, hemispherical, 3-
4 cm in dia. Receptacle hairy; bracts with wide brown border (usually
rusty on outside); outer bracts lanceolate, inner oblong-linear, obtuse
or subobtuse, indistinctly carinate on outside, somewhat rusty to scarious
at apices and densely fimbriate, or rarely incised. Ligules white, almost
as long as disk. Receptacular scales narrow-linear to lanceolate,
terminating into apically brown and uncinate cusp, longer than disk
florets. Flowering July to August. (Plate III, Fig. 1).

On limestone rocks, in mountains, at about 2,000—2,100 m.—
Caucasus: Western Transcaucasia. Endemic. Described from Abkhazia
(Pshitsa Mountains, cliffs and stony places). Type in Leningrad.

Note. D.I. Sosnowsky (1. c., 1927) was the first to change arbi-
trarily the spelling of the specific epithet “zyghia,” given by its author
J.N. Woronow (Il. c.), to “zygia”. But since the initial spelling of the
epithet is grammatically correct, it should be used from now on de-
spite the fact that all botanists (after Woronow) mentioning this spe-
cies in one connection or another used Sosnowsky’s spelling.

The specific epithet “zyghia,” used by J.N. Woronow, corresponds
to the ancient name of a locality in Abkhazia along the Bzyb River
near Pshitsa. This region was also known as “Dzighetia.”” Making an
adjective from the word “zigia” for uniformity of nomenclature, we
call this species the Zyghian chamomile.”

42. A. melanoloma Trautv. in Bull. Soc. Nat. Mosc. XLI, 2 (1868)
461, in Tr. Peterb. Bot. Sada, VIII, 392; Sosn. in Vestn. Tifl. Bot.
Sada, Ser. 2, 3, 157, pro min. p.; Sosn. in Fl. Gruzii, VIII, 308, pro
min. p.; Grossh. Fl. Kavk. IV, 117, pro min. p.—A. tinctoria var.
melanoloma Trautv. in Tr. Bot. Sada, IV (1876) 377.—A. rigescens
auct. non Willd.: Boiss. Fl. or. II (1875) 281, p. min. p.

Perennial. Stems erect, simple or with branches spreading from
middle, to 40 cm high, slender, unicapitulate, appressed-pubescent,
leafless in lower half. Basal leaves possibly caducous; cauline leaves
elliptical, sessile, glabrous above, appressed-pubescent beneath,
pinnately cut; segments erect, oblong-linear, pectinate-pinnate, divided
into linear-lanceolate and entire lobes, terminating into cusps; rachis
narrow-winged. Involucral bracts linear-lanceolate, subobtuse or obtuse,
appressed-pubescent, sometimes lanate on margin, with blackish
membranous border, green in middle; inner bracts very finely incised-

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53

fimbriate at apex. Receptacle ovoid-conical; scales linear, dense, longer
or shorter than florets, gradually narrowed into somewhat strong cusp.
Ligules oblong-linear. Corona conical, very short, brownish,
membranous. Flowering June.

In subalpine zone.—General distribution: Balkans-Asia Minor
(east). Possibly, found also in the Lesser Caucasus. Described from
Saganlug Range in Turkey. Type in Leningrad.

Note. Study of the holotype of this species and comparison of it
with others indicate that it has not yet been found in the Caucasus;
other species partly overlap this species: A. macroglossa Somm. and
Lev. and probably also A. abagensis m., in part.

At the time of Trautvetter, who described this species, the Saganlug
Range (locus classicus) was within the limits of Turkey, then (until the
October Revolution) Saganlug became the border range between Turkey
and Russia. However, now again it is in Turkey (i.e., in Asia Minor).
The comment of D.I. Sosnowsky (I. c., 1927) that “the occurrence of
this species in Asia Minor has not yet been reported, but theoretically
it is entirely possible,” was apparently based on a misunderstanding.

The type specimen (holotype) was selected by Trautvetter from
the collection of Lagovsky, who subsequently became notorious for
the extreme confusion (or even very strange and inexplicable juggling)
of the labels of his Caucasian and Ciscaucasian collections. As the
label indicates, the authentic specimen originates from Saganlug Range,
but it is not certain if the plant was collected in fact from this place.

Boissier annotated the type specimen as follows: “. . . mihi videtur
Anthemidis rigescentis forma involucri phyllis itensius nigrescentibus.”
Despite this, it must be recognized that A. melanoloma is a very good
species, because of not only its black-bordered involucral bracts but
also its leaves which are deeply divided, with divergent segments,
differing sharply from A. rigescens as well as all other species of this
group. The author himself of this species, Trautvetter, of course, quite
mistakenly referred his species to the A. tinctoria complex and later
unnecessarily (perhaps under the influence of Boissier’s authority)
reduced his own species A. melanoloma to the rank of a variety.

Series 4. Altissimae Fed.—Involucral bracts monochromatic, pale;
ligules white. Stems strongly branched, erect. Leaves green, pubescent;
hairs erect; segments linear, pinnatifid. Annuals.

43. A. altissima L. Sp. pl. II (1753) 893; C. Koch in Linnaea,
XXIII, 316; Boiss. Fl. or. II], 282; Schmalh. Fl. II, 61; Grossh. FI.
Kavk. IV, 118; Sosn. in Fl. Gruzii, VIII, 311; B. Fedtsch. Rast. Turk.
735; Fedtsch. and Fler. Fl. Evrop. Ross. 968.—Chamdemelum cota

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54

C.A. Mey. Verzeichn. (1831) 75, non A. cota L.—Cota altissima J.
Gay ex Guss. Fl. Sic. Syn. II (1842-1844) 867.—Ic.: Rchb. Icon. t.
1008.

Annual. Plants sparsely pubescent or subglabrous with erect, tall,
30—40(60) cm long, strong and upright stem, branched above. Leaves
ovate-oblong, 2-3 pinnately cut into short, lanceolate, acerate to acutely
conical lobes; rachis slightly serrate. Peduncles of inflorescence clavate
at blooming. Capitula large, 3.0—3.5 cm in dia. Outer involucral bracts
short-lanceolate, inner oblong, more scarious than outer. Receptacle
hemispherical; scales oblong-lanceolate, truncate or with apical notch,
abruptly narrowed into somewhat strong cusp as long as disk florets or
larger. Ligules white, elliptical-linear, as long as disk. Achenes dark
brown, compressed-tetraquetrous, slightly winged along margins, with
10 thin ribs on both sides and short apical border, but lacking corona.
Flowering May to September.

In dry places, partly in ruderal conditions.—European part: Crimea;
Caucasus: All regions; Soviet Central Asia: Kara-Kum (southwest);
mountainous Turkmenia. Described from Mediterranean Region. Type
in London.

44. A. coelopoda Boiss. Diagn. pl. or. II, 11 (1849) 12; Boiss. Fl.
or. III, 283; Grossh. Fl. Kavk. IV, 118.—Cota coelopoda Boiss. Diagn.
pl. or. II, 3 (1849) 21.

Annual. Plants weakly pubescent or subglabrous. Stems erect or
divaricately branched below middle, about 40 cm high. Leaves ovate,
twice pinnately cut into segments, in turn divided into very fine oblong-
linear corny-tipped, highly divaricate lobes. Peduncles of inflorescence
strongly clavate after flowering, fistular; capitula large, 3—4 cm in dia.
Outer involucral bracts oblong-lanceolate, acute; inner obtuse.
Receptacle hemispherical; scales oblong-linear, carinate, terminating
into cusp. Ligules white, elliptical linear, as long as disk. Achenes
compressed tetraquetrous, with indistinct vein on each side, or
sometimes 3-veined, with acute apical border, but without corona.

In lower and middle mountain zones.—Caucasus: Talysh (reported
by A.A. Grossheim from the Talysh Mountains, Zuvant). General
distribution: Balkans-Asia Minor. Described from Asia Minor (Smyrna
Plains, near Geir). Type in Geneva; isotype in Leningrad.

45. A. austriaca Jacq. Fl. Austr. Icon. et descr. V (1877-1878)
22; DC. Prodr. VI, 11, excl. var.: C. Koch in Linnaea, XVII, 44; Ldb.
Fl. Ross. II, 525; Boiss. Fl. or. III, 284; Schmalh. Fl. II, 61; Fedtsch.
and Fler. Fl. 968; Grossh. Fl. Kavk. IV, 118; Zefirov in Bot. Mat.
Gerb. Bot. Inst. Akad. Nauk SSSR, XVIII, 254.—A. hyrcana Sosn. ex
Grossh. Opred. Rast. Kavk. (1949) 453, nom. nud.—A. hyrcanica Sosn.

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55

(1947) in sched.—A. zangelana Sosn. ex Grossh. Opred. Rast. Kavk.
(1949) 453, nom. nud.—A. zangelanica Sosn. (1947), in sched.—Cota
austriaca J. Gay. ex Guss. Fl. Sic. Syn. II (1842-1844) 866.—Ic.:
Jacq. loc. cit. t. 444.

Annual. Plants grayish-pubescent. Stems erect, about (15)20—40(60)
cm high, branched. Leaves oblong, pinnately cut into pectinate, linear
segments with small, linear-oblong, acerate acute lobes; rachis toothed.
Peduncles of inflorescence not clavate; capitula medium, 2.5—3.0 cm
in dia. Involucre hispid; outer bracts lanceolate, inner linear-oblong
rather narrow, scarious at tip. Receptacle conical; scales oblong, at-
tenuate into short acumen, almost as long as disk. Ligules elliptical-
linear, white, as long as disk. Achenes tetraquetrous, slightly com-
pressed, with 3 indistinct grooves on each side, and short-toothed lobes
of inconspicuous corona along upper margin. Flowering June to Au-
gust.

In dry, somewhat weedy places.—European part: Crimea, Upper
Dnieper, Upper Dniester, Bessarabia. Caucasus: Ciscaucasia, Eastern
Transcaucasia (south), Talysh. General distribution: Central Europe.
Described from Austria. Type in Vienna.

Note. The species A. hyrcana Sosn. ex Grossh. and A. zangelana
Sosn. ex Grossh., inserted in A.A. Grossheim’s (1. c.). Opred. Rast.
Kavk., apparently represent small specimens of A. austriaca. The char-
acters by which these “species” are separated Grossheim’s key (corona
rudimentary, sometimes cylindrical, sometimes auriculate), possibly are
the result of variation within the same species, i.e., A. austriaca. It
is important to note that these “species” thus remained in the status
“nomina nuda,” and D.J. Sosnowsky himself did not try to describe
them, leaving the names given to them only on the herbarium
labels.

Series 5. Wiedemannianae Fed.—Involucral bracts with white
border. Stems branched, sinuate. Leaves pinnate, weakly pubescent;
hairs curly; leaf segments sharply toothed. Annuals.

46. A. wiedemanniana Fisch. and Mey. in Ind. Sem. II (1835) 27;
DC. Prodr. VI, 10; Ldb. Fl. Ross. II, 523; Boiss. Fl. or. III, 286;
Grossh. Fl. Kavk. IV, 118; Sosn. in Fl. Gruzii, VIII, 312.—A.
ormenioides Boiss. Diagn. pl. or. I, 4 (1844) 5.—Cota ormenioides
Boiss. ibid. II, 3 (1849) 21.

Annual. Plants grayish from curly and appressed hairs, 10-15 cm
high. Stems numerous, short, branched from base. Leaves long- and
narrow-linear, pectinately pinnately cut into deltoid, acicular, basally
toothed lobes. Peduncles of inflorescences very short, thick. Involucre

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56

compact and hispid; bracts oblong, white-bordered; outer bracts acute,
inner somewhat obtuse. Receptacle conical; scales oblong-lanceolate,
carinate, somewhat obtuse and slightly truncate, mucronate. Ligules
white, linear, truncate-3-toothed, shorter than disk. Achenes small,
black, sharply tetraquetrous, compressed, punctate (under lens), with
very short apical border in place of undeveloped corona. Flowering
May to June.

On cultivated lands, ruderal and weedy places.—Caucasus: Western
Transcaucasia (south). In the USSR, found only within the limits of
Adzharia. Described from Asia Minor (mountains near Angora, near
Safranbol and Mersivan). Type in Leningrad.

Section 4. Maruta (Cass.) Boiss. Fl. or. III (1875) 280.—Sect.
Euanthemis DC. Prodr. VI (1837) 7 p. p.—Maruta Cass. in Dict. Sc.
Nat. XXIX (1823) 174, pro gen.—Sect. Cotula Rupr. Fl. Ingr. (1860)
591, nom. absque diagn.—Achenes apically truncate, with or without
distinct and sometimes many-toothed corona. Receptacular scales
cuspidate or narrow-lanceolate. Annual herbs with finely divided pinnate
leaves and usually branched stems.

Type of section: A. cotula L.

Series 1. Cotulae Fed.—Leaves twice pinnate, lobes, linear or
filiform, numerous; stems usually tall, 15—30(40—60) cm high, strongly
branched. Plants sparsely pubescent or subglabrous.

47. A. cotula L. Sp. pl. II (1753) 894; Rupr. Fl. Ingr. 591; C.
Koch in Linnaea, XXIII, 315; Boiss. Fl. or. III, 315; Schmalh. FI. II,
62, excl. var.; Fedtsch. and Fler. Fl. 969 p. p.; Krascheninn. Sorn.
Rast. SSSR, IV, 223; Grossh. Fl. Kavk. IV, 116; Sosn. in Fl. Gruzii,
VIII, 317; B. Fedtsch. Rast. Turk. 735.—A. foetida Lam. FI. fr. I
(1778) 164.—Martua foetida Cass. in Dict. Soc. Nat. XXIX (1823)
174.—M. cotula DC. Prodr. VI (1837) 13; Ldb. Fl. Ross. II, 526.—
M. vulgaris Bluff and Fing. Fl. germ. II (1825-1833) 392.—
Chamaemelum cotula All. Fl. Pedemont. (1785) 676.—Cotula foetida
S.G. Gmel. Reise, I (1770-1784) 137.—ic.: Rchb. Icon. t. 1000;
Syreistsch. Fl. 261.—Exs.: Fl. Polon. Exs. No. 179; Herb. FI. Ingriae,
No. 310; Callier. Iter Taur. (1900) No. 629 (sub. A. arv.).

Annual. Plants scabrous from sparse hairs or almost glabrous. Stems
cymosely branched, 15—20(30) cm high. Leaves ovate-oblong, twice
pinnately cut into narrow-linear, entire or 2—3 toothed, short-acute,
erect lobes. Peduncles of inflorescence not clavate. Involucral bracts
oblong, obtuse, narrow-scarious along margin. Receptacle oblong-
conical; scales absent in lower part, linear above and at tip, cuspidate,

61

Plate IV.
1—Anthemis cotula L., habit; involucre; outer and inner involucral bracts;
peripheral (ligulate) florets, disk florets; 2—A. candidissima Willd., habit;
involucre; involucral bract, peripheral (ligulate) floret, disk floret.

57

64

58

not exceeding disk florets. Ligules oblong-elliptical and present in
peripheral sterile florets; tubes of disk florets basally swollen. Achenes
roundish in cross section, more or less finely tuberculate, lacking corona,
sunken, but usually slightly crenate from truncate ribs along margin.
Flowering May to August. (Plate IV, Fig. 1).

In weedy places.—European part: All regions except north;
Caucasus: All regions; Soviet Central Asia: Reported from extreme
northwest. General distribution: Central Europe. Described from “weedy
places of Europe, especially Ukraine.” Type in London.

Note. Based on the citation from Species plantarum of Linnaeus,
the Ukrainian plants can, to a certain extent, be considered topotypes.
Therefore, for the quite sharply isolated race growing more to the
north and west and described from Lithuania (A. lithuanica),
distinguishing itself as a separate species, there is a basis to call it by
the name proposed by Besser, without doubting that the Linnaean type
could be more similar to the northwestern race than to the southern.

48. A. lithuanica (DC.) Trautv. in Tr. Peterb. Bot. Sada, VII
(1883) 392.—A. lithuanica Bess. ex DC. Prodr. VI (1837) 13, pro
syn.—A. cotula B. lithuanica (DC.) Schmalh. FI. II (1897) 62.—Maruta
cotula B. lithuanica DC. ibid.; Ldb. Fl. Ross. II.

Annual. Stems strongly divergent, 50-60 cm high; internodes long.
Plants glabrous or very sparsely pubescent. Leaves twice pinnately
cut, ovate, sessile, divided into ovate segments, with very thin, hairy,
oblong lobes. Capitula small, not more than 1.5 cm in dia; peripheral
florets with ligule up to 5-6 mm long, as long as disk. Receptacle
ovoid-conical; peduncles of inflorescences not clavate. Involucral bracts
oblong, somewhat obtuse, scarious along margin. Receptacular scales
linear, shorter than florets. Achenes roundish in cross section, lacking
corona. Flowering May to July.

On borders of fields, in weedy places.—European part: Upper
Dnieper (north). Endemic? Described from Lithuania. Type in
Leningrad.

Series 2. Odontostephanae Fed.—Leaves pinnate, lobes very small,
few. Stems short, 5-10 cm high, slender, simple or sparingly branched.
Plants pubescent.

49. A. odontostephana Boiss. Diagn. pl. or. I, 6 (1845) 85; Boiss.
Fl. or. III, 319; B. Fedtsch. Rast. Turk. 735.

Annual. Plants short, 5-10 cm high, ash-gray-pubescent. Stems
usually numerous, thin, simple or weakly branched in lower part. Leaves
rather small, petiolate, ovate, pinnately cut into 2-3 somewhat acute,

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59

3-5 lobed segments, or segments very small. Peduncles of inflorescences
on terminal branches, long, clavate at apex. Capitula small, few-
flowered. Involucre compact, hispid; bracts oblong, obtuse, brownish-
scarious at apex. Ligules of peripheral pistillate florets oblong, slightly
longer than disk. Receptacle short-conical; scales few, setosely pointed.
Achenes terete, ribbed; outer achenes tuberculate-scabrous, with truncate
apex and 10-15 divergent teeth of indistinctly developed corona; inner
achenes usually almost smooth, with unilateral short-toothed corona.
In dry places, in middle mountain zone.—Soviet Central Asia:
mountainous Turkmenia (Greater Balkans). Described from Iran
(Persepolis Ruins). Type in Geneva; isotype in Leningrad.

Series 3. Microcephalae Fed.—Leaves simple, linear or pinnate,
but with fewer lobes. Stems short, filiform, 5-10 cm high, simple or
weakly branched. Plants almost wholly glabrous.

50. A. microcephala (Schrenk.) B. Fedtsch. in Rast. Turk. (1915)
735; Pavlov, Fl. Tsentr. Kazakhst. III, 238.—Maruta microcephala
Schrenk in Ind. Sem. Horti. Petrop. X (1845) 53; ibid., XI, 77; Trautv.
in Tr. Peterb. Bot. Sada, VIII, 393; ej. in Bull. Soc. Nat. Mosk. II, 39;
O. and B. Fedtsch. Perech. Rast. Turk. IV, 180.

Annual. Plants small, usually not more than 10 cm high, but
reaching 20 cm or more in cultivation (in botanical gardens), almost
entirely glabrous, with very thin, sometimes almost filiform stem (to 1
mm in dia in cultivation), divaricately branched almost from base,
sparsely leafy, with leafless long peduncles of inflorescences terminal
on stem branches. Leaves sessile, pinnatisect, cuneate-ovate; segments
narrow, linear, lobate or palmatipartite. Capitula very small, 0.3-0.5
cm in dia; peripheral (ligulate) florets with small, 2-toothed, almost
linear, white limb. Receptacle ovoid-conical, covered with somewhat
setaceous, small scales. Achenes very small, brownish, roundish, slightly
winged and finely sulcate, very indistinctly tuberculate, lacking corona.
Flowering June to August.

In dry, sandy and silty soils, dry stream beds, floodplains of rivers,
near irrigation channels, and in steppes (as a spring ephemeral).—
Soviet Central Asia: Lake Balkhash Region, Aralo-Caspian Region
(east). Endemic. Described from Dzhungaria (“in mountain valleys of
the Maibshalyrgan’’). Type and isotypes in Leningrad.

51. A. deserticola Krasch. and M. Pop. in Tr. Bot. Inst. Akad.
Nauk SSSR, Ser. I, 1 (1938) 178; Zakirov, in Tr. Uzb. Univ. Nov. Ser.
No. 89 (1957) 147.

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60

Annual. Plants glabrous. Stems 10-12 cm high, roundish in cross
section, longitudinally striate, simple or branched; branches long,
unicapitulate, filiform, but slightly clavate below inflorescence. Leaves
somewhat fleshy, entire, narrow-linear, 1-3 cm long, 1-2 mm wide,
obtuse, sometimes twice or thrice pinnately cut or irregularly pinnately
cut, with narrow-linear lobes. Capitula solitary, small, 3-5 mm in dia.
Involucre imbricate; outer bracts lanceolate, narrow-scarious, inner
ovate, obtuse, apically membranous. Ligules white, oblong, without
apical border, 2 mm long. Receptacle obtusely conical; scales
acuminate. Achenes slightly curved, obconical, 1 mm long, with 4—5
white, slightly winged ribs, truncate apex, and short corona incised on
margin. Flowering April.

Sandy deserts and saline soils.—Soviet Central Asia: Kyzyl-Kum.
Endemic. Described from Kazakhstan (Kyzyl-Kum Sands between Turd-
Kuduk and Kutun). Type and paratype (Uzbekistan, “Hungry Steppe’)
in Leningrad.

GENUS 1514. Chamaemelum Mill.!:?

Mill. Gard. Dict. Abridg. ed. 4 (1754); Adans. Fam. II (1763)
128; Cass. in Dict. Sc. Nat. XXIX (1823) 179, 185, non Vis. 1845.

Capitula on rather long erect peduncles, solitary on apices of main
stem and its branches, but usually many on a single plant, heterogamous,
with rather numerous peripheral pistillate ligulate florets, in one row
(in cultivated forms, often in several rows), and with numerous bisexual,
tubular disk florets, or homogamous, with only bisexual tubular disk
florets. Involucre broadly scutellate, 6-12 mm in dia and 3-6 mm
long; bracts herbaceous, with wide, light colored, membranous border
appendiculately broadened at apex, imbricate in 3-4 irregular rows,
with inconspicuous midrib; outer bracts lanceolate, with narrower
membranous border; inner bracts oblong and oblong-linear 1.5—2 times
as long. Receptacle slightly convex, obtusely conical, 2—2.5 times as
long as wide, fistular, punctate-alveolate, squamose; scales oblong-
linear, obtuse, with appendiculately broadened, light colored,
membranous border at apex. Corolla of ligulate florets (sometimes
present) white; ligules linear-oblong, 7-14 mm long, with punctate
glandular hairs below, decurved after anthesis; tube about 2.5 mm
long, strongly flattened dorsally, almost like wing with more or less

'Treatment by N.N. Tzvelev.
*From the Greek words: Chamai—small, and melon—apple; the smell of the
capitula of the species of this genus distinctly resembles that of a ripe apple.

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61

distinct, downward directed appendages at base, covering ovary apex;
corolla of tubular florets yellow 1.8-2.5 mm long, with 5 deltoid,
curved teeth; corolla tube, densely covered with punctate glandular
hairs, gradually broadened toward throat and base, with thickened scale
on lower side, more or less covering ovary apex by downward directed
appendages. Filaments rather long, glabrous, anthers without distinct
basal appendages, but with lanceolate-ovate-obtuse apical appendages.
Style bifid; branches narrow-linear, truncate. Achenes 1.2—1.5 mm long,
about 0.6 mm in dia, prismatically cylindrical, narrowed toward base,
glabrous, with 3(4) prominent veins, lacking pappus. Perennial herbs,
strongly aromatic with more or less branched, erect or ascending stems.
Leaves alternate; laminas 2—3-pinnatisect in very narrow lobes.

A few (2-3) species of this genus are distributed mainly in Southern
Europe and North Africa; one species is widely cultivated as a medicinal
and ornamental plant and often grows wild.

Type of genus: Chamaemelum nobile (L.) All.

Note. Following R. Visiani, Osserv. Matric. 1845, p. 12 and FI.
dalmat. II, 1847, p. 84, the generic name Chamaemelum has been
wrongly used in many botanical works up to the present time for a
large group of species belonging to the genus Tripleurospermum Sch.
Bip. However, the species Anthemis nobilis L., which has a whole
series of significant differences from other Anthemis L. species, un-
doubtedly should be considered the type species of the genus
Chamaemelum, established by Tournefort and subsequently restored
by Miller and Adanson; as a result, we also are adopting the generic
name Chamaemelum for it.

C. nobile (L.) All. Fl. Pedem. I (1785) 185.—Anthemis nobilis L.
Sp. pl. (1753) 894; DC. Prodr. VI, 6; Schmalh. Fl. I, 60; Taliev,
Opred. Vyssh. Rast. (1907) 486; Fedtsch. and Fler. Fl. Evorp. Ross.
III, 968.—Ic. Komarov, Sbor. Sushka i Razved. Lekarstv. Rast. Izd. 3
(1917) Plate 34; Hegi, Ill, Fl. VI, 1 (1918) 536, f. 272; Zemlinsky,
Lekarstv. Rast. SSSR (1951) 435.

Perennial. Plants 10-30 cm high, strongly aromatic, usually strongly
branched from base, grayish-green from rather long, simple, sparse,
appressed hairs. Stem erect or ascending, more or less branched, densely
leafy. Leaves, except lowermost, sessile, but always with small lateral
lobes near distinctly broadened base; lamina twice or thrice pinnately
cut, 1-6 cm long and 4-15 mm wide, oblong or linear-oblong, apical
lobes very narrow, linear or lanceolate-subulate, with rather long
cartilaginous cusp. Capitula solitary at apices of stem and branches.
Involucre 6-12 mm in dia and 3—6 mm long; bracts with wide, light
colored, membranous border. Ligulate florets numerous, white; disk

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62

florets tubular, yellow. Achenes 1.2—1.5 mm long and about 6 mm
wide with 3(4) prominent veins, lacking pappus. Flowering July to
September.

Cultivated as a medicinal and ornamental plant and now. and then
found in fields, by roadsides, as an escaped or naturalized plant.—
European part: Baltic Region, Upper Dnieper, Upper Dniester, Middle
Dnieper, Bessarabia, Black Sea Region, Crimea. General distribution:
Atlantic Europe, Central Europe, Mediterranean Region, Balkans.
Introduced into other regions. Described from Southern Europe. Type
in London (Kew Botanical Garden).

Economic Importance. Long used as a medicinal plant. The bloom-
ing capitula are used for this purpose, the medicinal effect of which is
very similar to that of medicinal chamomile (Matricaria recutita L.)
but, according to S.E. Zemlinsky (op. cit.), stronger; also cultivated as
an ornamental plant (usually the “double flowered” forms with tubular
florets more or less modified into ligulate ones).

GENUS 1515. Anacyclus L.':*

L. Spr pli 753) 892: |

Capitula many-flowered; outer florets ligulate, pistillate, with short-
winged tube; other florets tubular, bisexual, 5-toothed, very rarely all
florets tubular; achenes smooth, compressed, at least outer with wide
transparent wings on both sides, extended above into auricles; involucre
membranous; receptacle with bracts. Annual herbs.

1. Peduncles not thickened; petals yellow; corolla tubes of all florets
(especially outer) setose-ciliate; bracts oblanceolate, as long as
corolla tubes; achenes subglobose or ObOVOI ..............:eeeeeeseeeee
RRA TE Re ee 1. A. ciliatus Trautv.

+ Peduncles thickened; petals white above, reddish below; corolla
tubes of all florets glabrous, bracts cuneate-obovate, shorter than
corolla tubes; achenes deltoid-obovoid ...... 2. A. officinarum Hayne

1. A. ciliatus Trautv. in Bull. Soc. Nat. Mosc. XLI, 2, (1868) 461;
Grossh. Fl. Kavk. IV, 121.—Anthemis ciliata (Trautv.) Boiss. FI. or.
III (1875) 312.

Annual. Stem branched, flexuous, velutinous, 30-50 cm high.
Leaves weakly pubescent or glabrous, oblong, twice or thrice pinnately

'Treatment by C.S. Afanasiev.
*From the Greek words: anai—without, and cyclos—circle; named for sterile
outer ligulate florets, which are only pistillate.

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63

cut into thin, narrow-linear, short-cuspidate, divergent, 3-7 mm-long
lobes. Capitula 1.0-1.5 cm in dia, solitary at apices of main stem and
branches, on long, not thick, short-villous peduncles. Involucre short
pilose; bracts oblong, outer acute, inner somewhat obtuse with narrow
white-membranous margin. Receptacles hemispherical; scales
membranous, oblanceolate, subacute, as long as corolla tubes. Outer
florets pistillate, yellow, with oblong-elliptical ligules, 7-10 mm long
and setose-ciliate tubes; disk florets bisexual, tubular, with more or
less setose hairy tubes. Achenes subglobose or obovoid, 3-4 mm long
and 2.5-3.0 mm in dia, flat, with wide transparent wings, extended
above in more or less globos auricles in outer achenes. Flowering June
to July.

Along edges on dry slopes in lower mountain zone.—Caucasus:
Eastern Transcaucasia. Endemic. Described from Georgia. Type in
Leningrad.

2. A. officinarum Hayne Artzn. Gew. IX (1825) 46; Ldb. FI.
Ross. II, 2, 527; Schmalh. II, 63; Stankov Opred. Rast. Evrop. Ch.
625.—A. puncher Bess. ex DC. Prodr. VI (1837) 16.—Ic.: Hayne, op.
cit. t. 46; Hegi, Ill. Fl. VI, 2, fig. 382.

Annual. Stem erect or ascending, branched, weakly hairy,
sometimes almost glabrous in lower part, to 40 cm high. Leaves weakly
pubescent, oblong, twice pinnately cut into more or less sparse, entire,
2-3-fid, acute, linear, and linear-subulate, divergent, 4-10 mm long
lobes. Capitula about 1 cm in dia, solitary at apices of main stem and
branches, on long, thickened, short pilose peduncles. Involucre
hemispherical, short-pilose; bracts oblong, with white-membranous
margin; outer bracts acute, inner somewhat obtuse. Receptacle convex,
with membranous cuneate-obovate, short cuspidate scales, shorter than
corolla tubes. Outer florets with elliptical ligules, 8-13 mm long,
sometimes absent (var. discoidea), white above and reddish below;
disk florets bisexual, tubular, lemon-yellow, corolla tubes of all florets
glabrous. Achenes flat, deltoid-obovate, 2.5-3.0 mm long and 2.0—2.5
mm in dia, with wide wings extended above into broadly deltoid auricles
in outer achenes, much narrower and lacking auricles in inner achenes.
Flowering July to August.

Reported (in the literature) as a wild plant (escaped from botanical
gardens, where it is sometimes cultivated) from Podolia (vicinity of
Kremenets), and from the Volyn and Poltava regions.—General
distribution: Cultivated on a small scale in Bohemia and Thuringia.
Described from Germany from cultivated specimens. Type in Berlin.

Economic Importance. A medicinal plant. Its root (Radix pyrethrii
germanici s. communis. Die Deutsche Bertramwurzel) containing

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64

essential oil is used. In German folk medicine it is used as an analgesic
for toothaches, for diseases of the tongue, as a sialogogue and an
external stimulant. Its action is similar to that of the root of A. pyrethrum
DC. (Radix purethree romani) used in Austrian pharmacopeia, which
contains inulin, essential oil, and alkaloids—pyrethrin and pellitorin,
only its action is slightly stronger.

Note. Besides A. officinarum Hayne, other Anacyclus species are
also sometimes cultivated in botanical gardens, as for example, A.
pyrethrum DC., A. clavatus Pers. (=A. tomentosus DC.) and a few
others. Apparently, the Schmalhausen’s report (Flora, II, 63) on the
occurrence of A. tomentosus DC. in Crimea (Sevastopol) refers to
cultivated specimens.

GENUS 1516. Achillea L.'?

Li Sp--pl. (1753) 896:

Capitula small, many-flowered, in corymbs, less often solitary.
Involucre oblong, ovoid, oblong-cylindrical, turbinate or hemispherical;
involucral bracts imbricate, with white-membranous, yellow, or brown-
membranous margin. Receptacle flat, convex, conical or very rarely
long-cylindrical, with membranous scales, as long as tubular florets or
much shorter. Outer florets pistillate, usually 1-seriate, 5—23, ligulate;
ligules white, pink, red, or yellow, small, 1/6—1/2 as long as involucre,
equal or larger, sometimes deformed, less often absent; disk florets
bisexual, tubular, 5-toothed, corolla tube compressed, with more or
less swollen base, covering ovary apex. Achenes compressed dorsally,
oblong, cuneate, rarely oblong-obovoid, or oblanceolate, lacking or
with somewhat thick spongy wing, apically truncate, lacking pappus or
corona. Perennial herbs, sometimes woody at stem base, with entire,
pinnately lobed, pinnately cut, or pinnately parted leaves.

Type of genus: A. millefolium L.

The genus Achilles includes more than 20 species, distributed
almost exclusively in the temperate zone of the Old World, mainly in
the mountains and in the Mediterranean Region.

L.isyAchenes wingless 20e\::. o)4. en. eet eden) eee L.
+ Achenes winged; wing somewhat thick, spongy ...........:see+ 26.

'Treatment by C.S. Afanasiev; section Ptarmica (DC.) Koch by V.P. Botschantzev.
*The name of the plant used by ancient authors originated from Achilles, hero of
the Trojan War.

71

P+. fee

ee

65

Leaves narrow-linear, with very small, 1-2(3) mm long, transverse,

Semen inatermectmerien 6155.0.) cteeeeiebes, crt ethene ttt: pte la tits, .f6i else Vde(danasss Ze.
Leaves wider; segments larger, neither transverse nor imbricate ......
I I ta isd aananangniensousss 3.
Plants entirely glabrous ................::ccsseeeee 17. A. glaberrima Klok.
Plant more prlesa pubescent) iiiiis..<it tease swaths. wer caneddndesitiebsessees 4.
LGAVES POGUE (bk 2c iercden day su decdadalee ae: 15. A. ochroleuca Ehrh.
Leaves pinnately cut or pinnately parted, but not pectinate ........ ee
Ligules ‘wiite, pinkson ted rz..c5c cre AS) ot ont id BRO seen 6.
Ligules. yellownor alder wsgiczts:..a0e. 263. lek BSR... LS.
Leaf segments wide, ovate, sharply toothed ...............ceseeeseeseeeeee
LASS «a Sad cite SBR eeilaen IR, ety sl Meret be. 12. A. latiloba Ldb
Leal SCQMeENts, OGRER WIRE i cisascanapsnnsnnnasinannannecthEbe SIAN EES +0ess q:
Rhizome short, many-headed, plants lacking underground shoots;
leaf segments lineatisrs2).. cues ..7i...ceh abet leo 1. A. nobilis L.
Rhizome branched, plants with underground shoots; leaf segments
not lnéani broader 32hts.tes, gers sche cierrcd bh, eae A nano 8.
Rachis of cauline leaves always with intermediate (between basal
scoments ighes eritcetnk.. Lied. . vovgats, .22i..2el etd. es AGEs 9.
Rachis of cauline leaves lacking intermediate lobes, sometimes with
SOON MRO ATE FECTED a cissexgens DRL Gs octal Laken) aati. 10.

Rachis of leaves 1.0—2.5(4.0) mm wide, with incised-toothed lobes
or teeth throughout length; lobes of segments broadly lanceolate or
lanceolate, 0.8—2.0 mm wide; involucre usually 2.5—3.0 mm in dia;
ligule 2.5—3.0 mm wide .............. 10. A. distans Waldst. and Kit.
Rachis 1.0—-1.5(2.0) mm wide, with pinnately parted or entire
intermediate lobes from middle or almost from base; lobes of
segments lanceolate, 0.3-1.0 mm wide; involucre usually 3.0—3.5
mm in dia; ligules 2.0-2.4 mm wide ......... 11. A. stricta Schleich.
Rachis of leaves 1-2 mm wide; lobes of leaf segments 0.5—1.5 mm
NIGOh aie late. wik .. deh tide ce egect, wi 3. A. millefolium L.
Rachis 0.5—1.0 mm wide; lobes of leaf segments narrower, 0.1—0.6
PUES) «AV L008 dS eGaeh Vasyl, APR ORC I EES enes 11.
Rhizome thick, woody, many-headed. Plants densely lanate-villous
Bh, Setiidn BLD. RAS). AED, MO Blarase 9. A. bucharica Winkl.
Rhizome slender, not woody; pubescence different .................. 12,
Rachis of leaves about 1 mm wide; leaf segments somewhat distant;
involucral bracts with wide dark reddish-brown or reddish-brown
membranous Dorden il, ..golases5i, asic Bisickds sect lene 4. A. sudetica Opiz.
Rachis 0.5—1.0 mm wide; leaf segments approximate, membranous
border of involucral bracts not wide, usually light colored ....... 13.
Ligules pink, red, or very rarely white ........... 8. A. asiatica Serg.
DiigalesiyeHowish- White OF. WIC. ...,....<...-.scccsnesnersscanananeansacceees 14.

72

73

66

15.

16.

22.

Lobes of leaf segments lanceolate or linear-lanceolate, up to 0.5—
0.6 mm wide; involucre oblong, (2.0)2.5—3.0 mm in dia; ligules
1,522.0 mm: long and !15=2:S timmy Wide. Pee ee etek ene
BE BRUNE iL NOR TS AE 6. A. pannonica Scheele.
Lobes of leaf segments linear-setose, linear, less often linear-
lanceolate or lanceolate, usually 0.1-0.2 mm wide; involucre ovate-
cylindrical, less often oblong, 1.5—2.5(3.0) mm in dia; ligules 0.9-
1.5 mm long and 1-2 mm wide ........ 7. A. setacea Waldst. and Kit.
Plants tall, up to 120 cm high; middle cauline leaves broad oblong-
lanceolate, 8—18 cm long; involucre oblong-obconical .................0.
ils Ail on schva dee EA: Sees 13. A. filipendulina Lam.
Plants small; cauline leaves narrower, 2-6 cm long; involucre of
Gifferent Shape tincicscscestsecessovecesatos DMG Ie Mebetedc ere Rena aR ce 16.
Whole plant densely sericeous-tomentose; capitula on thick (from
pubescence) peduncles, in very dense, strongly convex, often

hemisphericalicorymbs 2221)..20. 2212g..2ae 14. A. coarctata Poir.
Pubescence different; peduncles not thick; corymbs less dense and
CORVER. | Nees, DAW RRNA ARR ee Lar
Rachis of leaves in upper half with intermediate teeth between
basal segments :2ts..emain eee ae ee ee ee 18.
Rachis lacking intermediate teeth, ..jcc2:2. ee eee ee. jee

Leaf segments linear; capitula in dense corymbs; ligules 1.0—1.5
mm long; achenes cuneate-oblong, 1.1—1.2 mm long. Plants growing
in Transcaucasia and Crimea ..............::00000 2. A. neilreichii Kern.
Leaf segments oblong; capitula in somewhat lax corymbs; ligules
0.8-0.9 mm long; achenes oblong, about 1.5 mm long. Plants
growing in the south of Ukraine ........ 21. A. micranthoides Klok.
Middle cauline leaves petiolate. Arenicolous plants ................:0:0+
Ns BE ies Riewsde dence donskes UIE ARO 18. A. micrantha Willd.
Middle cauline leaves sessile. Steppe and mountainous plants....20.
Stem divaricately branched from base, very rarely simple. Involucre

more or less hemispherical ...................005 16. A. leptophylla MB.
Stem simple or sometimes weakly branched above. Involucre oblong
OP OVOIE! Ph tA SR ER, Ee hs a Be) PAN

Lobes of leaf segments not cuneate, linear, oblong-linear, less often
lanceolate; capitula on shorter peduncles in dense corymbs; ligules
golden or bright yellow ............ eee 19. A. biebersteinii Afan.
Lobes of leaf segments linear-cuneate; capitula on more or less
peduncles in lax corymbs; ligules light yellow .................0..
Cee TRCHNY. «SAAS, RT 20. A. cuneatiloba Boiss. and Buhse.
Stem sulcate-ribbed; lobes of leaf segments almost cylindrical;
involucral bracts weakly pubescent to more or less glabrous ............
inewiiecesttysenvnivenSvavesne wn resiityieedess a ROOM: Met eee 25. A. tenuifolia Lam.

74

ty

30.

67

Stem cylindrical; lobes of leaf segments more or less flat, not
cylindrical; involucral bracts more or less densely pubescent .......

TRAE ERE Ree ONL EET eee ON Caile ss MAD oad GEA BRIM de cas esees 23.
Involucre (4)5—6 mm in dia, hemispherical or subglobose .............
SOR SRR BOR AER FORE 24. A. vermicularis Trin.
Involucre narrower, different in Shape ..............ccescceseeeeeeeeneeeees 24.
Involucre ovoid-cylindrical, 3-4 mm in dia leaf segments 0.5—1.0
mim long 28. SSS Ee na kB 26. A. schischkini Sosn.
Involucre obovoid, 2-3 mm in dia; leaf segments 1.0—1.5(2.0) mm
LOT ER a EPG thas Bah isdeleileaes 25.

Involucre appressed-hairy, 2.5—3.0(3.5) mm long; stem appressed-
hairy, finely tomentose; leaf segments more or less glabrous, usually
approximate, with rotund-obovate or obovate, toothed lobes ..........
rei tides teehee Ree tete s Map teae ee ate es AAR RE 22. A. wilhelmsii C. Koch.
Involucre somewhat sparsely tomentose, 3.0-3.5(4.5) mm long; stem
and leaves more or less densely pubescent; leaf segments somewhat
distant, with narrower, spatulate or oblong-obovate, prickly-toothed

ls) pple col be eieeeeie beh LEM REY er Be Eo 23. A. kermanica Gand.
CAVES TCRNITE 205 SL RUN GRR TS RR Acts Zr.
Leaves pinnately lobed, pinnately parted or pinnately cut ......... 38.
Leaves punctate-glandular hairy on both sides (glandular hairs in
oe) a oy ee trea a A gt Bea eo operas Pepe Ee «04 Can wie PARE 28.
Leaves not punctate-glandular-hairy; if glandular-hairy, then hairs
AOt IT EPC NOH AEN NY. GANS, AONE TR GON 30.

Involucral bracts densely pubescent, yellowish-green in middle, with
yellow membranous margin, or margin yellow with narrow dark
brown Border NAAN RO 35. A. salicifolia Bess.
Involucral bracts sparsely pubescent, green in middle with dark
bIOWN thembranGus POTEET ANF NR. 4 aR wai NE SAAR aN. 29.
Leaves grayish-green, densely crisped-hairy, involute. Transcaucasia
Perit trcet re Ree ee eT a ee) ot tte ae 36. A. sedelmeyeriana Sosn.
Leaves flat, green, less densely crisped-hairy. European Part, north
of Soviet Central Asia, Siberia .................. 37. A. cartilaginea Ldb.
Cauline leaves semiamplexicaul, obtuse. Carpathians....................
Sivcestedcasn devdec Crate eee EME sinc pienOD 27. A. lingulata Waldst. and Kit.
Cauline leaves sessile, not amplexicaul, acute .............ccsceeeeeeees St.
eaves covered * with “shartcrisped Haire sia... ici ets sd len.
AN nh Di be as 38. A. septentrionalis (Serg.) Botsch.
Leaves glabrous or covered with long straight hairs ................. a2.
Plants grayish-green from dense pubeScence..............:ccscceeeeeeees
Poe ae ella ec etee eeeLL at eS Rents ae 29. A. griseo-virens Alb.
Plants green, glabrous or with scattered hairs .............:cccessseeees 33.

aS

68

Bi.

38.

39.

42.

Plants entirely glabrous; stem branched, all branches terminating in

eapitula. Transcaucasia) 2% di. chy et cao 33. A. sachokiana Sosn.
Plants pubescent; stem simple, branched only in inflorescence; if
branched; then,axillary- branches Sterile..c...40.d:s4-)-srcsessaraupeetos-eo 34.

Leaves hairy (at least beneath, and along veins above) with
rubiginous, long, straight haws.Caueasus) :..0:icicyssascssacenerss< coor agetnee
PE REAR ETS ile BRS ates al 28. A. biserrata MB.
Leaves glabrous, or with colorless, shorter hairs ....................- 35.
Stem branched, axillary branches sterile, long. Caucasus ..................
sje tec cee Rn i 32. A. ptarmicifolia (Willd.) Rupr. ex Heimer!
Stem simple, branched only in inflorescence; if branched in other
part also, then axillary branches sterile and short ..................... 36.
Involucral bracts green in middle, with wide yellowish membranous
margin, or membranous part yellow, but with narrow dark brown
border; receptacular scales yellow at apex; ligulate florets white.
Eastern Siberia, Far East .......... 31. A. acuminata (Ldb.) Sch. Bip.
Involucral bracts green in middle, with dark brown membranous
margin; receptacular scales dark brown at apex; ligulate florets

White or white with, VIOlet Stripe: .-.°s1;..0.-0--sesqeeeete eater ree a7;
Rhizome short; leaves lanceolate. Kamchatka, Commander and Kuril
Slandsis 3. 2h12 Beek Aa. ie eee) 34. A. macrocephala Rupr.

Rhizome creeping, leaves linear or linear-lanceolate. European Part
etc ce OLB SENG S RL Gy wok Eup asec naa es al a a 30. A. ptarmica L.
Capitula solitary; leaves twice or thrice pinnately cut. Carpathians
PEE, LUIS ans, ee, eee 45. A. schuri Sch. Bip.
Capitula in corymbose; leaves pinnately lobed, pinnately parted, or
pinnately cut, sometimes twice pinnately parted, or twice or thrice
pinniatel yr cut sisi", kia seee as ret re aE etnias Ce
Leaves thrice pinnately cut; secondary lobes entire ...........0......e0
STR cl ews eal LNG, seca pear Beinn a vate, She ae 5. A. borealis Bong.
Leaves pinnately lobed, pinnately parted and pinnately cut to twice
pinnately part or twice pinnately cut; all lobes and segments inequally

SCRPAGEEMEATE Ys scxsvsnno ote teecheee ctor bos Sew ne ath ane eee ata 40.
Membranous part of involucral bracts dark-brown ................... 41.
Membranous part of involucral bracts yellow or yellow with dark
brown Dorderiitins.f citadel es See 43.

Leaves punctate-glandular hairy above. Far East, Kamchatka .....
ai Setalniinn nh eter ae Mee 44. A. camtschatica Rupr. ex Heimer
Leaves not punctate-glandular hairy .............ccsccssssessesesscesecceeenees 42.
Lowermost leaf segments with 1-2 large teeth or lobes on each side
besides small teeth. Mainly plant of plains, sometimes extending
Into MOUNtaINGH 25 Leese an Le 39. A. impatiens L.

69

+ Lowermost leaf segments, with 1 large tooth along upper margin
(besides small teeth), or lacking large tooth but only with similar
small teeth. Subalpine or alpine plants................cceeeseeseeseeseeeeees
_ gtr A pte a NG RE 40. A. ledebouri Heimer!

43. Leaves pinnately cut; upper leaf segments with equal small teeth,
sometimes some segments also with 1 large tooth or lobe along
upper margin; lower segments, always with 1-2 large teeth or lobes
on each side besides small teeth .................... 39. A. impatiens L.

+ Leaves pinnately lobed, pinnately parted, or pinnately cut, but then
their segments unequally toothed or pinnately lobed, pinnately parted,
or pinnately cut, with large number of secondary lobes or segments
PRE. UR CESS CEG RO eee, BC a beer eee, 44.

44. Capitula large (up to 7.5 mm long and 7 mm in dia), hemispherical;
ligules large (up to 3.5 mm long and 3.5 mm wide), white or pink
FELT. ease TROL, RLS 42. A. japonica Heimer!

+ Capitula smaller, semiovoid or oblong; ligules smaller or weakly
GeVelo ped) Witte) i AAG UN. Leaded anAlnia aud Be, eg. ee 45.

45. Leaves pinnately cut (often twice pinnately cut), densely punctate-
glandular hairy; capitula oblong; membranous part of involucral
bract yellow or yellow with narrow dark brown border; florets
covered with capitate glandular hairs; ligules exceeding disk or
somewhat undeveloped (in subtubular flowers) and not exserted
Om them with lat ger teeth atiapex iii sii. iien. eQuideaaatnss
REALE IRL, BING. EAR 43. A. ptarmicoides Maxim.

+ Leaves pinnatilobate or pinnately parted (sometimes twice-pinnately
lobed or twice-pinnately parted), not punctate-glandular hairy or
with few such hairs; capitula semiovoid; membranous part of
involucral bracts yellow with wider dark brown border; capitate
glandular hairs on florets very few or lacking; ligules exceeding
disk, with small apical teeth «00.0.0... cece eeeeeees 41. A. alpina L.

Section 1. Millefoliatae DC. Prodr. VI (1837) 24.—Millefolium

76 C. Koch, Syn. Fl. Germ. (1837) 372.—Ligules white, pink, red or light

yellow; leaves pinnately cut or pinnately parted, segments neither
transverse nor imbricate.

1. A. nobilis L. Sp. pl. (1753) 899 and (1763) 1268; MB. FI.
taur.-cauc. II, 339 p. p.; DC. Prodr. VI, 26 p. p.; Ldb. FI. Ross. II, 2,
533 p. p.; Boiss. Fl. or. III, 257 excl. B. ochroleuca; Schmalh. FI. I,
65, excl. B. ochroleuca; O. and B. Fedtsch. Perech. Rast. Turk. IV,
181; Grossh. Fl. Kavk. IV, 124; Kryl. Fl. Zap. Sib. XI, 2725.—A.
odorata Pall. Reise, I (1774) 379, II, 103 and III, 556, non L.—A.
corymbifera S. Gmelin, Reise, I (1774) 137.—Ic.: S. Gmelin, Reise, I,

Tif

70

tab. XXV, f. 1; Rchb. Ic. Fl. Germ. XVI, f. 1024, II; Syreistsch. III.
Fl. Mosk. Gub. III, fig. on page 264; Javorka and Csapody, Iconogr.
Fl. Hung. f. 3731 s, f.—Exs.: GRF No. 1069; Fl. pol. exs. No. 644a,
b; Fl. Ital. exs. No. 1778.

Perennial. Rhizome short, many-headed; plants grayish-green, more
or less densely lanate-flocculose, sometimes subglabrous; stems few,
3-6, less often up to 12, or solitary, (15)25—35(50) cm high, erect or
slightly ascending, usually sinuate, simple or branched near top, usually
densely leafy, finely sulcate, weakly angular and more strongly
pubescent below. Leaves ovate or oblong-elliptical, twice pinnately
cut, punctate-alveolate on both sides, somewhat antrorse, flat, (2)3-6
cm long, sessile, only lower cauline leaves and those on nonflowering
branches petiolate; midrib of leaves narrow, with more or less frequent
intermediate cuspidate lobes between middle part and apex (between
main segments), lanceolate, deltoid or linear, entire or pinnatifid; leaf.
segments linear, up to 10-15 mm long, few, distant (especially in
lower part of lower cauline leaves), divergent, pinnately cut into entire
or often pinnatifid lobes, with small, 0.5-1.0(2.0) mm long, lanceolate
or linear, acute lobes, terminating in cartilaginous cusp. Capitula in
dense, convex, compound corymbs; receptacle more or less convex,
often conical at fruiting. Involucre ovoid, less often oblong-ovoid,
2.0—3.0(3.5) mm long, 1.5—2.0(2.5) mm in dia; involucral bracts pale,
carinate, oblong, with narrow white or less often brown (var. marginata
C. Koch) membranous margin; bracts membranous, lanceolate, white,
transparent, slightly shorter than tubular florets. Ligules of outer florets
white or yellowish-white, reniform-rotund, subelliptical or semicircular,
truncate at apex, unequal 3-toothed, (0.6)1.0-1.6(1.9) mm wide, 1/3-
2/7 as long as involucre. Achenes oblong, about 1 mm long. Flowering
June to September.

In steppe and forest-steppe zones on chernozems and saline soils,
solonetzes, chalky and stony outcrops, in feathergrass and sheep’s fescue
steppes, steppe meadows, old fields, by roadsides, more rarely in thick-
ets of shrubs, along forest edges and in glades, as well as in riverine
meadows. In forest and desert zones as an introduced and weed plant,
often on railway embankments and in vacant lots and squares in cities.—
European part: Volga-Dnieper, Bessarabia, Middle Dnieper, Black Sea
Region, Crimea, Volga-Don, Lower Don, Trans-Volga Region, Volga-
Kama (south); Caucasus: Ciscaucasia, Dagestan, Western
Transcaucasia; Western Siberia: Upper Tobol, Irtysh, Altai; Soviet
Central Asia: Aralo-Caspian, Lake Balkhash Region (north). As an
introduced plant in Karelia-Lapland, Ladoga-Ilmen, Eastern Trans-
caucasia, mountainous Turkmenia, Syr-Darya, Tien Shan. General
distribution: Atlantic and Central Europe, western Mediterranean

78

71

Region (north), Balkans. Described from Central Europe. Type in
London.

Note 1. Varies markedly in the division of leaf segments, degree
of pubescence and other characters; hybridizes with A. neilreichii Kern.,
A. leptophylla MB., A. biebersteinii Afan., A. setacea Waldst. and
Kit., A. millefolium L. s. |. and a few other species of this genus.

Note 2. E. Boissier (op. cit. p. 256) reports the distribution of A.
odorata Koch in the Caucasus “between Vladikavkaz and Alagir
(Rupr.!)”. Following Boissier, this species was also reported for the
Caucasus by Trautvetter (Tr. Peterb. Bot. Sada, VIII, 2, p. 451). The
plant collected by Ruprecht in October 1860, between Vladikavkaz
and Alagir and identified by Boissier as A. odorata Koch, is preserved
in the herbarium of the Botanical Institute, Academy of Sciences of
the USSR. This plant differs from A. odorata Koch by dentate and not
entire lobes of the leaf segments and by narrower capitula. Apparently, it
is a hybrid between A. nobilis L. X A. setacea Waldst. and Kit.

2. A. neilreichii Kern. in Oest. Bot. Zeitschr. XXI (1871) 141;
Afan. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX, 363.—A.
ochroleuca auct. fl. Ross. non Ehrh.: Grossh. Fl. kavk. IV, 124.—A.
nobilis auct., p. p.: MB. Fl. taur.-cauc. II, 339; Ldb. Fl. Ross. II, 2,
533.—A. nobilis var. neilreichii (Kern.) Beck. Fl. Nied. Oest. (1890)
1198; Hayek in Hegi, Ill. Fl. VI, 2, 567 (subvar.).—A. nobilis B.
ochroleuca Boiss. Fl. or. I], 257, non A. ochroleuca Ehrh. Schmalh.
Fl. II, 65.—Ic.: Mag. Bot. Lapok, XII, tab. VI, f. la, b, c; Javorka and
Csapody, Iconogr. Fl. Hung. f. 3731s.—Exs.: Fl. Hung. exs. No. 578;
Fl. exs. Austro-Hung. No. 997; Fl. Bohem. et Morav. exs. No. 574; FI.
exs. Reip. Bohem. Sloven. No. 1184.

Perennial. Rhizome short, branched; plants grayish-green, usually
densely lanate-floccose; stems few, usually 2—4 or solitary, (20)25—50
cm high, erect or slightly ascending, straight or weakly flexuous stem,
simple or branched above, more or less sparsely leafy, finely sulcate,
cylindrical, often more weakly pubescent and sometimes almost glabrous
below, weakly angular, and strongly pubescent above. Leaves oblong-
elliptical, less often ovate, twice pinnately cut, punctate-alveolate on
both sides, somewhat antrorse, flat, 3-7 cm long, sessile, only lower
cauline leaves and leaves on nonflowering branches petiolate; midrib
narrow, usually with distant, cuspidate lobes (between large basal
segments) from middle to apex; lobes lanceolate or linear, entire or
pinnatifid; leaf segments linear, 15(20) mm long, few, distant (especially
in lower part of lower cauline leaves), divergent, pinnately cut into
entire or pinnatifid lobes, with small, 0.5-1.5(2.0) mm long, linear,
oblong-linear or lanceolate, subacute lobes, often terminating into very

79

72

short cartilaginous cusp. Capitula in dense, convex, compound corymbs.
Receptacle conical, less often somewhat convex, sometimes oblong-
conical at fruiting. Involucre oblong-ovoid, less often ovoid, 3.0—3.5
mm long and about 2 mm in dia; involucral bracts pale, carinate,
oblong, with narrow, brownish or brown, less often white membranous
border. Receptacular bracts membranous, lanceolate, white, transparent,
slightly shorter than tubular florets. Ligules of outer florets light yellow,
semicircular, subelliptical or rotund-reniform, truncate at apex,
unequally obtusely 3-toothed, less often almost entire, (1.0)1.4—2.0
mm wide, 1.0—1.5 mm long. Achenes 1.1—1.2 mm long, cuneate-oblong.
Flowering June to August.

Along open stony and stony-clayey slopes of mountains and hills
in the foothills, as also on stony outcrops, in shrub thickets, along
forest edges, thin forests and roadsides.—European part: Crimea
(mountains); Caucasus: Eastern and Southern Transcaucasia. General
distribution: Central Europe, Balkans-Asia Minor. Described from
Austria. Type in Vienna.

Note. It is a yellow-flowered mountain race of A. nobilis L. group,
related to the latter; it is well distinguished from it by the yellow-
colored ligules of the outer florets. Other characters: larger and narrower
capitula, larger ligules, less acuminate lobes of the leaf segments, fewer,
more distant, intermediate teeth of midrib, slightly larger achenes, etc.,
are not always clearly expressed, and are also found sometimes in A.
nobilis L. in one or another combinations. Intermediate forms, appar-
ently partly of hybrid origin, are also found at places of contact be-
tween the ranges of these two races.

It hybridizes with A. coarctata Poir. (A. dobrogensis Prod.) and a
few other species.

3. A. millefolium L. Sp. pl. (1753) 899; MB. FI. taur.-cauc. II,
338; Dc. Prodr. VI, 24, p. p.; Ldb. Fl. Ross. II, 2, 531 p. p.; Boiss. Fl.
or. III, 255 excl. 8. Janata; Schmalh. FI. II, 64 p. p.; O. and B. Fedtsch.
Perech. Rast. Turk. IV, 181 p. p.; Grossh. Fl. Kavk. IV, 124; Kom.
and Alis. Opred. Rast. Dal’nevost. Kr. II, 1027; Sugawara, Ill. Fl.
Saghal. IV, 1735 excl. synonymo; Kryl. Fl. Zap. Sib. XI, 2721.—A.
millefolium subsp. millefolium (L.) Hayek in Hegi, Ill. Fl. VI, 2, 569.—
A. millefolium subsp. eu-millefolium (L.) Hayek Prodr. Fl. Balcan.
639; Szafer, Kulcz., Pawl. Rosl. Polskie, 684 (forma typ.).—A.
millefolium var. macrophylla Serg. in Kryl. op. cit. 2722.—A. magna
auct. fl. Ross. non L.—A. ossica C. Koch, in Linnaea, XXIV (1851)
323 sec. Boiss. 1. c.; Trautv. in Tr. Peterb. Bot. Sada, VIII, 2, 45.—
A. subhirsuta Gilib. Fl. Lithuan III (1831) 217.—Millefolium officinale
and M. vulgare Giildenst. Reise. II (1791) 39, 166, etc.—Ic.: Syreistsch.

80

73

Ill. Fl. Mosk. Gub. III, Fig. on p. 264; Fedtsch. and Fler. Fl. Evrop.
Ross. Fig. 970; Kom Sbor. Sushka i Razved. Dek. Rast., 3rd ed., Fig.
39.—Exs.: Fl. exs. Austro-Hung. No. 988; Fl. Bohem. et Morav. exs.
No. 1281; Fl. exs. Bohem. Sloven. No. 941; Pl. Finn. exs. No. 338;
Eest. taimed. No. 135; Fl. pol. exs. No. 642; fl. Lithuan exs. No. 100.

Perennial. Rhizome slender, creeping, branched; whole plant more
or less covered with fine white hairs; stems few or solitary, usually
weakly pubescent (finely floccose), (5)20-—60(120) cm high, erect or
ascending from base, erect, less often flexuous, simple or branched
above, cylindrical, finely sulcate, with short leafy branches in axils of
upper and middle cauline leaves. Leaves lanceolate, oblong-lanceolate,
or almost linear, punctate-alveolate, twice or thrice pinnately cut, with
numerous more or less remote segments (1.5—10.0 mm apart); lower
cauline leaves and leaves of nonflowering branches 10-40 cm long,
0.8-5.0 cm wide, rachis 1-2 mm wide, leaves usually in upper part
with solitary teeth between basal segments; lobes and teeth lanceolate,
less often linear, 0.5—-1.5 mm long, 0.3—0.4(0.5) mm wide, terminating
in short cartilagenous cusp. Capitula in numerous, unequal, compound
corymbs, 2-15 cm in dia. Involucre oblong to almost ovoid, 3—4(6)
mm long, (2)3—4(5) mm in dia; involucral bracts green, carinate, with
prominent midrib, membranous along margin, often brownish; bracts
ovate to oblong-elliptical, membranous, floccose above, with scattered
hairs on dorsal surface. Ligules of outer florets white, pink, or red.
(1)2-4 mm long, 1.5-3.0(4.5) mm wide, more or less rotund, 2-3-
toothed at apex, limb a half as long as involucre; tubular florets up to
20, glandular-hairy on outside. Flowering July to October.

In forest, forest-steppe and steppe zones in dry-bottom forest-
meadows, in steppe meadows and meadow-covered mountain slopes,
in open thin forests, on deposits, along borders of fields, in wastelands
and along roads.—European part: All regions; Caucasus: All regions,
except Talysh; Western Siberia: Ob River Area, Irtysh, Upper Tobol,
Altai, Eastern Siberia: Angara-Sayans, Yenisei, Dauria, Lena-Kolyma;
Far East: Introduced in Kamchatka, Sakhalin, Ussuri; Soviet Central
Asia: In the north of Aralo-Caspian Region, Lake Balkhash Region,
Tien Shan, introduced in the southern mountains and in oases. General
distribution: Scandinavia, Atlantic, Central and Southern Europe,
Mediterranean Region, Balkans-Asia Minor, Iran, Dzhungaria-
Kashgaria, Mongolia, introduced to other continents. Described from
Western Europe. Type in London.

Note. A highly polymorphic species, strongly variable in size,
degree of pubescence, division of leaves, size and shape of capitula,
shape and size of ligules of outer florets, etc. There are many forms
of varying taxonomic significance, usually vaguely delineated from

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each other. Their correct assessment is possible only through detailed
study in nature and a monographic study of all members of section
Millefoliatae. In this context we are refraining from distinguishing and
describing individual races, even those like A. magna auct. Fl. Ross.,
and we present here as separate races, only a few which are already
included in a flora, for example, A. sudetica Opiz. and A. pannonica
Scheele. It hybridizes with A. distans W. and K.., A. stricta Schleich.,
A. biebersteinii Afan. and other species.

4. A. sudetica Opiz. in Andre, Hesperus (1812) and in Flora, V
(1822) 635; Szafer, Kulcz. Pawl. Rosl. Lolskie, ed. I, 624; Klok. in
Vizn. Rosl. URSR, 543.—A. millefolium var. alpestris Vimm. and Grab.
Fl. Sil. II (1829) 169; Koch, Syn. Fl. Germ. ed. I, 373.—A. millefolium
subsp. sudetica (Opiz.) Weiss in Koch, Syn. Fl. Germ. ed. III, II (1902)
1404; Hayek in Hegi, Ill. Fl. VI, 2, 571 Szafer, Kulcz. Pawl. op. cit.
ed. II, 684.—Ic.: Javorka and Csapody, Iconogr. Fl. Hung. f. 3739.—
Exs.: Fl. exs. Austro-Hung. No. 989; Pl. Pol. exs. No. 174.

Perennial. Rhizome slender, creeping; plants weakly pubescent with
fine white hairs; stems solitary or few, erect or ascending from base,
18-40 cm high, simple or weakly branched above, erect or slightly
flexuous, cylindrical, finely sulcate. Leaves light green, lanceolate or
oblong-lanceolate, midrib about 1 mm wide, entire or less often with
isolated teeth between main segments; middle and upper cauline leaves
sessile, upward directed 5—7(9) cm long, 1-2 cm wide, thrice pinnately
cut; segments slightly distant, pinnately cut into linear-lanceolate, acute,
entire or subentire, up to 0.5 mm-wide lobes terminating into short
cartilaginous cusp, lowermost segments semi-amplexicaul (auricles);
lower cauline leaves and leaves of nonflowering branches petiolate,
larger, latter up to 25-30 cm long and 3(6) cm wide, with linear lobes.
Capitula in compact, unequal, dense, convex compound corymbs,
sometimes almost globose on flowering. Receptacle conical. Involucre
oblong, (4)5-6 mm long, (2.0)2.5-3.0 mm in dia; involucral bracts
more or less pubescent, greenish, with wide dark brown border, weakly
carinate; outer bracts oblong-ovate or oblong, others oblong-lanceolate,
with prominent yellowish vein on dorsal surface; bracts membranous,
slightly shorter than tubular florets, dark brown at apex and along
margin in upper part, sparsely white-hairy along margin on outside.
Ligules of outer florets white or pink, (2.0)2.5-3.5 mm long,
subelliptical or semicircular, truncate at apex, obtusely 3-toothed.
Flowering VII-IX.

In alpine and subalpine meadows, stony and sandy places, less
often in beech forests.—European part: Upper Dniester. General dis-
tribution: Alps, Sudeten, Carpathians. Described from Central Europe.
Type in Prague?

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Note. Extremely variable in the degree of pubescence of the stem and
leaf, as well as in the intensity of the color of the involucral bracts.

5. A. borealis Bong. in Mém. Acad. Sc. St.-Pétersb. ser. 6, II
(1832) 149; Torr. a. Gray, Fl. N. Americ. II, 409; Rydb. in N. Americ. —
Fl. 34, 220; Hulten, Fl. Kamtch. IV, 166.—A. millefolium auct. non L.:
Less. in Linnaea, VI (1831) 167; Bongard, op. cit. 148.—A. millefolium
B. occidentalis DC. Prodr. VI (1837) 24; Ldb. Fl. Ross. II, 2, 532;
Herder in Bull. Soc. Nat. Mosc. XXXVIII, 2, 404.—?A. anethifolia
Fisch. in herb.; Trautv. in Tr. Peterb. Bot. Sada, VIII, 2, 450.—Ptarmica
borealis DC. |. c. 21; Ldb. 1. c. 527.

Perennial. Rhizome short, creeping; stems solitary, less often few,
17-27(45) cm high, erect or ascending from base, straight or weakly
flexuous, simple, strong, thicker, usually more than 2.5 mm in dia,
sulcate, cylindrical below, weakly angular above. Leaves thrice
pinnately cut, oblong-lanceolate or oblong-linear, more or less densely
pubescent; midrib entire, 0.7-1.0 mm wide; segments numerous,
somewhat distant, middle and upper cauline leaves sessile, 4-8 cm
long and 0.6-1.0(1.5) cm wide, lower leaves and leaves of non-
flowering branches petiolate, latter 12—15(20) cm long and to 2 cm
wide; segments twice pinnately cut into lanceolate-linear or linear lobes,
1.5-3.0 mm long and 0.2-0.4 mm wide, terminating into short
cartilaginous cusp. Capitula on densely villous peduncles, in dense,
many-flowered, compound unequal corymbs. Involucre ovoid, (4)5—6
mm long and about 3 mm in dia, usually densely villous-pubescent;
involucral bracts light colored, yellowish-green, carinate, with thick
prominent midrib and wide dark-brown membranous border; outer bracts
ovate, others oblong-ovate; bracts membranous, oblong-lanceolate,
acute, mostly brownish at apex. Ligules white, from roundish-obovate
to subelliptical, (2.5)3.0-4.0(4.5) mm long and about 2.5 mm wide,
indistinctly obtuse 3-toothed at apex or almost entire. Achenes about
2.5 mm long, oblong or oblong-cuneate. Flowering June to August.

Far East: Kamchatka, Kuril Islands. General distribution: Aleutian
Islands, Alaska, northern Canada. Described from Sitka Island. The
type ought to be in Leningrad.

Note. Only two specimens of A. borealis Bong., collected from
the USSR, are preserved in the herbarium of the Botanical Institute,
Academy of Sciences of the USSR, one with the label “Petropawlowsk.
Dr. Mertens” and the other with “Igir 1833.” Besides, one more milfoil
specimen, collected by Merck on the Kuril Islands, is also preserved
there with Fischer’s identification “anethifolia m.,” which is possibly
the type of this species, named but not described by Fischer. This
plant, like some of the specimens from Sitka Island, differs from A.

76

borealis Bong. in the shape and division of the leaves, weak pubes-
cence and other characters. Further study is required to solve the
question of whether A. anethifolia Fisch. in herb. is a separate species
or should be considered a synonym of A. borealis Bong., as suggested
by some botanists.

6. A. pannonica Scheele in Linnaea, XVIII (1844) 471; Szafer,
Kulcz. Pawl. Rosl. Polskie, ed. I, 624; Klok. in Vizn. Rosl. URSR,
544.—A. millefolium ssp. pannonica (Scheele) Hayek in Hegi, Ill. Fl.
VI, 2, 571; Szafer, Kulcz. Pawl. Rosl. Polskie, ed. II, 684.—A.
millefolium y. lanata Koch, Syn. Fl. Germ. ed. I (1837) 373.—A.
millefolium var. setacea Ldb. FI. Ross. II (1845-1846) 532, p. min. p.;
Schmalh. FI. II, 64, p. p.—Ic.: Javorka and Csapody, Iconogr. FI.
Hung. f. 3740.—Exs.: Fl. exs. Austro-Hung. No. 992; Fl. Hung. exs.
No. 575; Dorfler, Herb, Norm. No. 4540; Fl. Bohem. et Morav. exs.
No. 798; Fl. Polon. exs. No. 47.

Perennial. Rhizome slender, creeping, branched; plants more or
less densely covered with long, fine, white hairs; stems solitary or few,
grayish, usually lanate-villous in lower part, less often turning red,
(15)25-50(70) cm high, erect or ascending from base, straight or weakly
flexuous, simple, cylindrical, finely sulcate, sometimes with short and
densely leafy branches in axils of middle and upper cauline leaves.
Leaves twice pinnately cut, linear-oblong or linear-lanceolate, often
grayish from especially dense pubescence along midrib in lower part,
0.5—1.0 mm wide; middle and upper cauline leaves up to 8 cm long
and 1 cm wide, with numerous approximate segments, pinnately cut
into linear-lanceolate or lanceolate up to 0.5—0.6 mm wide lobes,
terminating into cartilaginous cusp, lower segments larger,
semiamplexicaul (auricles); lower cauline leaves long-petiolate, up to
12 cm long, 1.5—2 cm wide, leaves of nonflowering branches up to 35
cm long and 2 cm wide. Capitula in dense, compact, unequal corymbs.
Receptacle conical. Involucre oblong, 4-5 mm long, (2)2.5-3 mm in
dia; involucral bracts hairy, light colored, greenish-yellowish, weakly
Carinate, with prominent midrib and narrow, usually light brown
membranous border; outer bracts oblong-ovate, others oblong or
lanceolate-oblong; bracts membranous, whitish, oblong-lanceolate,
slightly shorter than tubular florets, with fine hairs on outside, mainly
in upper part. Ligules of outer florets yellowish-white or white, 1.5—
2.0 mm long, 1.5-2.5 mm wide, rotund-reniform or semicircular, less
often subelliptical, apically truncate, obtusely toothed. Flowering June
to August.

In steppes and forest-steppes, on herb and stony siopes of ravines,
hills, and mountains, by roadsides, in vineyards.—European part:

77

83 Upper Dniester, Bessarabia, Black Sea Region. General distribution:
Central Europe. Described from Hungary (vicinity of Pesta). Type in
Vienna.

Note. The species is close to A. setacea Waldst. and Kit., from
which it differs mainly by the broader lobes of the leaf segments and
the larger involucre and ligules.

It hybridizes with Chrysanthemum wmillefoliatum L.
(Chrysanthemoachillea carmen-sylvae Prod. Fl. Rom. 1000) and some
species of the genus Achillea.

7. A. setacea Waldst. and Kit. Pl. rar. Hung. I (1802) 82; Willd.
Sp. pl. III, 3, 2212; MB. FI. taur.-cauc. II, 338; DC. Prodr. VI, 25;
Boiss. Fl. or. III, 256; Hayek in Hegi, Ill. Fl. VI, 2, 573; Grossh. FI.
Kavk. IV, 124; Kryl. Fl. Zap. Sib. XI, 2275; Klok. in Vizn. Rosl.
URSR, 1544; Szafer, Kulcz. Pawl. Rosl. Polskie. ed. II, 684.—A.
millefolium var. setacea (Waldst. and Kit.) Koch, Syn. Fl. Germ. ed.
I (1837) 373; Ldb. FI. Ross. II, 2, 532, p. p.; Schmalh. Fl. II, 64 p. p.;
Weiss in Koch, Syn. Fl. Germ. ed. III, 2, 1404 (subsp.); O. and B.
Fedtsch. Perech. Rast. Turk. IV, 181; Majevsky, Fl. ed. 8, 578.—A.
odorata a. Willd. Tract. de Achilleis (1789) 42.—A. odorata Rchb.
Fl. exs. (1830) 229, non L.—? A. mollis Andr. in Rogovicz, Obozrenie
(1869) 302, nom. nud.; Trautv. in Tr. Peterb. Bot. Sada, VIII, 2, 451.—
Ic.: Walldst. and Kit. loc. cit. tab. 80; Rchb. Ic. Fl. Germ. XVI, f.
1028; Hegi, loc. cit. tab. 294 a, b, c; Javorka and Csapody, Iconogr.
Fl. Hung. f. 3737f; Fl. Yugo-Vost. Fig. 677.—Exs.: GRF No. 3987;
Rchb. Fl. Germ. exs. No. 43; Fl. exs. Austro-Hung. No. 993; Fl. Hung.
exs. No. 81; Fl. Bohem. et Morav. exs. No. 1162; Fl. exs. Reip. Bohem.
Sloven. No. 939, I-V.

Perennial. Rhizome slender, creeping, branched; stems less
numerous, less often solitary, grayish or grayish-green from fine, long,
intertwined, white hairs, usually lanate-vilous below, (10)20—35(70)
cm high, erect or ascending from base, straight, sometimes weakly
flexuous, simple or rarely weakly branched above, cylindrical, finely
sulcate often with short leafy branches in axils of middle and upper
cauline leaves. Leaves twice or thrice pinnately cut, linear-lanceolate,
less often linear-oblong, grayish or grayish-green, less often gray,
usually densely pubescent, especially in lower part of narrow (0.5—
0.75 mm wide) midrib; leaves of nonflowering branches petiolate, S—
8 cm, less often to 35 cm long and 0.5—0.7(1.6) cm wide; lower cauline
leaves petiolate or subsessile, 6-10(15) cm long and 0.4—1.0 cm wide;
middle and upper leaves sessile, 1.5—5.0(10.0) cm long, 0.2—0.8(1.0-
1.5) cm wide; leaf segments numerous, antrorse, usually approximate,
often 1.0—-1.5 mm apart, less often to 3 mm; segments of middle cauline

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78

leaves 2-4(9) mm long and 2-5 mm wide, usually longer at base of
middle and upper leaves, semiamplexicaul (auricles), gradually reduced
and remote in leaves of nonflowering branches and in lower part of
lower cauline leaves; segments twice pinnately cut, each lobe with 2—
5(7) lobules or teeth, 0.1-0.2 mm wide and 0.5—1(1.5) mm long, setose,
linear, linear-lanceolate or less often lanceolate, terminating into more
or less cartilaginous cusp. Capitula in dense, convex, compound,
sometimes unequal corymbs, often subglobose in beginning of
flowering. Receptacle convex to conical. Involucre oblong-cylindrical,
less often oblong or ovoid-oblong, often weakly angular, 3—4(5) mm
long and 1.5-2.5(3) mm in dia; involucral bracts pale, greenish-
yellowish, more or less pubescent, weakly carinate, with prominent
midrib; outermost involucral bracts deltoid or oblong-ovate, others
oblong, with light brownish, less often brown or white, narrow,
membranous border; bracts oblong-lanceolate or lanceolate, shorter
than tubular florets, whitish, membranous, transparent along margin,
glabrous or with appressed white hairs in upper part on outside. Ligules
of outer florets yellowish-white above, whitish below, 0.9-1.5 mm
long, and 1-2 mm wide, subelliptical, less often almost rotund or
elliptical, apically truncate and more or less distinctly obtusely-3-
toothed. Achenes oblong-cuneate, about 2 mm long, truncate at apex.
Flowering June to July (August); fruiting July to August.

In steppes, less often in meadows, among shrubs, along forest
edges, open glades, on grazing grounds, old fields, by roadsides, and
as a weed in crops, from forest-steppe to semidesert zone, and in the
Caucasus up to subalpine zone.—European part: Upper Volga (south),
Volga-Kama (south), Upper Dnieper (south), Middle Dnieper, Volga-
Don, Trans-Volga Region, Upper Dniester, Bessarabia, Black Sea
Region, Crimea, Lower Don, Lower Volga; Caucasus: Ciscaucasia,
Dagestan, Eastern, Western, and southern Transcaucasia, Talysh;
Western Siberia: Upper Tobol, Irtysh, Altai; Soviet Central Asia: Aralo-
Caspian Region, Lake Balkhash Region, Dzhungaria-Tarbagatai, Syr-
Darya, Tien Shan. General Distribution: Central and Atlantic Europe,
Mediterranean Region, Balkans-Asia Minor, Armenia and Kurdistan,
Iran. Described from Hungary. Type in Vienna.

Note 1. It hybridizes with A. leptophylla MB., A. coarctata Poir.,
A. nobilis L. and some other Achillea species, as well as with
Chrysanthemum millefoliatum L. (Chrysanthemoachillea borzae
Prodan). In the northern part of its range and even:in more southern
places, where in contact with A. millefolium L. s. 1., it is connected
through intermediate forms, and in northern Kazakhstan, where in
contact with A. asiatica Serg., intermediate forms between A. setacea
Waldst. and Kit and the latter species, apparently also partly of hybrid
origin, are observed.

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79

Note 2. A.A. Grossheim (Opred. Rast. Kavk. 456) and I.P.
Mandenova (in Fl. Georgii, VIII, 330) reported from the Caucasus A.
nabelekii Heim. [Pubi. Fac. Univ. Masaryk, 52 (1925) 12], described
from Kurdistan. A. nabelekii Heim., from the A. setacea Waldst. and
Kit. group, differs from the latter mainly by larger plant size, less
densely appressed segments of the cauline leaves with the segments up
to 0.5 mm wide, a larger, broadly ovoid involucre (5.5 mm long and
almost 4 mm in dia) and white ligules 1.75—2.0 mm long and 2.5-—3.0
mm wide. A. nabelekii Heim. is not preserved in the herbarium of the
Botanical Institute, Academy of Sciences of the USSR, and it is not
clear whether it was really found in the Caucasus. It is possible
that the most robust specimens of A. setacea Waldst. and Kit. were
described under this name.

8. S. asiatica Serg. in Sistemat. Zam. Gerb. Tomsk. Univ. 1 (1946)
6; Kryl. Fl. Zap. Sib. XI, 2723.—A. setacea auct. fl. Sib., non Waldst.
and Kit.—Exs.: Gerb. Fl. SSSR, No. 3984.

Perennial. Rhizome slender, creeping, branched; stems less
numerous, less often solitary, grayish from long, fine, intertwined white
hairs, (8)25-50(90) cm high, erect or ascending from base, straight or
flexuous, cylindrical, finely sulcate, simple or branched above, often
with short leafy branches in axils of middle and upper cauline leaves.
Leaves twice pinnately cut, oblong, less often oblong-linear, oblong-
or linear-lanceolate, green or grayish-green, more or less densely hairy,
especially along narrow (0.5-0.75 mm wide); midrib with long, fine,
white hairs, leaves of nonflowering branches long-petiolate, to 25 cm
long and 0.5-1.5(5) cm wide; lower cauline leaves petiolate or
subsessile, 7-20 cm long, 0.5—1.0(2.0—3.0) cm wide; middle and upper
leaves sessile, 1-2 to 6(15) cm long and 0.4—1.2 cm wide; leaf segments
numerous, approximate, usually 1.0—-1.5 mm apart, less often to 3 mm;
segments of middle cauline leaves 2-6(9) mm long and 2.5-5.0 mm
wide, usually slightly longer, semiamplexicaul (auricles), gradually
reduced and more distant in leaves of nonflowering branches and lower
part of lower cauline leaves; segments divided in 2—4 toothed lobes,
teeth, or lobules 0.5—1.0(2.0) mm long, and 0.1-0.3 (0.5) mm wide,
linear, lanceolate-linear, less often almost lanceolate, terminating in
cartilaginous cusp. Capitula in lax, convex, usually unequal, compound
corymbs. Receptacle convex to conical. Involucre poculiform, 3-4 mm
long, 2.5-3 mm in dia; involucral bracts pale, yellowish to pale green,
ovate, outer involucral bracts oblong or lanceolate-oblong, with
prominent midrib, weakly carinate, with brown, dark brown, less often
pale brownish membranous border; bracts oblong-lanceolate, slightly
shorter or almost as long as tubular florets, whitish, membranous,

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80

transparent along margin, sparsely hairy in upper part on outside.
Ligules of outer florets pink, less often purple, very rarely white, 1.0—
3.0(3.5) mm long, 1.5—2.5(3.0) mm wide, subelliptical or almost rotund,
truncate at apex, somewhat distinctly obtusely 3-toothed. Achenes
cuneately oblong, 2 mm long, apically truncate. Flowering July to
August; fruiting August to September.

In flooded and dry bottom meadows, shrub thickets, forests, forest
edges and glades, open mountain slopes, and less often in steppes, in
old fields and pastures, by roadsides and in fields, rarely in crops,
from forest to northern part of the steppe zone.—Western Siberia:
Upper Tobol, Irtysh, Ob River Area, Altai; Eastern Siberia: Yenisei,
Angara-Sayans, Dauria, Lena-Kolyma; Far East: Zeya-Bureya, Ussuri,
Uda River Area, Okhotsk; Soviet Central Asia: Lake Balkhash Region
(north and east), Dzhungaria-Tarbagatai. General distribution:
Mongolia. Described from the Chulym River valley. Type in Tomsk.

Note. A. asiatica Serg. is close to A. setacea Waldst. and Kit. and,
like the latter, is related to A. millefolium L. through transitional forms
(var. intermedia Serg. |. c. etc.).

9. A. bucharica Winkl. in Tr. Peterb. Bot. Sada. XI (1890) 163;
QO. and B. Fedtsch. Perech. Rast. Turk. IV, 182; B. Fedtsch. Rast.
Turk., 736.—A. schugnanica Winkl. op. cit. 164; O. and B. Fedtsch.
op. cit. 183; B. Fedtsch. op. cit. 736.

Perennial. Rhizome thick, many-headed, branched, woody; plants
densely villous-lanate with white, divergent hairs; stems few, erect or
ascending from base, (5)10—25(40) cm high, simple, erect or weakly
curved, cylindrical, finely sulcate. Leaves usually grayish-green from
more or less dense pubescence, with distinct midrib, 0.6—1 mm wide;
middle and upper cauline leaves sessile, upward directed, linear-
lanceolate or lanceolate, (1)2—-4(6) cm long, 3.0-0.6 cm wide, twice
pinnately cut, with slightly distant segments, lower larger
semiamplexicaul, divided in almost equal, entire, linear-lanceolate or
lanceolate lobes, 0.3—0.5(0.7) mm wide, terminating into rather long
and thin cartilaginous cusp; lower cauline leaves and those of
nonflowering branches 10-15 cm long, petiolate. Capitula on lanate
peduncles in dense, unequal, strongly convex, more often hemispherical,
compound corymbs, 1.5—3.0(4.0) cm in dia. Receptacle convex.
Involucre ovate-campanulate or oblong, about 4 mm long and 2.5 mm
in dia; involucral bracts more or less densely pubescent; outer bracts
ovate-oblong, others oblong-lanceolate, with dark-brown membranous
border, and with weakly developed yellowish midrib; bracts finely
membranous, white or brownish at apex and along margin, glabrous,
oblong-lanceolate, slightly shorter than light yellow tubular florets.

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81

Ligules of outer florets white (yellowish-white?); rotund-reniform,
(0.8)1.0—1.2(1.5—2.0) mm long, (1.5)2.0-2.5 mm wide, more or less
distinctly 3-toothed at apex. Achenes cuneate-oblong, 1.8—2.0 mm long.
Flowering July to October.

In meadows and on stony slopes in alpine and subalpine zones.—
Soviet Central Asia: Pamiro-Alai. General distribution: Known from
the Lake Shiva in Afghanistan. Described from Sagyrdasht (Darvaz
Range). Type in Leningrad.

Note. It varies significantly in the degree of pubescence as well as
the shape and toothing of the ligules of the outer florets.

10. A. distans Waldst. and Kit. ex Willd. Sp. pl. II, 3 (1804)
2207; Szafer, Kulcz. Pawl. Rosl. Polskie, ed. I, 623; Klok. in Vizn.
Rosl. URSR, 541.—A. dentifera DC. Fl. Fr. V (1815) 485 and Prodr.
VI, 25.—A. dentifera B. distans (Waldst. and Kit.) Heuff. Enum. pl.
Banat. (1858) 98.—A. tanacetifolia var. a. dentifera (DC.) Koch. Syn.
Fl. Germ. I (1837) 374.—A. tanacetifolia var. B. distans (Waldst. and
Kit.) Rochel, Pl. Banat. rar. XXIII (1828) 74, fig. 71; Hayek in Hegi,
Ill. Fl. VI, 2, 575; Szafer, Kulcz. Pawl. op. cit. ed. II, 683 (sybsp.).—
A. tanacetifolia auct. mult. Fl. Ross. non All.—Ic.: Rchb. Ic. Fl. Germ.
XVI, f. 1025; Javorka and Csapody, Iconogr. Fl. Hung. f. 3735 f.—
Exs.: Fl. exs. Austro-Hung. No. 995; Schulz Herb. norm. Nos. 1388,
2000.

Perennial. Rhizome slender, branched; plants more or less
pubescent, with fine white hairs; stems solitary or few, erect, 30-
60(90) cm high, simple, erect or weakly flexuous, cylindrical, finely
sulcate. Leaves green; middle and upper cauline leaves sessile, upward
directed, oblong-lanceolate, 10-12 cm long and 3-4 cm wide, twice
pinnately cut, segments linear- or oblong-lanceolate, divided in
lanceolate or broadly lanceolate acute lobes, usually with cartilaginous
cusp, lower segments semi-amplexicaul; lower cauline leaves and leaves
of nonflowering branches petiolate, twice or thrice pinnately cut, with
longer segments, lanceolate; leaves on nonflowering branches 25(45)
cm long and 5—6(13) cm wide; midrib 1.0-2.5(4.0) mm wide, usually
toothed throughout with intermediate (between main segments), crenate-
toothed lobes or teeth; young leaves sericeous, others weakly pubescent
with scattered hairs. Capitula on hairy, slender peduncles in many-
flowered, unequal, dense, convex, compound corymbs; inflorescence
up to 12 cm in dia. Receptacle conical. Involucre oblong, (4)5—6 mm
long (2)2.5—3 mm in dia; involucral bracts more or less pubescent,
greenish, with narrow, membranous, dark-brown border, weakly
carinate; outer bracts ovate or deltoid-ovate, others oblong-lanceolate,
with prominent yellowish midrib on outside; bracts lanceolate,

8

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82

membranous, white, transparent, slightly shorter than tubular florets,
with long white hairs on outside. Ligules of outer florets white, less
often pink, (1.8)2.5-—3.0(3.5) mm long, subelliptical, apically truncate,
indistinctly obtusely 3-toothed. Flowering July to August.

On open herb and stony mountain slopes, in subalpine meadows,
on polonines*, in forest glades, less often in beech forests.—European
part: Upper Dniester. General distribution: Central Europe, Balkans.
Described from Western Europe. Type in Vienna.

Note 1. Highly variable in the shape, size, and division of leaves,
width and degree of toothing of the leaf midrib, color intensity of the
margin of the involucral bracts, and other characters. It hybridizes
with A. millefolium L. and some other species of the genus Achillea.

Note 2. M.V. Klokov (op. cit.) reported A. tanacetifolia All. from
Western Ukraine (Carpathians), which differs from A. distans Waldst.
and Kit. mainly in the width of the leaf midrib, broadly ovate lobes of
the leaf segments and red (very rarely white) outer florets. There are
no specimens of A. tanacetifolia All. s. s. from this region in the
herbarium of the Botanical Institute, Academy of Sciences of the USSR.

Long ago C. Koch (1. c.) reported that the species of the A.
tanacetifolia s. 1. group differ from each other only in the width and
toothing of the leaf midrib, as well as in the form of toothing of the
leaf segments and their lobes. These characters are highly variable in
A. tanacetifolia All. s. s., A. distans Waldst. and Kit., and A. stricta
Schleich., and in many cases their identification is very tentative.
Perhaps, it is more appropriate to consider them subspecies of A.
tanacetifolia All. s. 1., as was recently done by different authors, for
example, Hayek (op. cit.) and others. In the absence of sufficient
herbarium material, including authentic specimens, and given the lack
of information on the ecology and ranges, and also not having the
opportunity to observe them in nature, we cannot resolve this complex
problem and describe them here as species.

11. A. stricta Schleich. Cat. omn. pl. in Helv. nasc. (1821) 5,
nom. nud.; Heimerl in Kern. Schedae ad FI. exs. Austro-Hung. III
(1884) 118; Szafer, Kulcz. Pawl. Rosl. Polskie, ed. I, 623; Popov,
Ocherk Rast. i Fl. Karpat. 247; Klok. in vizn. Rosl. URSR, 523.
—A. tanacetifolia var. angustifolia Weihe in Flora, VI (1823) 749.—
A. millefolium 8. lanuginosa Gaud. Fl. Helv. V. (1829) 374.—A.
tanacetifolia var. Y. stricta Koch, Syn. ed. I (1837) 374; Hayek in
Hegi, Ill. Fl. VI. 2,575; Dostal, Kvetena, 1596 (subsp.); Szafer, Kulcz.

*High-mountain pastures in East Carpathian Mountains—General Editor.

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83

Pawl. op. cit. ed. II, 683 (subsp.).—A. distans subsp. stricta (Schleich.)
Janchen sec. Soo-Javorka, A. Magyar nov. kez. II (1951) 680.—A.
distans var. lanuginosa (Gaud.) Mausf. Verz. (1940) 249.—Ic.: Javorka
and Csapody, Iconogr. Fl. Hung. f. 3734 p.—Exs.: Fl. exs. Austro-Hung.
No. 994; Schultz, Herb. norm. No. 1999; Fl. Rom. exs. No. 1381.

Perennial. Rhizome slender, branched; plants more or less pubescent
from fine white hairs; stem solitary, ascending or erect, 30—60(100)
cm high simple, erect or flexuous, cylindrical, finely sulcate. Leaves
bright green, upward directed, twice pinnately cut; segments distant,
oblong, decurrent, pinnatifid in small cuspidate, narrow-lanceolate or
lanceolate lobes and teeth, 0.3-0.5 to 1.0 mm wide, terminating into
short cartilaginous cusp; middle and upper cauline leaves sessile, oblong
or oblong-lanceolate, up to 12 cm long and 2-3 cm wide, with lower
segments semiamplexicaul; lower cauline leaves and leaves on
nonflowering branches petiolate, lanceolate, larger, latter up to 40 cm
long and 10 cm wide; midrib 1.0—1.5(2.0) mm wide, with intermediate
(between main, distant segments) cuspidate, lanceolate, entire or less
often pinnatifid lobes in middle and upper cauline leaves; young leaves
densely sericeous with white hairs, others weakly pubescent with split
hairs. Capitula on more slender, long peduncles, in dense, convex,
unequal, compound corymbs. Receptacle conical. Involucre oblong,
(4)5—6 mm long, (2.5)3.0-3.5(4.5) mm in dia; involucral bracts weakly
pubescent, pale, slightly carinate, with light or dark brown membranous
border; outer bracts ovate or ovate-oblong, others lanceolate-oblong,
light green, with prominent yellowish midrib; bracts lanceolate, slightly
shorter than tubular florets, white-membranous, usually with occasional
hairs outside. Ligules of outer florets white or pink, (1.8)2—2.4 mm
long, subelliptical or almost rotund, apically truncate, obtusely 3-
toothed. Achenes 2 mm long, oblong, roundish at apex. Flowering July
to September.

On open, clayey, herb and rocky mountain slopes, on polonines*,
in forest glades and among shrubs.—European part: Upper Dniester.
General distribution: Central Europe, Balkans. Described from
Switzerland. Type in Vienna.

Note. It is extremely variable in the shape, size and division of the
leaves, density of the pubescence, intensity of the color of the
membranous border of the involucral bracts and other characters; it
comprises a series of forms (cf. Hegi 1. c.). Hybridizes with A.
millefolium L. and some other species.

-*See footnote on p. 82—General Editor.

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84

12. A. latiloba Ldb. ex Nordm. in Bull. Acad. Sc. St. Pétersb. II
(1837) 312 and FI. Ross. II, 2, 534; Boiss. Fl. or. III, 259; Grossh. FI.
Kavk. IV, 123.—A. szowitzii Fisch. and Mey. in DC. Prodr. VI (1837)
26.—A. dschimilensis C. Koch in Linnaea, XXIV (1851) 324.

Perennial. Rhizome slender, creeping; plants more or less densely
covered with appressed, long, soft hairs; stems solitary or few (?) 30—
50 cm high, erect or weakly ascending, simple, less often sparingly
branched above, straight or somewhat flexuous, sulcate, cylindrical.
Leaves pinnately cut; middle leaves 4-10 cm long, upper cauline leaves
lanceolate, sessile, with wide, ovate, cuspidate-incised, decurrent, up
to 1 cm-long segments; upper cauline leaves smaller, pectinate-pinnately
parted or pinnatifid, with narrower, entire or cuspidate segments; leaves
on nonflowering branches 22-30 cm long; lower cauline leaves
petiolate, narrow-lanceolate, with numerous segments, up to 1.5 cm
long, lower segments smaller, remote, cuneate or lanceolate, toothed
or entire; middle segments somewhat distant, ovate, decurrent on wide
midrib, pinnatifid into entire or usually 2—4-toothed incised lobes with
acute teeth, upper segments slightly smaller, connate. Capitula
aggregated in dense, convex, compound, many-flowered corymbs.
Receptacle convex. Involucre oblong, 4.5-5.5 mm long, about 3.5 mm
india; involucral bracts reddish-brown on margin; outer bracts deltoid-
ovate, others ovate-oblong, obtuse, weakly carinate, with prominent
midrib; bracts oblong-lanceolate, acute, membranous, transparent on
margin, slightly shorter than tubular florets, with scattered hairs outside.
Ligules of outer florets white, rotund or rotund-reniform, obtusely 3-
toothed at apex, 3.0(3.5) mm wide, 2.5—3.0 mm long. Achenes cuneate-
oblong, about 2 mm long. Flowering July to August.

In subalpine meadows, along forest edges.—Caucasus: Western
Transcaucasia. General distribution: Asia Minor, Armenia. Described
from Georgia. Type in Leningrad.

Note. The prior name is A. latiloba Ldb. ex Nordm., which was
published on July 31, 1837. The sixth volume of De Candolle’s
Prodromus, where A. szowitzii Fisch. and Mey. was published, was
finished with printing in December of the same year.

Section 2. Filipendulinae DC. Prodr. VI (1837) 37; Ldb. Fl. Ross.
II, 2, 535; Boiss. Fl. or. III, 254.—Ligules yellow or golden; leaves
pinnately parted or pinnately cut, leaf segments not membranous.

13. A. filipendulina Lam. Encycl. meth. I (1783) 27; DC. Prodr.
VI, 28; Ldb. Fl. Ross. II, 2, 535; Boiss. Fl. or. III, 259; O. and B.
Fedtsch. Perech. Rast. Turk. IV, 181; Grossh. Fl. Kavk. IV, 124.—A.
eupatorium MB. Tabl. prov. Casp. (1798) 119 excl. synon. Tourn.; FI.

5

_

85

taur.-cauc. II, 337; III, 585; Willd. Sp. pl. Il, 3, 2205.—A. filicifolia
MB. FI. taur.-cauc. II (1808) 338 and III, 585.—Tanacetum angulatum
Willd. Tract. de Achilleis (1789) 52, 2, f. 3.—Ic.: Gartenflora, XXVIII,
tab. 264.—Exs.: GRF Nos. 1874, 3985.

Perennial. Rhizome woody; whole plant more or less densely
pubescent from slightly appressed hairs; stems less numerous, less
often solitary, (25)40—70(120) cm high, erect, ribbed-sulcate, straight
or weakly flexuous, simple, less often sparingly branched, densely leafy,
very rarely with short branches in axils of cauline leaves. Leaves
pubescent, with frequent punctate-alveolate glandular hairs on both
sides, wide, oblong-lanceolate, pinnately parted, with decurrent oblong-
lanceolate, or oblong, crenulate and obtusely toothed large segments;
midrib of leaves serrate-dentate; leaves on nonflowering branches long-
petiolate, up to 40 cm long; lower cauline leaves petiolate, middle 8-
18 cm long, with (0.5)1.5—2.0(3.0) cm-long middle segments, lower |
segments smaller, more remote; upper leaves sessile, smailer, uppermost
about | cm long, filiform-linear, serrate-dentate or entire. Capitula
with few or many flowers, in dense, large (to 10 cm in dia), compound,
convex, unequal corymbs. Receptacle convex to oblong-cylindrical.
Involucre oblong-obconical, often with elongate base, or oblong-
cylindrical, 3—4 to 9-10 mm long and about 2 mm in dia; involucral
bracts deltoid-oblong, pubescent, pale; bracts oblong-lanceolate, much
shorter than florets. All flowers tubular, or outer flowers short-ligulate,
often irregular, with small (up to 1 mm long) three-lobed reniform-
rotund, bright yellow ligules. Achenes oblong, 1.5-1.75 mm long.
Flowering June to July (September).

On gravel beds in river valleys, on stony, clayey, clayey-sandy
soils along irrigation channels, springs and streams, as well as in old
fields and open dry mountain slopes, in glades and along edges of
mountain forests and shrub thickets. From foothill plains to upper part
of the tree belts (Caucasus) and tree-shrub (Soviet Central Asia) veg-
etation.—European part: Crimea (?); Caucasus: Ciscaucasia, Eastern
and Southern Transcaucasia; Soviet Central Asia: Tien Shan, Syr-Darya,
Pamiro-Alai. General distribution: Armenia-Kurdistan, Iran. Described
from collections of Tournefort from “Levant.” Type in Paris.

Note. It hybridizes with A. millefolium L. It varies greatly in the
shape of the receptacle, involucre, number of florets in the capitula,
density of pubescence, shape of the leaf segments and some other
characters. It forms several varieties. These varieties are found
sporadically in different parts of the general range of A. filipendulina
Lam., do not have separate ranges, and are connected by transitional
forms.

86

Var. leptoclina DC. |. c.—Receptacle elongated, oblong-cylindri-
cal; capitulum many-flowered (30-40 flowers); involucre oblong-cy-
lindrical, up to 9-10 mm long; all flowers tubular, outer sometimes
slightly deformed.

Var. eupatoria (MB.) DC. |. c. [A. eupatoria MB (I. c.)].—
Receptacle not elongated; capitula few-flowered (8-10 flowers); all
flowers tubular.

Var. filicifolia (MB.) DC. 1. c. [A. filicifolia M.B. (1. c.)].—
Receptacle not elongated; capitula few-flowered (8-10 flowers); outer
flowers with short, almost trilobate ligule.

Var. szowitziana DC. |. c.—Receptacle not elongated; capitula
few-flowered (8-10 flowers); outer flowers short-ligulate; leaves
sericeous, with linear-lanceolate acute segments.

14. A. coarctata Poir. in Lam. Encycl. meth. Suppl. (1810) 94;
Prodan. Fl. Rom. 1003.—A. compacta Willd. Sp. pl. Ill, 3 (1804)
2206, non Lam. Encycl. meth. I (1783) 27; DC. Prodr. VI, 28; Ldb. FI.
Ross. II, 2, 536; Boiss. Fl. or. IV, 261; Schmalh. Fl. II, 65; Grossh.
Fl. Kavk. IV, 125; Klok. in Vizn. Rosl. URSR, 544.—A. glomerata
MB. FI. taur.-cauc. III (1819) 585.—A. sericea Janka in Linnaea, XXX
(1859) 579.—A. tomentosa Stankov, Opred. Rast. (1949) 626, non
L.—Ic.: Rchb. Sc. Fl. Germ. XVI, f. 1024; Javorka and Csapody,
Iconogr. Fl. Hung. f. 3730s.—Exs.: Rchb. Fl. Germ. exs. No. 2343; Fl.
Graeca, exs. No. 751; Balansa, Pl. D’orient. No. 244; Dorfler, Herb.
norm. No. 3043; Fl. exs. Austro-Hung. No. 1825; Fl. Hung. exs. No.
80.

Perennial. Rhizome vertical, long, thickened in upper part, many-
headed; plants densely sericeous (ferruginous-sericeous on long
storage); stems few, less often solitary, (12)25-—40(75) cm high,
cylindrical, finely sulcate, slightly ascending from base or erect, straight
or slightly flexuous, simple, sometimes weakly branched, often with
undeveloped, short, leafy branches in axils of middle cauline leaves.
Leaves linear-lanceolate; middle cauline leaves (2)4—6(9) cm long,
upward directed, usually slightly sinuate or arcuate, sessile, pinnately
cut, with numerous, approximate, oblong segments, 4—6 mm long, larger
at leaf base, semiamplexicaul; segments pinnately parted, with linear,
entire, less often 2—3-toothed lobes, obtuse or sometimes short-
cuspidate; segments of upper-most leaves usually entire, usually only
near leaf base; midrib narrow, usually with teeth between main segments
in upper part; leaves on nonflowering branches petiolate, (6)15—20 cm
long, with distant, unequal segments, 2—5 to 10-35 mm long. Capitula
on short peduncles, thickened from dense pubescence, aggregated in
many-flowered, very dense, strongly convex, often almost semiglobose
corymbs. Receptacle at beginning of anthesis almost flat or weakly

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87

convex, at fruiting usually oblong-conical. Involucre turbinate, 3-4
mm long, 2.5-3.5 mm in dia; involucral bracts membranous, outermost
oblong-narrow deltoid, others oblong, obtuse, convex dorsally, with
narrow membranous border in upper part, sometimes brownish at apex;
bracts slightly shorter than tubular flowers, finely membranous below,
white-membranous in upper half, oblong, obtuse, sinuate-toothed at
apex, glabrous or with occasional hairs on outside. Ligules of outer
florets small, a fifth to a third as long as involucre, reniform, obtusely
3-toothed, often undeveloped or absent. Achenes cuneate-oblong, about
1 mm long. Flowering May to July (August).

In dry stony places.—European part: Bessarabia. According to
Klokov (op. cit.), distributed in the Black Sea Region, which has not
been confirmed. General distribution: Balkans-Asia Minor, Armenia-
Kurdistan (north), Central Europe (Romania, Bulgaria). Type in Paris.

Note. A. coarctata Poir. hybridizes with A. setacea Waldst. and
Kit. (A. romanica Prod. op. cit. 1003), A. neilreichii Kern. (A.
dobrogensis Prod. op. cit. 1004), A. leptophylla MB. (A. nyaradyana
Prod. op. cit. 1003), as well as with several other species of the genus.

15. A. ochroleuca Ehrh. in Beitr. z. Naturk. VII (1792) 166, non
Willd. Sp. pl. III, 3 (1804) 2210; Waldst. and Kit. Pl. rar. Hung. 1, 33;
Afanasiev in Bot, Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX, 365.—
A. pectinata Willd. op. cit. 2197, non Lam. Encycl. meth. I (1783) 28;
DC. Prodr. VI, 28; Ldb. Fl. Ross. II, 2, 535; Boiss. Fl. or. Ill, 262;
Schmalh. Fl. II, 65; Prodan, Fl. Rom. I, 2, 1001; Dostal, Kvetena,
CSR, 1592; Klok. in Vizn. Rosl. URSR, 544.—Ic.: Waldst. and Kit.
op. cit. tab. 34; Rchb. Ic. Fl. Germ. XVI, f. 1041; Javorka and Csapody,
Iconogr. Fl. Hung. f. 3729f.—Exs.: Fl. exs. Austro-Hung. No. 999; Fl.
Hung. exs. No. 579.

Perennial. Rhizome woody, many-headed; stems grayish from
appressed tomentose pubescence, few, ascending from base, less often
straight, (10)25-—35(60) cm high, finely sulcate, cylindrical below,
weakly angular above, simple, sometimes sparingly branched above,
often with short leafy branches in axils of middle cauline leaves. Leaves
with appressed pubescence or sub-glabrous, usually punctate-alveolate
on both sides, upward directed, linear, pinnately parted, 2-3(4) cm
long, with broad (up to 1.5—2.0 mm wide) midrib, segments pectinate,
arranged somewhat distant, antrorse, decurrent, 1—-3(5) mm long, linear
or subulate-linear, acute, with very short white-cartilaginous cusp, entire
or some 1—3-toothed; middle cauline leaves sessile, basally with longer,
often linear-filiform segments, slightly amplexicaul; lower cauline leaves
as well as leaves on nonflowering branches and that on short branches,
and often also uppermost cauline leaves long-petiolate, with segments

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88

usually only in upper half and near base (auricles). Capitula on densely
tomentose peduncles, in compound, convex, usually dense and inequal
corymbs. Receptacle flat at flowering, convex at fruiting. Involucre
ovate or oblong-ovate, 2.5—3.0(3.5) mm long, 2.0—2.5(3.0) mm in dia;
involucral bracts membranous, weakly carinate, with thick midrib;
outermost bracts deltoid-ovate, other oblong, obtuse, white-membranous
on margin, sometimes light brown at apex; bracts oblong-lanceolate,
much shorter than tubular florets, almost membranous, with wide white
border, glabrous or with occasional hairs outside. Ligules pale yellow
(sometimes white?), almost a half as long as involucre, oblong-rotund,
with 3 indistinct obtuse apical teeth. Achenes cuneately oblong, about
1.25 mm long. Flowering June to August.

On sands, less often on sandy loam soils and stony outcrops.—
European Part: Black Sea Region, Bessarabia. General distribution:
Balkans; Central Europe, Romania, Hungary, Czechoslovakia.
Described from Hungary. Type in Vienna?

Note. It varies considerably in the pubescence and dissection of
the leaves.

The Caucasian botanists A.A. Grossheim, D.I. Sosnowsky, I.P.
Mandenova and others, following E. Boissier, mistakenly accepted as
A. ochroleuca Ehrh. the yellow-flowered race of yarrow close to A.
nobilis L., the correct name of which is A. neilreichii Kern. (cf.
Afanasiev, l.c.).

1. A. leptophylla MB. FI. taur.-cauc. II (1808) 335; DC. Prodr.
VI; 29; Ldb: FY Ross:-Il,-2,:536; Boiss: Fl.sor. TII,:262; Grossha Fr
Kavk. IV, 125; Klok. in Vizn. Rosl. URSR, 544; Majevsk. FIl., Ed. 8,
579.—A. leptophylla f. granilica and f. desertorum Paczoski in Vestn.
Russk. Fl. III, 4. (1917) 155.—A. gerberi B. leptophylla (MB.) Schmalh.
Fl. II (1897) 66.—A. taurica MB. FI. taur.-cauc. II (1808) 336 and III
(1819) 584; DC. op. cit., 29.—A. taurica f. villosior Boiss. op. cit.
262.—?A. leptophylloides Prod. in Contrib. bot. Cluj, I, fasc. 17 (1928)
8.—Ic.: Vizn. Rosl. URSR, Fig. 292; Stankov and Taliev, Opred. Fig.
333.—Exs.: GRF No. 820; Herb. Fl. Ucr. No. 98.

Perennial. Rhizome slender, short, vertical, strongly branched,
grayish-green, tomentose-villous, very rarely plants subglabrous; stems
numerous, less often solitary, (6)12—25(35) cm high, cylindrical, usually .
divaricately branched from base, less often simple, sometimes with
short leafy branches in axils of cauline leaves. Leaves linear-lanceolate;
lower leaves oblong-linear, (1.5)2.0-4.0(7.0) cm long, pinnately cut,
usually with thin, narrow, toothless midrib; segments antrorse, in upper
leaves usually entire, toothlike or short-linear, in other cauline leaves
3—4(5) mm long, incised or less often divided in 3-5 oblong, oblong-

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89

spatulate or short-linear lobes, (0.5)1.0-2.0 mm long, obtuse or
subactue; lower leaves petiolate, with somewhat distant segments and
with 3-4 pairs of larger segments near base, more or less amplexicaul.
Capitula on longer, slender peduncles, in somewhat lax, convex, few-
flowered (usually about 10 capitula) compound corymbs. Receptacle
flat or slightly convex. Involucre short-ovate or almost semiglobose,
densely appressed-hairy, 3-4 mm long, (2.0)2.5-3.5 mm in dia;
involucral bracts greenish, with prominent midrib; outer bracts acute-
ovate, others oblong, white-membranous along margin; bracts slightly
shorter than tubular florets, oblong or almost ovate, membranous with
wide white border, more or less glandular on outside, glabrous or
sparsely appressed hairy in upper part. Ligules of outer florets bright
yellow (at beginning of flowering, according to published reports,
almost orange), 1.5-2.0 mm long, rotund-reniform or subelliptical,
apically truncate, more or less distinctly obtusely 3-toothed, a third to
half as long as involucre. Achenes cuneate-oblong, slightly roundish at
apex, 1.0—1.25(1.5) mm long. Flowering May to July; fruiting June to
August.

Usually grows as isolated individuals on clay and clayey loam,
less often on stony-rubbly soils and stony outcrops in semideserts and
steppe zones.—European part: Black Sea Region, Crimea, Lower Don,
Lower Volga, Trans-Volga Region; Caucasus: Ciscaucasia. Reported
from Balkans (Romania, Bulgaria). Described from the south of the
European Part of the USSR. Type in Leningrad.

Note. The plants on stony outcrops (granite, slate, limestone) of-
ten differ significantly in the dissection of the leaves with more exten-
sive, lax and weak pubescence. Paczosky (op. cit.) separated such
specimens on granite as f. granitica Pacz.

Forms transitional to A. micrantha Willd., partly of hybrid origin,
are found in the western part of the range of A. leptophylla in
Ciscaucasia.

It forms hybrids with A. setacea Waldst. and Kit., A. nobilis L., A.
glaberrima Klok., A. coarctata Poir. and A. millefolium L.

Apparently, A. leptophylla MB. s. str. is distributed in the northern
and western parts of Crimea, and a closely related race grows on the
Kerch Peninsula and the southern coast.

17. A. glaberrima Klok. in Ind. sem. Hort. Bot. Charjkov. (1925)
6; in Ukr. Bot. Zhurn. 3 (1926) 20; Vizn. Rosl. URSR, 544.

Perennial. Rhizome reduced, many-headed; plants dark green,
entirely glabrous; stems few, less often to 15 or solitary, 16—35(50?)
cm high, cylindrical, finely sulcate, erect or ascending from base, with
short leafy branches in axils of cauline leaves, branched above. Leaves

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90

somewhat thick, punctate-alveolate on both sides, oblong-linear or lin-
ear, 2—4(7) cm long, petiolate, pinnately parted, uppermost leaves entire
or with isolated segments, antrorse or erect; segments entire or 1—2-
lobed, 1—2(3) mm long, entire, oblong-ovate, oblong or short-linear,
subobtuse, with very short, sometimes cartilaginous cusp, segments
longer at base. Capitula on slender, up to 5 mm-long peduncles, in
few-flowered, lax, compound corymbs. Receptacle convex. Involucre
globose-ovoid, about 4 mm long, 2.5—3.5 mm in dia; involucral bracts
membranous, light green, with wide, yellowish membranous border,
outer bracts ovate, others oblong; bracts oblong-lanceolate, membra-
nous, white, transparent, much shorter than tubular florets; ligules of
outer florets yellow, 1.25—1.5 mm long, almost rotund, truncate at
apex, with 3 unequal obtuse teeth. Achenes oblong-cuneate, about 1.25
mm long. Flowering June to July.

On granite outcrops.—European part: Black Sea Region. Endemic.
Described from southern Ukraine. Type in Kiev.

Note. Found only in the Stalino (Donetsk) Region in the village of
“Kamenny mogily” (Bestash-Gora) over an area of about 4 km’.
According to M.V. Klokov, it hybridizes with A. leptophylla MB.

18. A. micrantha Willd. Tract. de Achilleis (1789) 33, non A.
micrantha Willd. Sp. pl. III, 3 (1804) 2209; Afan. in Bot. Mat. Gerb.
Bot. Inst. Akad. Nauk SSSR, XIX, 361.—A. gerberi Willd. Sp. pl. III,
3, (1804) 2196; MB. FI. taur.-cauc. II, 334; DC. Prodr. VI, 29; Ldb.
Fl. Ross. II, 2, 536; Boiss. Fl. or. II, 263; Schmalh. Fl. II, 65; Grossh.
Fl. Kavk. IV, 125; O. and B. Fedtsch. Perech. Rast. Turk. IV, 182;
Szafer, Kulcz. Pawl. Rosl. Polskie, ed. II, 684; Majevsk. Fl. Ed. 8,
579; Kryl. Fl. Zap. Sib. XI, 2727.—A. gerberi B. subcristata DC. |.
c.—A. gerberi a. major and B. minor C. Koch, in Linnaea, XXIV
(1851) 326.—A. gerberi genuina Schmalh. |. c.—A. gerberi genuina
6. tenuifolia Schmalh.—A. gerberi var. leptophylla Serg. in Kryl. Fl.
Zap. Sib. XI (1949) 2727, non Schmalh.—A. aegyptiac S. Gmelin,
Reise, II (1774) 198, non L.—A. cancrini Grun. in Bull. Soc. Nat.
Mosc. XVI, 4 (1869) 417; Trautv. in Tr. Peterb. Bot. Sada, VIII, 2,
450.—A. santolina Falk, Beitr. II (1786) 243, non L. sec. Ldb. 1. c.—
2A. birjuczensis Klok. in Vizn. Rosl. URSR (1950) 541.—Ic.: S. Gmelin
op. cit. tab. 8, f. 2.—Exs.: GRF No. 673.

Perennial. Rhizome slender, with short underground branches;
plants grayish, more or less tomentose; stems 2-3, less often solitary,
20-40(50) cm high, indistinctly angular, usually weakly flexuous,
simple, less often weakly branched above, usually with short, densely
leafy branches in axils of cauline leaves. Cauline leaves petiolate,
linear or oblong, 3—5(10) cm long, often more or less falcate, with

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wide midrib, pinnately cut; segments usually complicate, often slightly
distant in lower and middle cauline leaves, punctate-alveolate on both
sides, ovate or oblong, somewhat decurrent, incised or divided in 3-
5 ovate, obovate, oblong, rarely lanceolate lobes, entire or sometimes
incised toothed, terminating in cartilaginous cusp, less often segments
entire. Capitula in compound, dense, somewhat convex, corymbs.
Receptacle conical, less often weakly convex. Involucre oblong or
oblong-ovoid, 3—-4(5) mm long, 2-3 mm in dia; usually appressed-
hairy; involucral bracts yellowish-green; outer bracts ovate, others
oblong, carinate, with narrow white membranous border; bracts
lanceolate, much shorter than tubular florets, membranous, with narrow
white margin, glabrous or glandular outside and with few appressed
hairs in upper part. Ligules of outer florets yellow, small, 0.8—1.25
mm long, a fourth to a third as long as involucre, rotund-reniform,
with 3 obtuse teeth. Achenes cuneate-oblong, apically truncate, finely
rugose, 1.25—-1.5 mm long. Flowering June to August; fruiting July to
September (October).

Grows in circular patches on sands, sandy soils, less often in steppe
pastures and alkaline meadows, from forest-steppe to desert zones.—
European part: Ladoga-IImen (introduced), Bessarabia (?), Middle
Dnieper, Black Sea Region, Crimea, Volga-Don, Lower Don, Lower
Volga, Trans-Volga Region; Caucasus: Ciscaucasia; Soviet Central
Asia: Aralo-Caspian Region, Kyzyl-Kum (north), Syr-Darya; Western
Siberia: Upper Tobol, Irtysh. Endemic. Reported as an introduced plant
several times in Western Europe (vicinity of Berlin and other places).
Described from the southeast of the European part of the USSR. Type
in Berlin.

Note 1. A quite polymorphic plant. It varies significantly in the
shape and dissection of the leaves, shape and size of the capitula, etc.,
on the basis of which different forms, having no importance as spe-
cies, have been segregated (see above). Forms transitional to A.
leptophylla MB. and A. biebersteinii Afan. are found, especially in the
western part of its range, apparently partly of hybrid origin. It also
hybridizes with A. millefolium L. and rarely with A. setacea Waldst.
and Kit. The form distributed in Crimea needs further study.

Note 2. This species was first described and validly published by
Willdenow in Tract. de Achilleis in 1789 under the name A. micrantha
Willd. Willdenow described the same plant in 1804 in Species
Plantarum as A. gerberi Willd. Therefore, according to the International
Rules of Botanical Nomenclature, unfortunately the long-established
and generally accepted name A. gerberi Willd. should be rejected as
invalid and replaced by the priority name A. micrantha Willd. (cf.
Afanasiev, op. cit.).

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19. A. biebersteinii Afan. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XIX (1959) 361, nom. nov.—A. micrantha Willd. Sp. pl.
III, 3 (1804) 2209; non 1789; MB. FI. taur.-cauc. II, 336 and III, 584;
DC. Prodr. VI, 29; Boiss. Fl. or. III, 264; O. and B. Fedtsch. Perech.
Rast. Turk. IV, 182; Hayek, Prodr. Fl. Balcan. 644; Grossh. Fl. Kavk.
IV, 125 and Opred. Rast. Kavk. 457.—?A. abrotanifolia Willd. op.
cit. 2213, p. p.—A. micrantha B. sulfurea DC. Prodr. VI (1837) 29.—
A. millefolium var. micrantha (MB.) Schmalh. Fl. II (1897) 64, p.
max. p.—A. pubescens auct. mult. fl. Ross. non L.—A. tomentosa auct.
fl. ross. non L.—A. decumbens auct. fl. Ross. non Lam.—A. odorata
Grossh. Fl. Kavk. IV, 123, non L. nec Koch.—Ic.: Boulomoy, FI.
Liban et Syrie, tab. 219, f. 1 and 221, f. 1—Exs.: Fl. cauc. exs. No.
298; Fl. Palestinae exs. No. 96; H.B.P. No. 49.

Perennial. Rhizome slender, branched, woody in upper part; plants
grayish-green, more or less densely covered with long, weakly
appressed, hairs; stems few, less often solitary, straight or weakly
flexuous, (12)20-—35(60) cm high, finely sulcate, cylindrical below,
weakly angular above, often with short leafy branches in axils of cauline
leaves, simple, sometimes branched above. Cauline leaves pinnatisect,
sessile, linear-lanceolate to oblong-linear, mostly upward directed,
straight or more or less falcately bent upward; segments numerous,
somewhat distant, especially in lower half of leaves, leaves longer at
base, semiamplexicaul (auricles), pinnately cut or parted in 3(5) narrow,
somewhat obtuse, linear, oblong-linear, less often lanceolate lobes,
terminating in short, usually cartilaginous cusp; midrib usually narrow,
not toothed; middle cauline leaves (2)3—4(10) cm long, with segments
(2)3-5(8) mm long and lobes (1)2—3(4) mm long; uppermost leaves
mostly with entire, narrow segments; leaves on nonflowering branches
twice pinnately cut, petiolate, (5)8—12(25) cm long, with distant smaller
segments in lower part. Capitula usually on short peduncles, in more
or less dense, convex, compound, usually unequal corymbs. Receptacle
convex or conical, Involucre oblong to ovate, 3-4 mm long, (2.0)2.5-
3.0 mm in dia; involucral bracts thin-membranous, light colored,
yellowish, usually glaucous, weakly carinate, with midrib prominent
dorsally, white-membranous along margin, obtuse; outer bracts ovate
or oblong, others oblong, bracts wide, oblong, less often broadly
lanceolate, short, almost a half as long as tubular florets, thin-
membranous, transparent. Ligules of outer florets golden or bright
yellow, 1/2—1/3(2/7) as long as involucre, rotund-reniform or semi-
circular, 1.0-1.5(1.8) mm long, 1.5—2.0(3.0) mm wide, truncate above
with 3-obtuse, unequal teeth. Achenes 1.0-1.25 mm long, cuneate-
oblong, roundish at apex. Flowering May to August.

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On clayey, stony, sometimes sandy soils, less often on stony out-
crops and gravel beds along riverbanks in foothill plains, foothills, and
along mountain slopes up to 2,500(3,000) m, in desert, semidesert,
steppe and less shrubby, forest and meadow vegetation. Also found in
irrigated and unirrigated fields, and on old fields, wastelands, along
roads, irrigation channels, etc.—European part: Lower Don (south);
Caucasus: Eastern, Western and Southern Transcaucasia, Ciscaucasia,
Dagestan; Soviet Central Asia: mountainous Turkmenia, Kara-Kum,
Amu-Darya, Kyzyl-Kum (south), Pamiro-Alai Region Syr-Darya, Tien
Shan, Lake Balkhash Region, Dzhungaria-Tarbagatai. General
distribution: Eastern Mediterranean Region, Balkans-Asia Minor,
Armenia-Kurdistan, Iran. Described from Asia Minor. Type in Berlin?

Note. It varies greatly in the degree of pubescence, size and divi-
sion of the leaves, shape of the involucre, color of the florets and some
other characters and comprises a series of more or less weakly separated
forms with presently unknown ranges. Differentiation of all these forms
and descriptions of some as separate geographical races is possible
only after observing them in nature and monographic treatment of the
section Filipendulinae.

Evidently, it hybridizes with A. leptophylla MB., A. setacea Waldst.
and Kit., and A. nobilis L.

According to the International Rules of Botanical Nomenclature
(Article 61), the name A. micrantha Willd. Sp. pl. Ill, 3 (1804) 2209,
under which it was first described, cannot be retained for this species,
as it is a latter homonym of A. micrantha Willd. Tract. de Achilleis
(1789)33. Therefore, we have to reject the commonly accepted name
“micrantha” for this species and replace it with another (cf. Afan. op.
cit. and Note to the previous species).

20. A. cuneatiloba Boiss. and Buhse in Nouv. Mém. Soc. Nat.
Mosc. XII (1860) 117; Boiss. Fl. or. III, 257; Trautv. in Tr. Peterb.
Bot. Sada, VIII, 2, 450; Grossh. Fl. Kavk. IV, 125 and Opred. Rast.
Kavk. 456.—Ic.: Boiss. and Buhse, |. c. tap. X, f. II.

Perennial. Rhizome slender, branched; plant light green, sparsely
covered with long, weakly appressed hairs; stems few or solitary, erect,
25-50 cm high, straight or weakly flexuous, simple, finely sulcate,
cylindrical in lower part, slightly angular above. Leaves oblong-linear,
punctate-alveolate on both sides; lower cauline leaves and leaves on
nonflowering branches petiolate, others sessile, twice pinnately cut,
with somewhat distant segments, decurrent over somewhat narrow
midrib and slightly amplexicaul, divided in linear-cuneate divergent
lobes, 2—4 mm long, slightly broadened above, obtuse, with very short,
cartilaginous cusp; uppermost leaves pinnatisect, with entire segments.

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Capitula on longer peduncles in compound, lax, unequal corymbs.
Receptacle short-conical. Involucre ovate, 3.5—4.0 mm long, 2.5—3.0
mm in dia, weakly pubescent; involucral bracts pale, yellowish, slightly
carinate, with prominent midrib, with white membranous border; outer
bracts deltoid or deltoid-ovate, others oblong, obtuse; bracts broadly
lanceolate, acute, membranous, white, transparent, glabrous, much
shorter than tubular florets. Ligules of outer florets light yellow, rotund-
reniform, a third to a half as long as involucre, with three obtuse teeth
at apex. Achenes cuneate-oblong, slightly roundish at apex, about 1
mm long. Flowering May to July.

On herb slopes, along edges of forests in middle mountain zone.—
Caucasus: Southern Transcaucasia. General distribution: Armenia-
Kurdistan. Described from Southern Transcaucasia. Type in Geneva.

Note. Sommier and Levier described A. cheilanthifolia Somm.
and Lev. in Tr. Peterb. Bot. Sada, XVI (1900) 239 from Transcaucasia
on the basis of a single specimen collected by Brotherus, “Achalczih
in Cartalinia, 7 Jun. 1881 fl. fr. juv. (No. 521 Brotherus).” We have
not seen the herbarium specimens of this species but, judging from the
description, it differs from A. cuneatiloba Boiss. and Buhse by having
smaller lobes of the leaf segments, 1.0-1.5 mm long and very short
ligules, slightly exserted from the involucre.

21. A. micranthoides Klok. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVI (1954) 360 and in Vizn. Rosl. URSR, 544.—A.
micrantha auct. fl. Ucr. non Willd. 1804.—A. millefolium var.
micrantha (MB.) Schmalh. FI. II (1897) 64, p. p.

Perennial. Rhizome short; plants grayish-green from sparse villous-
tomentose pubescence; stems few, sometimes solitary, (12)20-—30(42)
cm high, erect, sulcate, very weakly flexuous or straight, simple, with
short, densely leafy branches in axils of cauline leaves, rarely weakly
branched above. Leaves oblong-elliptical or oblong, (1.5)3.0-—5.0(9.0)
cm long, usually upward directed with frequent punctate-alveolate
glandular hairs on both sides, pinnately cut, with antrorse, somewhat
distant, oblong, incised or lobed segments, lobes linear or oblong,
acute, terminating in short cartilaginous cusp; segments of uppermost
cauline leaves usually entire; middle and upper cauline leaves sessile,
with oblong, slightly amplexicaul (auricles) segments; lower leaves
petiolate; midrib narrow, with few toothless or incised small
intermediate lobes in upper part between main segments, midrib of
uppermost leaves without intermediate lobes. Capitula in lax, compound,
often unequal, convex corymbs. Receptacle convex. Involucre ovate,
3—4 mm long, 2.0-—2.5 mm in dia; involucral bracts membranous, weakly
carinate, outer oblong-ovate, others oblong, with narrow membranous

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border; bracts oblong-lanceolate, glabrous or with isolated hairs outside,
along margin, especially at apex transparent. Ligules of outer florets
light yellow, rotund-reniform, 0.8—0.9 mm long, with 3 obtuse teeth.
Achenes oblong, apically truncate, about 1.5 mm long. Flowering June
to July.

On slopes of steppe depressions—heaths.—European part: Black
Sea Region. Endemic. Described from southern Ukraine. Type in
Leningrad.

Section 3. Santolinoideae DC. Prodr. VI (1837) 30; Ldb. FI. Ross.
II, 2, 538; Boiss. Fl. or. III, 254; Hoffm. in Pflanzenfam. IV, 5, 272.—
Leaves narrow, segments small, transverse, imbricate, ternate, less often
entire; ligules yellow, less often white.

22. A. wilhelmsii C. Koch in Linnaea, XXIV (1851) 328; Trautv.
in Tr. Peterb. Bot. Sada, VIII, 2, 451; Afan. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XV, 397.—?A. santolina L. Sp. pl. (1753)
896 p. p.; DC. Prodr. VI, 31 p. p.; Boiss. Fl. or. III, 266 p. p.—A.
santolina auct. fl. Cauc. non L.—A. eriophora auct. fl. Ross. non
DC.—A. terctifolia Ldb. Fl. Ross. I, 2, (1845-1846) 536, non Willd.—
Exs.: Pl. or. exs. No. 297.

Perennial. Rhizome slender; stems numerous, somewhat woody,
cylindrical, grayish from dense and appressed short-tomentum, erect
or ascending, straight or weakly flexuous, (10)15—20(25) cm high,
with numerous, short, densely leafy branches in axils of cauline leaves,
with upward directed branches, sometimes simple. Leaves subglabrous,
rarely somewhat pubescent, linear, (1.0)1.5—2.0(3.0) cm long, usually
upward directed, often falcately bent, pinnately cut, with numerous,
approximate, transverse, small, about 1 mm-long, ternate segments with
rotund-obovate or obovate, very finely toothed lobes, imbricate,
covering midrib in young leaves and leaves of short branches; segments
of lower leaves more distant, narrower, often entire, cuspidate. Capitula
in compound, rather dense corymbs. Receptacle, flat. Involucre obovate,
pointed, 2.5—3.0(3.5) mm long, 2.5—3.0 mm in dia; involucral bracts
appressed-hairy, oblong, obtuse, carinate, outer often brownish at apex,
others with narrow white-membranous border; bracts lanceolate, thin,
membranous, with wide white membranous border, appressed-hairy in
upper half on outside. Ligules of outer florets yellow, three-lobed,
small, a fourth to a third as long as involucre. Achenes oblong-carinate,
about 1 mm long. Flowering May to July.

On dry stony-gravelly and clayey slopes, in screes, along irrigation
channels, and as weed in crops.—Caucasus: Eastern and Southern
Transcaucasia, Talysh. General distribution: Armenia-Kurdistan, Asia
Minor. Described from Transcaucasia. Type in Berlin?

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23. A. kermanica Gand. in Bull. Soc. bot. Fr. 65 (1918) 37.—A.
krascheninnikovii Afan. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XV (1953) 395.—A. santolina auct. non L.: Boiss. Fl. or III (1875)
266 p. p. and auct. fl. Asiae Mediae.—Exs.: GRF No. 3986.

Perennial. Rhizome slender; stems (12)15—25(32) cm high,
numerous, somewhat woody, cylindrical, grayish-green from sparse,
more or less dense pubescence, ascending or erect, usually distinctly
flexuous, with short densely leafy branches in axils of cauline leaves
and with upward directed branches, sometimes simpie. Leaves grayish-
green, more or less densely pubescent, linear, (1.5)2.0—3.0(5.0) cm
long, usually falcately bent and often antrorse, pinnately cut, with
numerous, distant, small, 1-1.5(2) mm long, ternate segments with
lobate or oblong-obovate uncate lobes, often imbricate covering midrib
in young leaves and leaves of short branches; segments of lower leaves
more distant, sharp toothed. Capitula in compound lax corymbs.
Receptacle flat. Involucre obovate, pointed, 3.0—3.5(4.5) mm long and
2.5-3.0 mm in dia; involucral bracts loosely tomentose, oblong, obtuse,
carinate, outer often greenish and brown at apex, others with narrow
white-membranous border; bracts lanceolate, thin-membranous, with
white-membranous border, on outer side usually pubescent from base.
Ligules of outer florets yellow, three-lobed, small, a fourth to a third
as long as involucre. Achenes oblong-cuneate, 1.5—2.0 mm long.
Flowering April to July.

In deserts and semideserts in clayey and loessial, often saline or
gypseous soils, less often on takyrs and stony slopes of valleys and
ravines, often along irrigation channels and as weed in crops.—Soviet
Central Asia: mountainous Turkmenia, Pamiro-Alai, Amu-Darya, Syr-
Darya, Kara-Kum. General distribution: Iran. Described from Kerman.
Isotype in Leningrad; type in Paris.

24. A. vermicularis Trin. in Mém. Acad. Sc. St. Pétersb. VI (1818)
494; DC. Prodr. VI (1837) 32 errore “verniculata”; Ldb. Fl. Ross. II,
2, 539; Boiss. Fl. or. III, 266; Grossh. Fl. Kavk. IV, 123.—A. amoena
C.A.M. Verz. (1831) 76.—Ie.: Trin. op. cit. t. XIII.

Perennial. Rhizome thick, woody; stems numerous, hard,
cylindrical, finely sulcate, grayish from dense short and appressed-
tomentose pubescence, usually bent or flexuous, branched, 20—40(55)
cm high, with numerous, short, leafy branches in axils of distant cauline
leaves. Leaves glaucous, densely pubescent, pinnately cut, linear, 1-2
cm long, falcate or sinuate; upper and middle cauline leaves, as also
leaves on short branches, cylindrical, almost cylindrical, with numerous
approximate transverse, small, about 1 mm long ternate segments with
sharply toothed lobes imbricately covering midrib; lower leaves with

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distant, narrower and more strongly acute segments, not covering midrib
continuously. Capitula in few-flowered, mostly simple corymbs or
solitary at apices of branches. Receptacle convex. Involucre densely
pubescent, semiglobose or almost globose, 4-5 mm long and (4)5-6
mm in dia; involucral bracts with prominent midrib, and wide,
membranous, transparent, white border, sometimes brownish or brown;
outer bracts ovate or oblong-ovate, others oblong, bracts almost as
long as tubular florets, oblong-lanceolate, appressed-hairy on outside,
with wide, transparent, white, membranous, toothed border. Ligules of
outer florets light yellow, rotund-reniform, 2.2—2.5(3.0) mm wide and
1.5-2 mm long, indistinctly obtusely 3-toothed, 2/5—1/2 as long as
involucre. Achenes cuneately oblong, 1.5—2.0 mm long. Flowering June
to August.

On dry gravelly and stony mountain slopes from 1,500 m to the
alpine zone inclusively.—Caucasus: Eastern and Southern
Transcaucasia, Talysh. General distribution: Armenia-Kurdistan, Iran.
Described from northern Iran (Gilan). Type in Moscow?

25. A. tenuifolia Lam. Encycl. meth. I (1783) 26; DC. Prodr. VI,
31; Ldb. Fl. Ross. II, 2, 538; Grossh. Fl. Kavk. IV, 122 and Opred.
Rast. Kavk. 455.—A. tenuifolia var. albicaulis Trautv. in Tr. Peterb.
Bot. Sada, II (1873) 546.—A. albicaulis C.A.M. Verz. (1831) 76;
Boiss. Fl. or. III, 270.—Ic.: Takhtajan and Fedorov, Atlas Ris. K. FI.
Erevana, Fig. 108.—Exs.: Pl. or. exs. No. 321.

Perennial. Rhizome thick, woody many-headed; stems numerous,
hard, angular-sulcate, white or whitish, densely short- and appressed-
tomentose, densely white-tomentose at base; erect, simple or branched
(20)25-50(75) cm high. Leaves distant, more or less glabrous, linear,
(1.5)2.0-3.0(5.0) cm long, pinnately cut; lower and middle cauline
leaves with distant, usually ternate, less often entire, almost cylindrical
segments, 1-3 mm long, with oblong-linear lobes having short
cartilaginous apical cusp; segments in other narrow-linear leaves
approximate, short, tooth-like; uppermost leaves filiform, usually entire.
Capitula in compound, dense, usually many-flowered corymbs.
Receptacle flat or convex. Involucre oblong-ovate, 3—-4(5) mm long
and 2-3 mm in dia; involucral bracts carinate, with prominent midrib;
outer bracts sparsely floccose, ovate-deltoid, others glabrous or
subglabrous, oblong, glaucous, with wide, transparent, white border;
bracts almost as long as tubular florets, glabrous, lanceolate, with white-
membranous border. Ligules of outer florets yellow, three-lobed or
irregularly obtusely 3-toothed, small, a sixth to a fourth as long as
involucre. Achenes oblong-cuneate, about 2 mm long. Flowering May
to August (September).

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98

On dry gravelly, stony or sandy places from lower to middle
mountain zone.—Caucasus: Eastern and Southern Transcaucasia,
Talysh. General distribution: Armenia-Kurdistan, Iran. Described from
Levant. Type in Paris.

26. A. schischkini Sosn. in Zhurn. Russk. Bot. Obshch. VI (1921)
146; Grossh. Fl. Kavk. IV, 123.

Perennial. Rhizome, thick, woody; stems numerous, hard, finely
sulcate, weakly angular above, grayish from appressed, short, dense
tomentum, ascending, more or less flexuous, 10-25 cm high, with short,
leafy branches in axils of cauline leaves. Leaves greenish, short
pubescent, linear, 1.5-2.0 cm long, sinuate, cylindrical, pinnatisect,
with numerous, small 0.5-1.0 mm long, transverse, densely imbricate
segments, 3-partite or lobed in upper and middle cauline leaves, with
ovate-rotund, finely toothed lobes; segments in lower part of lower
leaves narrow, acute or with 2-3 teeth, midrib quite distinct beneath.
Capitula in lax, few-flowered, compound corymbs at apices of stems.
Receptacle almost flat. Involucre angular ovoid-cylindrical, 4-5 mm
long, 3-4 mm in dia, involucral bracts finely floccose, coriaceous-
membranous, with prominent midrib and narrow light brown
membranous broader; outer bracts ovate-elliptical, other oblong; bracts
almost as long as tubular florets, lanceolate, finely appressed-hairy on
outside, with narrow white-membranous border. Ligules of outer florets
grayish-yellow (in dry form), rotund-reniform, unequally obtusely 3-
toothed, 2—2.5 mm wide, 2/5—1/3 as long as involucre.

On stony and gravelly dry slopes and rocks in middle mountain
zone. Possibly in Transcaucasia. General distribution: Armenia-
Kurdistan, eastern Anatolia. Described from Melyazgerd. Type in
Leningrad.

Section 4. Ptarmica (DC.) Koch, Syn. (1837) 370.—Ptarmica DC.
Prodr. VI (1837) 19.—Leaves entire to thrice pinnately cut; capitula
usually in corymbose inflorescence, less often solitary, often
semiglobose, sometimes ovate, less often oblong; involucral bracts and
receptacular scales deciduous. Florets white, less often pink, with
flattened tubes; ligulate florets 6—23, often 8, usually with limb as long
as involucre, sometimes shorter, less often weakly developed, not
exserted from involucre. Achenes flattened, winged; wing somewhat
thick, spongy.

Note. The shape and size of the capitula, number and size of the
ligulate florets, division of the leaf and achene structure are the
characters used in separating the genera Ptarmica DC. and Achillea L.

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If, however, one studies both genera as a whole or just a fairly large
number of species of each genus, then the enumerated characters appear
typical to the one genus as well as to individual members of the other
genus. We will examine a few examples: most Ptarmica DC. species
have large semiglobose capitula, whereas P. ptarmicoides (Maxim.)
Worosch. has minute ovoid capitula and also usually very short ligules,
bringing it closer to Achillea L. However, the other characters—shape
and division of the leaf, winged achenes, link it inseparably to Ptarmica
DC. Many Ptarmica DC. species have entire or weakly divided leaves,
whereas P. schuri Sch. Bip. has ternately divided leaves, somewhat
resembling the leaves of Achillea L., but the size of the semiglobose
capitula, large ligules and winged achenes bring this species closer to
Ptarmica DC. Many Achillea L. species have 5 or less ligulate florets
each, but A. cretica L. and A. armenorum Boiss. and Hausskn. have 7-
8 and up to 13 ligulate florets. Both these species have semiglobose
capitula, whereas the capitula in most Achillea L. species are ovoid.
Besides, A. cretica L. is characterized by very large, white ligules and
winged achenes. All this brings these species close to Ptarmica DC.
However, the leaf structure and other characters of the two species
link them inseparably to the genus Achillea L. A. borealis Bong. has
large ligules and winged achenes but approaches Achillea L. in the
structure of the leaf and the fewer ligulate florets. Along with species
having greatly divided leaves, the genus Achillea L. has a fairly large
number of species with less divided leaves (for example, A. filipendulina
Lam.). In other words, there is not a single character that would
demarcate Ptarmica DC. as a separate genus from Achillea L. Not
incidentally, therefore, Ptarmica DC. is recognized as a separate genus
by very few taxonomists, mostly by those dealing with a small number
of the species of this group. Therefore, I follow Koch and many other
authors, and consider Ptarmica DC. only as a section of the genus
Achillea L.

The distinguishing features of some species of section Ptarmica
(DC.) Koch are not always well defined; the ranges of these species
often overlap to considerably. Therefore, it would be more appropriate
to consider them only as subspecies; however, according to the traditions
established in the “Flora of the USSR,” binomial names have been
retained for them. Here, we have in mind on the one hand A. salicifolia
Bess., A. sedelmeyeriana Sosn., A. cartilaginas Ldb. and A.
septentrionalis (Serg.) Botsch., on the other, A. biserrata MB. and A.
griseo-virens Alb.; yet another, A. alpina L., A. ptarmicoides Maxim.,
A. japonica Heimerl and some others.

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Series 1. Lingulatae Botsch.—Leaves entire, toothed, obtuse;
cauline leaves semiamplexicaul, basal and lowermost cauline leaves
petiolate. Inflorescence corymbose, pubescence consisting of long,
straight ferruginous hairs.

27. A. lingulata Waldst. and Kit. Descr. et Ic. Pl. rar. Hung. I
(1799) 2, tab. 2; Boiss. Fl. or. III, 276; Heimerl in Denkschr. Acad.
Wien. XLVIII, 151 (Monogr. sect. Ptarmica, 39); Popov, Ocherk Rast.
i Fl. Karpat, 247; Klok. in Vizn. Rosl. URSR, 543.—Ptarmica lingulata
DC. Prodr. VI (1837) 24.—Exs.: Fl. Hung. exs. No. 581.

Perennial. Rhizome creeping, with few stems; stems 20-50 cm
high, erect, simple, branched only in inflorescence, leafy, glabrous
below, sparsely pubescent above, in inflorescence densely covered with,
long, ferruginous, soft, straight, squarrose hairs. Leaves green, entire,
toothed, obtuse with ferruginous, long, soft, straight, squarrose hairs
on margin and midrib beneath; basal leaves withering before flowering,
obovate, petiolate, 2-4 cm long and 1.0—1.7 cm wide; petioles winged;
lower cauline leaves oblanceolate, petiolate, 4-5 cm long and about
1.1 cm wide, with winged petioles; middle and upper cauline leaves
oblong, semiamplexicaul. 1-4 cm long and 0.2-1.0 cm wide.
Inflorescence compact, corymbose; capitula semiovoid, about 7 mm
long and 7 mm in dia (excluding ligules). Involucral bracts imbricate,
3-seriate, lanceolate, 3.5-5.0 mm long, 0.75-1.5 mm wide, in middle
part herbaceous, yellowish-green, with prominent midrib and rather
densely pubescent with ferruginous, long, soft, erect, squarrose hairs;
wide membranous border of involucral bracts dark brown, glabrous,
fimbriate-toothed along margin. Receptacular scales like involucral
bracts, reduced from periphery to center. Outer florets 7-9 pistillate,
ligulate, about 6 mm long, with similar capitate glandular hairs along
tube; ligules white, oval, about 4 mm long and 3.2 mm wide, with 3
obtuse teeth at apex; disk florets white, tubular, bisexual, about 3 mm
long, covered with scattered capitate glandular hairs, five-lobed.
Achenes oblanceolate, about 2 mm long and 1 mm wide, glabrous.
Flowering June to August.

On peaks of the Carpathians, among rocks and subalpine
meadows.—European part: Upper Dniester. General distribution:
Central Europe, Balkans-Asia Minor. Described from several places in
Romania. Type in Prague.

Series 2. Biserratae Botsch.—Leaves lanceolate, entire, biserrate.
Inflorescence corymbose. Pubescence of long, straight, ferruginous
hairs.

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101

28. A. biserrata MB. FI. taur.-cauc. II (1808) 334; Boiss. FI. or.
III, 275; Heimerl in Denkschr. Acad. Wien, XLVIII, 184 (Monogr.
sect. Ptarmica, 72); Grossh. Fl. Kavk. IV, 122; Mandenova, FI. Gruzii,
VIII, 322.—Ptermica biserrata DC. Prodr. VI (1837) 23; Ldb. FI.
Ross. II, 2, 529.—Exs.: Fl. Cauc. exs. No. 372.

Perennial. Rhizome creeping, with solitary stems; stems 35-100
cm high, straight, simple, branched only in inflorescence, leafy, sparsely
pubescent below, rather densely above with ferruginous, long, soft,
straight hairs. Leaves green, lanceolate, sessile, entire, biserrate, sparsely
hairy along midrib on upper and lower surfaces like stem and also with
numerous, more or less sessile, capitate glandular hairs on lower surface;
lower leaves 0.6—5.0 cm long and 2.0-0.7 cm wide, withering before
flowering; upper leaves 1.5-12.0 cm long and 0.2-1.8 cm wide.
Inflorescence corymbose; capitula semiglobose, 5-8 mm long, 6-9 mm
in dia (excluding ligules). Involucral bracts imbricate, 3-seriate, broadly
lanceolate, 3.5-6.0 mm long and 1.5-2.2 mm wide, rather densely
hairy like leaves, herbaceous in middle, yellowish-green with prominent
midrib and wide membranous yellow border, dark brown fimbriate-
dentate along margin. Receptacular scales like involucral bracts,
reducing from periphery to centre. Outer florets 8, pistillate, ligulate,
7-8 mm long; ligules white, oval, 6-8 mm long and 4.0-6.5 mm wide,
with 3 obtuse teeth at apex; disk florets about 3.5 mm long, white,
tubular, bisexual, 5-toothed, teeth acute. Achenes broadly oblanceolate,
about 2.3 mm long and 1.3 mm wide. Flowering May to August.

In montane forests.—Caucasus: Ciscaucasia, Eastern and Western
Transcaucasia. General distribution: Armenia-Kurdistan. Described
from eastern part of Greater Caucasus. Type in Leningrad.

29. A. griseo-virens Alb. Mat. dlya Fl. Kolkhidy (1895) 133;
Grossh. Fl. Kavk. IV, 122; Mandenova, Fl. Gruzii, VIII, 322, Fig.
388.—A. biserrata MB. var. griseo-virens Kolak. Fl. Abkhazia, IV
(1949) 229.

Perennial. Rhizome creeping, with solitary stems; stem 35-80 cm
high, leafy, branched in inflorescence and also with short nonflowering
branches in axils; stems, both surface of leaves, involucral bracts and
receptacular scales rather densely covered with ferruginous, long, soft,
straight hairs. Leaves green or grayish-green, lanceolate, sessile,
bidentate, with numerous, subsessile capitate glandular hairs beneath;
lower leaves 1.5—5.0 cm long and 0.3-0.8 cm wide, withering before
anthesis; upper leaves 1.5-7.0 cm long and 0.2-1.4 cm wide.
Inflorescence corymbose; capitula semiglobose, 5-10 mm long, 6—10
mm in dia (excluding ligules). Involucral bracts imbricate, 3-seriate,
broadly lanceolate, 3.5-6.0 mm long, 2-3 mm wide, herbaceous in

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middle, yellowish-green with prominent midrib and broad membra-
nous-yellow border, dark brown, fimbriate-toothed along margin.
Receptacular scales like involucral bracts, reducing from periphery to
center. Outer florets 8, pistillate, ligulate, 7-10 mm long; ligules white,
oval, 5-8 mm long and 4.0-5.5 mm wide, with 3 obtuse teeth at apex;
disk florets about 3.5 mm long, white, tubular, bisexual, 5-toothed,
teeth acute. Achenes broadly oblanceolate, about 2.5 mm long and 1.5
mm wide. Flowering May to September.

In subalpine and alpine meadows.—Caucasus: Ciscaucasia,
Western Transcaucasia. Endemic. Described from western part of
Greater Caucasus. Type in Tbilisi.

Note. A. griseo-virens Alb. is described from two localities. The
plant with the following label should be considered as the lectotype:
“Circassia: in pratis alpinis jugi Khag-Kytzyrkha (Khach-Katsyrka)
1894, No. 221, N. Alboff,” preserved in the herbarium of the Tbilisi
Botanical Institute, Academy of Sciences of the Georgian SSR.

A. griseo-virens Alb. is very close to A. bisserrata MB. Unlike the
latter, A. griseo-virens Alb. has pubescence on both sides of the leaves
and short nonflowering branches in the leaf axils.

Series 3. Ptarmicae Botsch.—Leaves linear or lanceolate, entire,
toothed or bidentate, glabrous or pubescent, hairs colorless, straight,
short. Inflorescence corymbose.

30. A. ptarmica L. Sp. pl. (1753) 898; Boiss. Fl. or. III, 274;
Klok. in Vizn. Rosl. URSR, 543.—Ptarmica integrifolia Gilib. FI.
Lithuan. III (1782) 216.—P. vulgaris Blakw. ex DC. Prodr. VI (1837)
23 p. p.—P. vulgaris Clus. ex Ldb. Fl. Ross. II, 2 (1845) 529.—
Achillea ptarmica L. ssp. euptarmica Heimerl var. genuina Heimerl in
Denkschr. Acad. Wien, XLVIII (1884) 173 (Monogr. sect. Ptarmica,
61).—Ic.: Syreistsch. Ill. Fl. Mosk. Gub. III, 262; Hegi, Ill. Fl. VI, 2,
fig. 290; tab. 264, fig. 1.—Exs.: GRF No. 672.

Perennial. Rhizome creeping, with solitary stem; stem 20—125 cm
high, leafy, simple, branched only in inflorescence or with short
nonflowering branches in leaf axils, glabrous below, sparsely pubescent
in middle with short, curly, colorless hairs, densely above. Leaves
green, linear or linear-lanceolate, sessile; bidentate; lower glabrous,
uppermost on both sides or only beneath hairy along midrib, hairs
long, soft, erect, colorless, scattered; lower leaves 1.0—8.5 cm long,
and 1-6 mm wide, withering before flowering, upper 0.7—9.0 cm long
and 1.0-6.5 mm wide. Inflorescence corymbose, capitula semiglobose,
5.0-6.5 mm long, 7-9 mm in dia (excluding ligules). Involucral bracts
imbricate, three-seriate, broadly lanceolate, 2.5—4.0 mm long and 1.2-

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1.5 mm wide, with broad green central part, bordered with dark brown,
membranous, ciliate-toothed frill, densely covered with colorless straight
or flexuous, short, appressed hairs. Receptacular scales apically dark
brown, like involucral bracts, reducing from periphery to center; outer
florets 6-10; pistillate, ligulate, 3.0-6.5 mm long; ligules white,
pigmented violet at base or less often white throughout, oval, 1.8-5.0
mm long and 1.2-4.5 mm wide, with 3 obtuse teeth at apex; disk
florets 2.2-2.5 mm long, white, tubular bisexual, 5-toothed, teeth acute.
Achenes broadly oblanceolate, about 1.5 mm long and 0.8 mm wide.
Flowering July to August.

In meadows, along riverbanks and among shrubs.—Arctic: Arctic
Europe; European part: Karelia-Lapland, Dvina-Pechora, Baltic Re-
gion, Ladoga-IImen, Upper Volga, Volga-Kama, Upper Dnieper, Volga-
Don, Upper Dniester; Western Siberia: Ob River Area (introduced?).
General distribution: Scandinavia, Central Europe, Atlantic Europe,
Western Mediterranean, Balkans. Described from temperate Europe.
Type in London.

Note. Sometimes specimens of A. ptarmica L. with double ca-
pitula are found where most of the florets are ligulate and only a few
tubular. Such plants are often cultivated.

31. A. acuminata (Ldb.) Sch. Bip. in Flora, XXXVIII (1855)
15.—Ptarmica acuminata Ldb. FI. Ross. II, 2 (1845) 529.—P. vulgaris
Blakw. ex DC. Prodr. VI (1837) 23 p. p.; Turez. in Bull. Soc. Nat.
Mosc. XXV, 431.—Achillea ptarmica L. ssp. euptarmica Heimerl var.
acuminata Heimerl in Denkschr. Acad. Wien. XLVIII (1884) 173
(Monogr. sect. Ptarmica, 61); Kitam. Compos. Japon. II, 322.—A.
ptarmica L. ssp. macrocephala Heimer! var. angustifolia Heimer] in
Denkschr. Acad. Wien, XLVIII (1884) 176.—A. acuminata Freyn in
Oest. Bot. Zeitschr. XLV (1895) 344.—A. ptarmica auct. non L. (1753);
Kom. and Alis. Opred. Rast. Dal’nevost. Kr. II (1932) 1027.—Ptarmica
speciosa auct. non DC. (1837); Popov, Fl. Sib. II (1959) 726.

Perennial. Rhizome short?; stems solitary or few, 25-130 cm high,
leafy, simple, branched only in inflorescence or with short nonflowering
branches in leaf axils, glabrous below, sparsely in middle and in part
densely pubescent above with short, curly, colorless hairs. Leaves green,
lanceolate or linear-lanceolate, sessile, biserrate; lower leaves glabrous,
uppermost scatteredly pubescent on midrib on both sides or only beneath
with short, soft, straight, colorless hairs; lower leaves 0.7-10.0 cm
long and 0.5-9.0 mm wide, withering before flowering; upper leaves
1-11 cm long and 4.5—9.0 mm wide. Inflorescence corymbose; capitula
semiglobose, 4.0-7.5 mm long and 4.5-9.0 mm in dia (excluding
ligules). Involucral bracts imbricate, 3-seriate, 2.3-4.5 mm long and

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1.2-1.7 mm wide, with narrow yellowish-green central part, bordered
with wide, yellow, ciliate-toothed frill along membranous margin,
sometimes membranous frill light brown, not densely pubescent with
colorless, straight or flexuous, short, appressed hairs. Receptacular
scales apically yellowish, like involucral bracts, reducing from periphery
to center. Outer florets 10—23, pistillate, ligulate, 4.0-9.7 mm long;
ligules white, oval, 3-8 mm long and 2.5-5.5 mm wide, with 3 obtuse
teeth at apex; disk florets 2-3 mm long, white, tubular, bisexual, 5-
toothed; teeth acute. Achenes broadly oblanceolate, about 2.5 mm long
and 1 mm wide. Flowering June to September.

In meadows, riverine sands, riverbanks and among shrubs.—Eastern
Siberia: Lena-Kolyma, Dauria; Far East: Zeya-Bureya, Uda River Area,
Ussuri, Sakhalin. General distribution: Mongolia, Japan, China.
Described from Trans-Baikal Region. Type in Leningrad.

Note. A. acuminata (Ldb.) Sch. Bip. is close to the European
species A. ptarmica L., replacing it in Asia. A. acuminata (Ldb.) Sch.
Bip. differs from A. ptarmica L. in the color and number of ray florets,
color of the involucral bracts and receptacular scales, as well as the
leaf shape.

32. A. ptarmicifolia (Willd.) Rupr. ex Heimerl in Denkschr. Acad.
Wien, XLVIII (1884) 127 (Monogr. sect. Ptarmica, 15) in clave.—
Pyrethrum ptarmicaefolium Willd. Sp. pl. III, 3 (1804) 2151.—
Chrysanthemum ptarmicaefolium Mus.-Puschkin ex Willd. |. c.; in
syn.—Anthemis ptarmicaefolia Adam in Weber and Mohr, Beitr.
Naturk. I (1805) 71.—Achillea leucanthema Persoon, Syn. II (1807)
467.—A. grandiflora MB. FI. taur.-cauc. II (1808) 333; Boiss. Fl. or.
III, 275; Grossh. Fl. Kavk. IV, 122; Mandenova, Fl. Gruzii, VIII, 325,
Fig. 390.—A. dracunculoides Desf. Hort. Paris (1829) 180.—Ptarmica
grandiflora DC. Prodr. VI (1837) 23; Ldb. Fl. Ross. I, 2, 531.—
Achillea ptarmica L. ssp. ptarmicaefolia Heimer] var. grandiflora
Heimer] op. cit. 178. (Monogr. 66).—A. ptarmica L. ssp. ptarmicaefolia
Heimerl var. ruprechtiana Heimer! op. cit. 78 (Mongr. 66).

Perennial. Rhizome creeping, usually with few stems; stems 15-60
cm high, leafy, branched inflorescence and with long nonflowering
branches in leaf axils, glabrous below, in middle part sparsely and
above rather densely pubescent with colorless, short, crisped hairs.
Leaves green, linear, scatteredly pubescent beneath with colorless
straight hairs; lower leaves 2.5—7.0 cm long, 1.5-4.0 mm wide, biserrate,
withering before flowering; upper leaves and those of axillary branches
1-10 cm long, 1-5 mm wide, toothed. Inflorescence corymbose; capitula
semiglobose, 4.5-7.0 mm long and 6.5—9.0 mm in dia (excluding
ligules). Involucral bracts imbricate, 3-seriate, broadly lanceolate, 2.3-

105

4.0 mm long and 1.0-1.8 mm wide, with scattered not very long crisped
colorless hairs, fimbriate-toothed along margin, yellowish-green in
middle, with wide, membranous, light brown frill along margin.
Receptacular scales like involucral bracts, reducing from periphery to
center. Outer florets 8-12, pistillate, ligulate florets 5.2-10.0 mm long;
ligules white, oval, 7-9 mm long and 2.5-3.7 mm wide, with three
obtuse teeth at apex, disk florets 2.7-3.0 mm long, white, tubular,
bisexual, 5-toothed; teeth acute. Achenes broadly oblanceolate, about
2.8 mm long and 1.6 mm wide. Flowering July to August.

On screes and rocks in subalpine and alpine zones.—Caucasus:
Ciscaucasia, Eastern and Western Transcaucasia. Endemic. Type in
Berlin; isotype in Leningrad.

Note. A. ptarmicifolia (Willd.) Rupr. ex Heimerl is characterized
by rather long non-flowering axillary branches. Sometimes some
branches in some specimens of A. ptarmicifolia (Willd.) Rupr. ex
Heimerl bear solitary capitula. These specimens would remove the
boundary between A. ptarmicifolia (Willd.) Rupr. ex Heimerl and A.
sachokiana Sosn., if it were not for the pubescence on the leaves,
branches, involucral bracts, and receptacular scales, typical of the
former species.

Sometimes A. ptarmicifolia (Willd.) Rupr. ex Heimer! hybridizes
with A. biserrata MB.

33. A. sachokiana Sosn. in Tr. Azerb. Otd. Zakavk. Fil. Akad.
Nauk SSSR, Sekt. Bot. (1933) 46, fig.; Grossh. Fl. Kavk. IV, 122.

Perennial. Entirely glabrous plants. Rhizomes creeping; stems
ascending, 15-30 cm high, leafy, branched; main stem with corymbose
inflorescence, cauline branches often with solitary capitula. Leaves
green, broadly lanceolate, toothed, sessile, 0.5—1.5 cm long and 1-5
mm wide; lower leaves withering early. Capitula semi-globose, about
7 mm long and 9 mm dia, elongate. Involucral bracts imbricate, 3-
seriate, broadly lanceolate, 3-5 mm long and 1.0—2.8 mm wide, green
in middle, with wide membranous dark-brown border, fimbriate-toothed
in upper part. Receptacular scales, like involucral bracts, reducing from
periphery to center. Outer florets 8, pistillate, ligulate, about 8 mm
long; ligules white, oval, about 6.5 mm long and 5.5 mm wide, with
3 obtuse teeth at apex; disk florets about 4 mm long, white, tubular,
bisexual, 5-toothed, teeth acute. Achenes broadly oblanceolate, about
3.2 mm long and 1.5 mm wide. Flowering July to August.

In alpine zone on stony places.—Caucasus: Eastern Transcaucasia.
Endemic. Described from mountainous pastures in the vicinity of
Shemakhi. Type in Tbilisi; isotype in Leningrad.

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Note. A. sachokiana Sosn. is known to me only from one locality,
the place of origin of the type specimen. It differs from A. ptarmicifolia
(Willd.) Rupr. ex Heimerl by a complete absence of pubescence, shorter
and broader leaves, and in that the axillary branches are not
nonflowering but usually terminate in solitary capitula.

34. A. macrocephala Rupr. in Mater. k Blizh. Pozn.
Prozyabaemosti Ross. Imper. 2 (1845) 41; ejusd. Fl. samojed. cisur.
(1846) 41; Heimerl in Denkschr. Acad. Wien, XLVIII, 127 (Monogr.
sect. Ptarmica, 15) in clave.—A. ptarmica L. var. speciosa Herd. in
Bull. Soc. Nat. Mosc. XXX VIII (1865) 407, non A. speciosa Henckel
(1806).—A. ptarmica L. ssp. macrocephala Heimer! var. angustifolia
Heimerl op. cit. 176 (Monogr. 64) p. p.—A. ptarmica L. ssp.
macrocephala Heimer! var. latifolia Heimer] op. cit. 177 (Monogr.
65); Hultén, Fl. Kamtch. IV, 167; Kitam. Compos. Japon, II, 322.—
A. ptarmica L. var. beringiana Trautv. in Tr. Peterb. Bot. Sada, IX, 2
(1886) 447.—Ptarmica macrocephala Kom. Fl. Kamch. III (1930) 137
and Izbr. Soch. VIII: II (1951) 437.

Perennial. Rhizome short; stems solitary or few, 25-100 cm high,
leafy, erect, branched in inflorescence and, also with short nonflowering
branches in leaf axils; glabrous below, in middle part sparsely and
above densely pubescent with short, curly, colorless hairs. Leaves green,
lanceolate or broadly-lanceolate, sessile, biserrate, lower leaves
glabrous, middle beneath, and upper on both sides pubescent on midrib
with scattered, colorless, erect hairs; lower leaves 2.5—8.0 cm long,
2.5-8.0 mm wide, withering before flowering; upper leaves 0.8—9.0
cm long, 2-10 mm wide. Inflorescence corymbose; capitula
semiglobose, 6-8 mm long and 8-11 mm in dia (excluding ligules).
Involucral bracts imbricate, 3-seriate, 3-5 mm long, 1.7—2.0 mm wide,
with broad green central part, bordered with dark brown membranous
margin, densely or sparsely covered dorsally with colorless, rather
long, curly hairs, margins fimbriate-toothed. Receptacular scales dark
brown at apex, like involucral bracts, reduced from periphery to center,
outer florets 8-14, pistillate, ligulate, 7.2-10.0 mm long; ligules white,
sometimes violet at base, oval, 5.6-9.0 mm long and 4.5-6.0 mm
wide, with 3 obtuse teeth at apex; disk florets 2.2-2.7 mm long, white,
tubular, bisexual, 5-toothed, teeth acute. Achenes broadly oblanceolate,
about 2.4 mm long and 1 mm wide. Flowering July to September.

Along riverbanks, in willow stands, along gravel beds and in
meadows near rivers.—Far East: Kamchatka (Commander and Kuril
islands). General distribution: Japan. Described from the vicinity of
Petropavlovsk-Kamchatsky (Martens). Type in Leningrad.

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Series 4. Salicifoliae Botsch.—Leaves lanceolate, entire, biserrate,
punctate-glandular hairy on both sides, or not; also covered with short,
curly, colorless hairs. Inflorescence corymbose.

35. A. salicifolia Bess. Suppl. Catal. pl. Jard. Botan. Krzemieniec
(1812) 3.—Ptarmica vulgaris DC. var. pycnocephala Trautv. in Bull.
Nat. Mosc. XXXIX, 2 (1866) 345.—P. vulgaris Clus. var. cartilaginea
DC. in B. and O. Fedtsch. Perech. Rast. Turkest. IV (1911) 181.—
Achillea borystenica Klok. in Vizn. Rosl. URSR (1950) 543, nom.
subnud.—A. cartilagines Ldb. ssp. borystenica Sakalo, in Vizn. Rosl.
URSR (1950) 543, in syn.—Ptarmica borystenica Klok. and Sakalo in
Bot. Mat. Gerb. Bot. Sada, XVI (1954) 357.

Perennial. Rhizome short, plants usually grayish-green, densely
covered with short crisped hairs; stems erect, 20-100 cm high, leafy,
branched; branches usually short, nonflowering, only uppermost, and
sometimes also middle branches, bearing capitula. Leaves 0.7-10.5 cm
long, 1-13 mm wide, reduced from base to stem apex, sessile, linear-
lanceolate, entire, biserrate, slightly involute, punctate-glandular hairy
on both sides; lower cauline leaves withering before flowering.
Inflorescence corymbose; capitula usually many-flowered, semiovate,
4.0-5.5 mm long and 3.5—5.0 mm in dia (excluding ligules). Involucre
imbricate, 3-seriate; involucral bracts 2-3 mm long, 1.25—1.5 mm wide,
broadly lanceolate, herbaceous in middle, yellowish-green, with
prominent midrib, membranous on margin, yellow or yellow with dark
brown border. Receptacular scales like involucral bracts, reduced from
periphery to center. Outer florets 8, pistillate, ligulate, 2.0-4.25 mm
long; tube with isolated capitate glandular hairs; ligules white or white
with violet base, oval, 1-3 mm long, 1.3-3.25 mm wide, with three
obtuse teeth at apex; disk florets 5-toothed, white, tubular, bisexual,
1.5—2.6 mm long, with solitary capitate glandular hairs. Achenes oblong
or oblanceolate, 1.0-1.5 mm long and 0.75-1.0 mm wide, glabrous.
Flowering June to August.

Along banks of rivers and lakes, in meadows and marshes, birch
stands.—European part: Dvina-Pechora (introduced), Upper Volga,
Upper Kama, Upper Dnieper, Middle Dnieper, Volga-Don, Trans-Volga
Region, Upper Dniester, Bessarabia, Black Sea Region, Crimea, Lower
Don, Lower Volga; Western Siberia: Ob River Area, Upper Tobol,
Irtysh, Altai; Eastern Siberia: Lena-Kolyma, Angara-Sayans; Soviet
Central Asia: Aralo-Caspian Region, Lake Balkhash Region,
Dzhungaria-Tarbagatai. General distribution: Central Europe. Described
from Dniester. Type in Leningrad.

Note. Ptarmica borystenica Klok. and Sakalo was described in
1954. The authors distinguished their species from P. cartilaginea

114

108

Ldb. by a series of characters. These species should be differentiated,
although the border between them is’not always distinct. In the note to
their species, M.V. Klokov and D.I. Sakalo say that while examining
the authentic material of Achillea salicifolia Bess. preserved in Kiev,
they could not distinguish it from Ptarmica cartilaginea Ldb., and
suggested that, apparently, preference should be given to the name
proposed by Besser as having priority. I cannot agree with this, as it
contradicts the facts. The plant from Dniester, which was labeled as
Achillea salicifolia Bess. in Besser’s own hand, is being preserved in
the Herbarium of the Botanical Institute, Academy of Sciences of the
USSR. This specimen should be designated as the lectotype of the
species. The lectotype of A. salicifolia Bess. differs from the type
specimen of Ptarmica cartilaginea Ldb. particularly in those traits by
which P. borystenica Klok. and Sakalo was characterized. Therefore,
the description of P. borystenica Klok. and Sakalo was superfluous,
and the name must be considered a synonym of Achillea salicifolia
Bess. As regards Ptarmica cartilagines Ldb., this is a separate, closely
related species of the genus Achillea L.

36. A. sedelmeyeriana Sosn. in Schedis herb. “Pl. orient. exs.,”
fasc. IX—XVI (1928) 7 and in Zhurn. Russk. Bot. Obshch. XIV, 1
(1929) 82; Grossh. Fl. Kavk. IV, 121; Mandenova, in Fl. Gruzii, VIII,
325, Fig. 389.—Exs.: Pl. or. exs. No. 221.

Perennial. Plants grayish-green, densely covered with short, curly
hairs; stems erect, 50-65 cm high, leafy, branched; lower branches
short, nonflowering, upper terminating in capitula. Leaves 0.6—6.0 cm
long and 1-6 mm wide, reducing from base to stem apex, sessile,
linear-lanceolate, entire, biserrate, somewhat involute, punctate-
glandular-hairy on both sides; lower leaves withering before flowering.
Inflorescence corymbose, capitula semiglobose, about 4 mm long and
4 mm in dia (excluding ligules). Involucre imbricate, 3-seriate;
involucral bracts 2.2-3.5 mm long, 1.0-1.5 mm wide, broadly
lanceolate, herbaceous in middle, green with prominent midrib,
membranous along margin, dark brown. Receptacular scales like
involucral bracts, reduced form periphery to center, outer florets 8,
pistillate, ligulate, about 4 mm long, with isolated capitate glandular
hairs along tube; ligules white, oval, about 2.5 mm long and 2.2 mm
wide, with 3 obtuse teeth at apex, disk florets white, 5-toothed, tubular,
bisexual, about 2 mm long, with isolated capitate glandular hairs.
Flowering June.

Humid places in middle mountain zone.—Caucasus: Western
Transcaucasia. Endemic. Described from Dzhavakhetia (village of
Gorolovka). Type in Tbilisi; isotype in Leningrad.

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109

Note. A. sedelmeyeriana Sosn. occupies an intermediate position
between A. cartilaginea Ldb. and A. salicifolia Bess. It is closely
related to the former species by the dark brown margin of the involu-
cral bracts, and to the latter species by the dense pubescence and
involute leaves. The range of A. sedelmeyeriana Sosn. is disjunct from
the ranges of both these species.

37. A. cartilaginea Ldb. in Rchb. Fl. Germ. exc. III (1832) 849.—
Ptarmica ircutiana DC. Prodr. VI (1837) 22; Ldb. Fl. Ross. II, 2,
529.—P. vulgaris Blackw. 5. ?cartilaginea DC. Prodr. VI (1837) 23.—
P. speciosa DC. Prodr. VI (1837) 23; Ldb. FI. Ross. II, 2, 530 p. p.—
P. cartilaginea Ldb. Fl. Ross. II, 2 (1845) 530; Sergievskaya, in FI.
Zap. Sib. XI, 2728.—Achillea cartilaginea Ldb. Ind. hort. dorpat. sec.
Ldb. Fl. Ross. II (1845) 530, in syn.; Boiss. Fl. or. III, 274; Heimer]
in Denkschr. Acad. Wien, XLVIII, 127 (Monogr. 15), in clave.—A.
speciosa Henckel £. occidentalis Rupr. in Mater. Blizh. Pozn.
Prozyabaemosti Ross. Imper. 2 (1845) 41; ejusd. Fl. samojed. cisur,
1846 (41).—Ptarmica lenensis Turcz. in Bull. Soc. Nat. Mosc. XXV,
4 (1852) 431. Achillea lenensis Turcz. in Bull. Soc. Nat. Mosc. XXV,
4 (1852) 431.—A. ircutiana Sch. Bip. in Flora, XXX VIII (1855) 15.—
A. cartilaginea Ldb. var. /Jatifolia Rupr. Fl. Ingr. (1860) 587.—A.
ptarmica L. ssp. cartilaginea Heimerl, op. cit. 174 (Monogr. 62).—A.
cartilaginea Boiss. ex Grossh. Fl. Kavk. IV (1934) 122 p. p. quoad
nom.; Klok. in Vizn. Rosl. URSR, 543.—A. speciosa auct. non Spreng.
(1806); Ldb. Fl. Alt. IV (1833) 122.—Ic.: Syreistsch. Ill. Fl. Mosk.
Gub. III, 263.—Exs.: GRF No. 822.

Perennial. Rhizome short; plants green, sparsely pubescent from
curly hairs; stems erect, 20-150 cm high, leafy, branched, lower
branches short, nonflowering, middle and upper long, terminating in
capitula. Leaves punctate-glandular hairy on both sides, sessile, flat,
lanceolate, entire, biserrate-dentate, reduced from stem base to apex,
0.2-12.0 cm long, 0.5-17.0 mm wide, lower leaves withering before
flowering. Inflorescence corymbose; capitula usually numerous,
semiglobose, 3-5 mm long and 3-5 mm in dia (excluding ligules).
Involucre imbricate, 3-seriate; involucral bracts 1.8—4.1 mm long and
1.0—2.2 mm wide, broadly lanceolate, herbaceous in middle part, green,
with prominent midrib, membranous on margin, dark brown.
Receptacular scales, like involucral bracts, reduced from periphery to
center. Outer florets 8, pistillate, ligulate, 4.7-5.7 mm long; tube with
isolated capitate glandular hairs; ligules white or white with violet
base, oval, 3.0-4.2 mm long and wide, with 3 obtuse teeth at apex;
disk florets 5-toothed, white, tubular, bisexual, 2.1-2.7 mm long, with
solitary capitate glandular hairs. Achenes oblong or oblanceolate, 1.8-
2.1 mm long and 1.0-1.1 mm wide. Flowering June to August.

110

Along banks of rivers and lakes, in meadows and marshes, coastal
scrubs and coastal sands.—Arctic: Arctic Europe, Arctic Siberia;
European part: Karelia-Lapland, Dvina-Pechora, Baltic Region,
Ladoga-Ilmen, Upper Volga, Volga-Kama, Upper Dnieper, Middle
Dnieper, Volga-Don, Trans-Volga Region, Bessarabia, Black Sea
Region, Lower Don, Lower Volga; Western Siberia: Ob River Area,
Upper Tobol, Irtysh, Altai; Eastern Siberia: Yenisei, Lena-Kolyma,
Angara-Sayans; Far East: Sakhalin (introduced?); Soviet Central Asia:
Aralo-Caspian Region, Lake Balkhash Region. General distribution:
Scandinavia, Central Europe. Described from Lithuania. Type in
Leningrad.

Note. This species should perhaps be more appropriately called A.
speciosa Spreng. in Henckel, Adumbr. plant (1806) 1. However, certain
facts go against it. All specimens of A. speciosa Spreng. preserved in
the herbarium of the Botanical Institute, Academy of Sciences of the
USSR, were raised in different botanical gardens in the first half of the
19th century. There are no plants collected under natural conditions,
and also in the description of the species it is stated that its native
country is unknown. The specimens are tall plants with thick angular
stems, large punctate-glandular leaves, and large capitula. There are
no such large plants among the materials of A. cartilaginea Ldb. pre-
served in the Herbarium of the Botanical Institute, Academy of Sci-
ences of the USSR, and collected under natural conditions. Besides, as
was mentioned above, all the plants identified as A. speciosa Spreng.
have leaves covered with punctate-glandular hairs, whereas this char-
acter is not mentioned in the original description of A. speciosa Spreng.,
and I did not have the type specimen at my disposal. Therefore, the
species under consideration is named A. cartilaginea Ldb., and the
species name A. speciosa Spreng. should be reserved for the large
plants cultivated in botanical gardens.

38. A. septentrionalis (Serg.) Botsch. comb. nova.—Ptarmica
cartilaginea Ldb. ssp. septentrionalis Serg. in Fl. Zap. Sib. XI (1949)
2728.—Achillea ptarmica L. speciosa Schmalh. FI. II (1897) 63, non
A. speciosa Henckel (1806).—Ptarmica speciosa Stank. in Stank. and
Taliev, Opred. (1949) 626.—P. speciosa auct. non DC. (1837); Ldb.
FiwRoss. si b2 5 30spmp:

Perennial. Rhizome short, plants green, sparsely pubescent from
curly hairs, or stem and leaves glabrous and inflorescence branches
and involucral bracts pubescent; stem erect, 30-125 cm high, leafy,
simple, branched only in inflorescence or throughout, and then lower
branches reduced, nonflowering, upper terminating in capitula. Leaves

116 0.3-11.0 cm long and 0.5—22 mm wide, reduced from the base to stem

111

apex, sessile, linear-lanceolate, entire, biserrate-dentate, flat, lacking
punctate glandular hairs; lower leaves withering before anthesis.
Inflorescence corymbose; capitula semiglobose, 6—7 mm long, 4-6 mm
in dia (excluding ligules), Involucre imbricate, 3-seriate; involucral
bracts 2.5-3.8 mm long, 1.2-1.7 mm wide, broadly lanceolate,
herbaceous in middle, green, with prominent midrib, membranous on
margin, dark brown. Receptacular scales like involucral bracts, reduced
from periphery to center. Outer florets 8, pistillate, ligulate, 4.7—6.0
mm long with isolated capitate glandular hairs along tube; ligules white
throughout or with violet base, oval, 3.2-4.0 mm long and 2.4—3.2 mm
wide; with 3 obtuse teeth at apex; disk florets white, 5-toothed, tubular,
bisexual, 2.5-3.0 mm long, with capitate glandular hairs. Achenes
oblanceolate, about 2.4 mm long, 1.2 mm wide. Flowering June to
August.

In flooded meadows, along riverbanks among shrubs and flood-
plain forests.—European part: Dvina-Pechora, Upper Volga, Volga-
Kama, Upper Dnieper, Volga-Don, Lower Volga; Western Siberia: Ob
River Area. Endemic. Described from several places in Western Sibe-
ria. Type in Tomsk.

Note. A. septentrionalis (Serg.) Botsch. is similar to A. cartilaginea
Ldb., especially to individuals that grew in very humid conditions and
have weakly pubescent leaves. However, unlike these plants, A.
septentrionalis (Serg.) Botsch. does not have punctate-glandular hairs
on leaves. Earlier it was believed that A. speciosa Spreng. had the
combination of characters listed above, and our plant was so identified.
However, it would now be more appropriate to consider that A. speciosa
Spreng. has punctate-glandular hairy leaves and is a unique garden
form close to A. cartilaginea Ldb. As for plants with eglandular leaves,
they are placed in the separate species—A. septentrionalis (Serg.)
Botsch.

Series 5. Alpinae Botsch.—Leaves pinnatifid or pinnately cut, with
punctate glandular hairs above or without them, but with long colorless
straight hairs. Inflorescence corymbose.

39. A. impatiens L. Sp. pl. (1753) 898; Ldb. Fl. Alt. IV, 123;
Heimerl in Denkschr. Acad. Wien. XLVIII, 127 (Monogr. sect.
Ptarmica, 15) in clave.—Ptarmica impatiens DC. Prodr. VI (1837)
22; Ldb. Fl. Ross. II, 2, 527; Turcz. in Bull. Soc. Nat. Mosc. XXV,
429; Sergievskaya in Fl. Zap. Sib. XI, 2731; Popov, Fl. Sr. Sib. II,
725.—Achillea impatiens L. ssp. euimpatiens Heimerl op. cit. 185
(Monogr. 73) p. p., excl. pl. transsilvanicae.—Exs.: GRF No. 1117.

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112

Perennial. Rhizome short; stem erect, 35-130 cm high, leafy,
branched only in inflorescence, glabrous below, in middle sparsely |
and in upper part and inflorescence densely covered with colorless,
rather long, appressed straight hairs. Leaves sessile, lanceolate, green,
glabrous above, with colorless, scattered, appressed, straight hairs
beneath, 1-7 cm long, 2-20 mm wide, pinnately cut, segments usually
distant, slightly longer than wide, linear; upper leaves entire, lower
with 1-2 linear teeth closer to base of upper margin and sometimes
(mainly in lowermost leaves) with 1-2 smaller deltoid teeth at base of
lower margin; segments and teeth serrulate-denticulate; teeth and apex
cartilaginous-cuspidate; lower leaves withering before flowering.
Inflorescence corymbose; capitula semiglobose, 4-7 mm long and 4—
7 mm in dia (excluding ligules). Involucre imbricate, 3-seriate;
involucral bracts 2.2-4.0 mm long and 1.3-1.6 mm wide, broadly
lanceolate, with scattered, rather long, appressed straight, colorless
hairs, in middle herbaceous, green with prominent midrib, membranous
on margin, dark brown or yellow, with dark brown border. Receptacular
scales, like involucral bracts, reduced from periphery to center. Outer
florets 8, pistillate, ligulate, 5.3-6.6 mm long, with isolated capitate
glandular hairs along tube; ligules oval, white throughout or with violet
base, less often violet, 3.5—4.7 mm long and 3.2-6.0 mm wide, with 3
rather large obtuse teeth at apex; disk florets white, 5-toothed, tubular,
bisexual, 2.7-3.2 mm long, with isolated capitate glandular hairs.
Achenes about 2.2 mm long and 1 mm wide. Flowering June to August.

In montane and plain forests, forest meadows.—Arctic: Arctic Si-
beria; Western Siberia: Ob River Area, Irtysh, Altai; Eastern Siberia:
Yenisei, Lena-Kolyma, Angara-Sayans, Dauria. General distribution:
Dzhungaria-Kashgaria?, Mongolia?. Described from Siberia. Type in
London.

Note. A. spinosa Schur, which is very closely related to the Sibe-
rian species A. impatiens L., grows near the city of Cluj in Romania.
It differs from A. impatiens L. by taller stems and a larger number of
teeth on the lower segments of the leaves (2—4 and not 1-2 on each
side), which was noted even in the original description. Most authors
have without basis combined A. spinosa Schur and A. impatiens L.

40. A. ledebouri Heimer! in Flora (1883) 389; ej. in Denkschr.
Acad. Wien, XLVIII, 127 (Monogr. sect. Ptarmica, 15) in clave.—A.
impatiens L. ssp. ledebouri Heimer] op. cit. (1884) 127, 184 (Monogr.
15, 74).—Ptarmica krylovii Serg. in Sistemat. Zam. Gerb. Tomsk. Univ.
1-2 (1949) 13 and in Fl. Zap. Sib. XI, 2730.—P. tenuisecta (Kryl.)
Serg. in Sistemat. Zam. Gerb. Tomsk. Univ. 1-2 (1949) 13 and in FI.
Zap. Sib. XI, 2730.—P. alpina DC. f. tenuisecta Kryl. sec. Serg. in

118

113

Sistemat. Zam. Gerb. Tomsk. Univ. 1—2 (1949) 13 and in Fl. Zap. Sib.
XI, 2731.—Achillea alpina auct. non L. (1753); Ldb. Fl. Alt. [V (1833)
123.—Ptarmica alpina auct. non DC. (1837); Ldb. Fl. Ross. II, 2,
528; Kryl. Fl. Alt. III, 614.

Perennial. Rhizome rather long, horizontal; stem erect, 20—75 cm
high, leafy, usually branched only in inflorescence, glabrous below, in
middle sparsely and in inflorescence densely covered with rather long,
appressed, straight, colorless hairs. Leaves 0.5-5.2 cm long, 2-11 mm
wide, sessile, lanceolate, pinnately lobed, pinnately parted, or pinnately
cut, with scattered, rather long, appressed, straight, colorless hairs on
both sides; lobes, or segments, linear, acute, distant, at a distance more
or less equaling their width, serrulate-denticulate, sometimes with a
large tooth on upper margin, cartilaginous-cuspidate at apices and teeth;
lower leaves withering before flowering. Inflorescence corymbose;
capitula fewer, semiglobose, 6-7 mm long and 6—7 mm in dia (excluding
ligules). Involucre imbricate, 3-seriate; involucral bracts 3.1-5.0 mm
long and 1.6-—2.0 mm wide, broadly lanceolate, with scattered, rather
long, straight, appressed colorless hairs, herbaceous in middle, green,
with prominent midrib, membranous on margin, dark-brown.
Receptacular scales, like involucral bracts, reduced from periphery to
center. Outer florets 8-pistillate, ligulate, 6.2-8.5 mm long, glabrous;
ligules white, oval, 4.5-6.0 mm long, 3.5-5.5 mm wide, with 3 obtuse
apical teeth; disk florets white, 5-toothed, tubular, bisexual, 3.2—3.3
mm long, glabrous. Achenes broadly lanceolate, 2.1-2.7 mm long and
0.8-1.0 mm wide. Flowering July to August.

In alpine and subalpine meadows.— Western Siberia: Altai. General
distribution: Dzhungaria-Kashgaria. Described from Altai. Type in
Vienna; isotype in Leningrad.

Note. It was customary to call this species A. alpina L. (see Ldb.
Fl. alt. 1V, 123). However, the description of the species by Linnaeus
did not correspond to the Altai plants, as indeed shown by Heimerl
(1. c. 1883), who described the Altai plants, accepted by Ledebour as
A. alpina L., as a new species—A. ledebouri Heimerl. Later, Heimer]
(1. c. 1884) considered his species as a subspecies of A. impatiens L.
and drew attention to the fact that the material used by him as type
material for describing the new species (Karelin and Kirilov, 1840,
No. 782) also included typical A. impatiens L.

L.P. Sergievskaya described Ptarmica krylovii Serg. and P.
tenuisecta Serg. (1. c.) from Altai. Besides the type specimens of these
species, I have studied a fairly large number of plants and am convinced
of the following:

1. Even the isotypes of A. krylovii Serg., contrary to the description,
show various degrees of leaf incision (from pinnatilobate to pinnatisect).

119

114

Besides, the segments of more divided leaves sometimes also have
large teeth together with minute teeth; in other words, even the isotypes
of P. krylovii Serg. have characters of both the “species” described by
L.P. Sergievskaya. Re-examination of the entire material would con-
vince one about the impossibility of dividing it into two species.

2. The plants described by L.P. Sergievskaya as two different
species are indistinguishable from the type species Achillea ledebouri
Heimerl.

A. ledebouri Heimerl, apparently, sometimes hybridizes with A.
impatiens L., which explains the large variation in the degree of leaf
incision in the former species.

41. A. alpina L. Sp. pl. (1753) 899; Willd. Sp. pl. III, 3, 2193
p. p. quoad pl. sibiricam.—A. mongolica Fisch. ex Spreng. Novi proven
hort. Acad. Halen. et Berol. (1818) 3; Heimerl in Denkschr. Acad.
Wien, XLVIII, 127 (Monogr. sect. Ptarmica, 15), in clave.—Ptarmica
alpina DC. Prodr. VI (1837) 22 p. p. quoad pl. sibiricam.—P.
mongolica DC. Prodr. VI (1837) 22; Turcz. in Bull. Soc. Nat. Mosc.
XXV, 430.—P. sibirica Ldb. Fl. Ross. II, 2 (1845) 528; Popov, FI. Sr.
Sib. II, 725.—Achillea sibirica Ldb. Ind. Sem. hort. Dorpat. (1811),
sec. Ldb. Fl. Ross. II, 2 (1845) 528 in syn.; Kom. and Alis. Opred.
Rast. Dal’nevost. Kr. II, 1027; Kitam. Compos. Japon. II, 324.—A.
sibirica ssp. mongolica Heimer] op. cit. (1884) 188 (Monogr. 76).

Perennial. Rhizome short; plants green, covered with rather long,
colorless, straight, appressed hairs; stem 30-75 cm high, erect, leafy,
branched only in inflorescence or in upper half. Leaves 0.9-9.5 cm
long, 1.8-13.0 mm wide, sessile, linear-lanceolate, pinnately lobed or
pinnately parted, punctate-glandular hairy or eglandular; lobes linear,
acute, unevenly serrate-dentate, pinnately lobed or pinnately parted;
secondary lobes and teeth cartilaginous-cuspidate, slightly involute;
lower leaves withering before flowering. Inflorescence corymbose;
capitula almost ovate, 4.5-6.5 mm long and 3.5-6.0 mm in dia
(excluding ligules). Involucre imbricate, 3-seriate; involucral bracts
2.5—4.3 mm long, 1.2-2.0 mm wide, broadly lanceolate, herbaceous in
middle, green, with prominent midrib, membranous along margin,
yellow, with dark brown border: Receptacular scales like involucral
bracts, reduced from periphery to center. Outer florets 8, pistillate,
ligulate, 3.3-4.3 mm long, sometimes with or without occasional
capitate glandular hairs along tube, ligules white, oval, 1.6—2.7 mm
long, 1.5—2.5 mm wide, with 3 weakly developed apical teeth; disk
florets white, 5-toothed, tubular, bisexual, 2.0-2.5 mm long, with or
without occasional capitate glandular hairs. Achenes broadly
oblanceolate, 2.2-2.4 mm long, 1.0-1.2 mm wide. Flowering July to
September.

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115

Along riverbanks, in meadows and marshes, in shrub thickets and
mixed forests.—Eastern Siberia: Yenisei, Lena-Kolyma, Angara-
Sayans, Dauria; Far East: Okhotsk, Zeya-Bureya, Uda River Area,
Ussuri. General distribution: Mongolia, China. Described from Siberia.
Type in London.

Note. A. alpina L., originating from Siberia, was not very clearly
described in the original description. The type of A. alpina L., pre-
served in London, was accessible with difficulty to the authors dealing
with the Siberian flora. Therefore, initially, A. alpina L. was wrongly
identified with plants later described as A. ledebouri Heimer! (cf. the
note to this species), and in recent times has been forgotten.

I had at my disposal a photograph of the type of A. alpina L. On
comparing the photograph of the type specimen of A. alpina L. with
its description in the catalogue of the London herbarium (S. Savage—
A Catalogue of the Linnaean Herbarium, 1945, 1956, No. 1017/13),
I came to the conclusion that A. alpina L. is identical with plants
identified as A. sibirica Ldb. (A. mongolica Fisch. ex Spreng.) in the
Soviet Union.

This is evident from the plant height, the shape, incision and size
of leaves, and the size and number of capitula in inflorescence.

Therefore, it is no longer necessary to resolve the question about
the nomenclatural preference of A. mongolica Fisch. ex Spreng. over
A. sibirica Ldb., nor about the futile attempts of Maximowicz (cf.
Mém. Acad. Sc. St.-Péterb., sav. étr. 1859, 154) to distinguish A.
mongolica Fisch. ex Spreng. and A. sibirica Ldb. as separate species.

Now one thing is clear: the prior name of the species under con-
sideration is A. alpina L.

42. A. japonica Heimer! in Denkschr. Acad. Wien, XLVIII (1884)
128 (Monogr. sect. Ptarmica, 16), in clave non A. japonica Sch. Bip.
(1854), nom. nudum.—A. sibirica Ldb. ssp. japonica Heimerl op. cit.
190 (Monogr. 78); Kitam. Compos. Japon. II, 327.—Ptarmica
mongolica DC. var. lilacina Fr. Schmidt in Mém. Acad. Sc. St.-Pétersb.
VII sér. XII, 2 (1868) 147.

Perennial. Rhizome long; plants green, with long, straight,
appressed, colorless hairs; stem 35-115 cm high, erect, leafy, branched
in inflorescence or in upper half. Leaves 0.5—12.5 cm long, 1-20 mm
wide, sessile, lanceolate, punctate-glandular-hairy, or eglandular,
pinnately parted or pinnately cut; lobes or segments linear, acute;
unevenly serrate-dentate, pinnately parted or pinnately cut; lobes,
segments and teeth cartilaginous-cuspidate, slightly involute; lower
leaves withering before flowering. Inflorescence corymbose; capitula
semiglobose, 5—7.5 mm long, 5-7 mm in dia (excluding ligules).

12

—

116

Involucre imbricate, 3-seriate; involucral bracts 2.6—4.3 mm long, l-
2 mm wide, broadly lanceolate, herbaceous in middle, green, with
prominent midrib, membranous on margin, yellow, with dark brown
frill. Receptacular scales like involucral bracts, reduced from periphery
to center. Outer florets 8, pistillate, ligulate, glabrous, 4.8-5.0 mm
long; ligules white or pink, oval, 3.3-3.5 mm long, 2.7—-3.5 mm wide,
with 3 obtuse teeth at apex; disk florets white, 5-toothed, tubular,
bisexual, 2.3-2.5 mm long, glabrous. Achenes oblanceolate, about 1.7
mm long, 0.6 mm wide. Flowering July to September.

Along riverbanks and sea coasts, in meadows and on herb slopes.—
Far East: Ussuri, Sakhalin. General distribution: Japan, China.
Described from Japan. Type in Leningrad.

Note. Several plants from different localities in Japan are cited in
the description of A. sibirica Ldb. ssp. japonica Heimer] (1. c.). I am
selecting as the lectotype of A. japonica Heimer! the well collected
plants with the following label: “Japonia, Insula Jesso, circa Hakodate,
Dr. Albrecht. 1861,” which are preserved in the Herbarium of the
Botanical Institute, Academy of Sciences of the USSR.

Sometimes hybrids with A. ptarmicoides Maxim. are found.

43. A. ptarmicoides Maxim. Prim. Fl. amur. (1859) 154; Heimer]
in Denkschr. Acad. Wien, XLVIII, 127 (Monogr. sect. Ptarmica, 15),
in clave; Kom. and Alis. Opred. Rast. Dal’nevost. Kr. II, 1027.—A.
sibirica Ldb. var. discoidea Rgl. in Mém. Acad. Sc. St.-Pétersb. sér.
VII, IV, No. 4 (1861) 87; Kitam. Compos. Japon. II, 325.—A. sibirica
Ldb. ssp. ptarmicoides Heimer] op. cit. 189 (Monogr. 77).—Ptarmica
ptarmicoides Worosch. in Delect. semin. hort. bot. princ. Acad. Sc.
URSS, 10 (1955) 21 and in Bot. Mat. Gerb. Bot. Inst. Acad. Nauk
SSSR, XIX (1959) 641.

Perennial. Rhizome short; plants green, sparsely covered with rather
long, straight, appressed, colorless hairs; stems 20-100 cm high, erect,
leafy, branched only in inflorescence. Leaves 0.6-10.0 cm long and
1.5-15.0 mm wide, punctate-glandular hairy, sessile, lanceolate,
pinnately cut; segments linear, acute, unevenly serrate-dentate, pinnately
cut; teeth and segments cartilaginous-cuspidate, slightly involute; lower
leaves withering before flowering. Inflorescence corymbose; capitula
usually numerous, 4-6 mm long, 3.0—5.5 mm in dia (excluding ligules),
subovoid. Involucre imbricate, 3-seriate; involucral bracts 1.5—-4.0 mm
long, 1.0-1.8 mm wide, broadly lanceolate, herbaceous in middle, green,
with prominent midrib, membranous on margin, yellow or yellow with
narrow dark brown border. Receptacular scales, like involucral bracts,
reduced from periphery to center. Outer florets 8, pistillate, ligulate,
2.2-3.3 mm long, with capitate glandular hairs along tube; ligules

122

117

white, oval, 0.7—1.5 mm long and 0.7—1.6 mm wide, sometimes erect,
with 3 rather deeply divided obtuse teeth, sometimes outer (ray) florets
1.8-2.0 mm long, tubular, with rudimentary ligule comprising 3 obtuse
teeth; disk florets white, 5-toothed, tubular, bisexual, 2.2-2.3 mm long,
covered with capitate glandular hairs. Achenes oblong or oblanceolate,
2.3-2.6 mm long and 1.0-1.3 mm wide. Flowering July to September.

Valley and mountain meadows, shrub thickets.—Eastern Siberia:
Dauria; Far East: Zeya-Bureya, Uda River Area, Ussuri, Sakhalin.
General distribution: Japan, China. Described from Amur. Type in
Leningrad.

Note. In describing A. ptarmicoides Maxim. (Il. c.), the author
included under it only those specimens at his disposal that had strongly
reduced ligules, not surpassing the disk of the capitulum. Plants that
had a well developed ligule but in other respects did not differ from
the former were identified as Ptarmica mongolica (Fisch.) DC. (lI. c.),
i.e., they were even put in a different genus. In order to be more
persuasive of such a separation of the material, he considers P. sibirica
Ldb. and P. mongolica (Fisch.) DC. (in the note to the latter) as an
independent species. However, after examining the authentic specimens
of P. sibirica Ldb. and P. mongolica (Fisch.) DC., one is convinced
that they belong to a single species (cf. above my note to Achillea
alpina L.) and also that this species differs from the material included
under it by Maximowicz.

All this discussion indicates that Maximowicz artificially separated
the specimens studied by him into A. ptarmicoides Maxim. and Ptarmica
mongolica (Fisch.) DC. on the basis of only a single character—ligule
length. If other characters are examined, then one is convinced that
this is a single species—Achillea ptarmicoides Maxim. in which the
length of ligules varies. However, other characters, in particular the
punctate-glandular hairs on the leaves, capitate glandular hairs on the
florets, nature of the division of the leaves, pubescence, etc., constitute
a set of more stable characters.

A. ptarmicoides Maxim. was described from several localities. The
following plant was selected as the lectotype: “Untern Amur, in
siidlichen Theile, bei Uchssumi, auf Mergelboden. nicht selten, 23 Juli
1855 (fl. pr.), C. Maximowicz.” preserved in the herbarium of the
Botanical Institute, Academy of Sciences of the USSR.

A. ptarmicoides Maxim. hybridizes with A. alpina L.

44. A. camtschatica Rupr. ex Heimerl in Denkschr. Acad. Wien,
XLVIII (1884) 127 (Monogr. sect. Ptarmica, 15), in clave.—A. sibirica
Ldb. ssp. camtschatica Heimerl, op. cit. 189 (Monogr. 77); Kitam.
Compos. Japan. II 325.—Ptarmica kamtschatica Rupr. ex Kom. FI.

123

118

Kamch. III (1930) 138 and Izbr. Soch. VIII, 437.—A. sibirica auct.
non Ldb.; Hultén, Fl. Kamtch. IV (1930) 169.

Perennial. Rhizome long; plants green, rather densely covered with
straight, appressed, long, colorless hairs; involucral bracts and branches
of inflorescence with ferruginous hairs; stems 12-85 cm high, erect,
leafy, branched only in inflorescence. Leaves 0.6—11.0 cm long, 2-15
mm wide sessile, lanceolate, punctate-glandular hairy, pinnately lobed,
pinnately parted, or pinnately cut; leaf segments and lobes linear, acute,
unequally sharply serrate-dentate and pinnately lobed, pinnately parted
or pinnately cut, teeth and apices of lobes and segments cartilaginous-
cuspidate, slightly involute; lower leaves withering before flowering.
Inflorescence corymbose; capitula semiglobose, 5.5—8.0 mm long, 5.5-
7.0 mm in dia (excluding ligules). Involucre imbricate, 3-seriate;
involucral bracts 2.5-4.5 mm long, 1.5-1.9 mm wide, broadly
lanceolate, herbaceous in middle, green, with prominent midrib,
membranous along margin, dark brown. Receptacular scales like
involucral bracts, reduced from periphery to center. Outer florets 8,
pistillate, ligulate, 4.8-7.0 mm long, with isolated, capitate glandular
hairs along tube; ligules white, oval, 3-5 mm long, 3.3—4.1 mm wide,
with 3 roundish teeth at apex; disk florets bisexual, white, tubular, 5-
toothed, 2.0-2.7 mm long, with isolated capitate glandular hairs.
Achenes oblong, about 3.3 mm long and 1 mm wide. Flowering July
to September.

In forests along meadow-covered slopes, on sands and gravel beds
along riverbanks, in willow stands, near lakes and on dunes near the
sea.—Arctic: Anadyr; Eastern Siberia: Lena-Kolyma; Far East:
Kamchatka (and Commander Islands), Okhotsk, Zeya-Bureya, Sakhalin
(Kuril Islands). General distribution: Beringia. Described from the
vicinity of Petropavlovsk-Kamchatskii. Type in Leningrad.

Series 6. Anthemoideae (DC.) Botsch.—Ptarmica sect.
Anthemoideae DC. Prodr. VI (1837) 19.—Leaves twice or thrice
pinnately cut, pubescence of long, scattered, straight, colorless hairs
and occasional punctate-glandular hairs beneath. Stems unicapitulate.

45. A. schuri Sch. Bip. in Oest. bot. Wochenbl. VI, No. 38 (1856)
300; Popov, Ocherk. Rast. i Fl. Karpat. 247; Klok. in Vizn. Rosl.
URSR, 543.—Anthemis tenuifolia Schur in Verhandl. Siebenburg.
Vereines II, No. 10 (1851) 171, non Achillea tenuifolia Lam. (1783).—
A. caespitosa Herbich in Flora XL (1857) 509.—Ptarmica tenuifolia
Schur, Enum. pl. Transsilv. (1866) 327.—Achillea schuri Heimer] in
Denkschr. Acad. Wien, XLVIII (1884) 125 (Monogr. sect. Ptarmica,
13), in clave.—A. oxyloba DC. ssp. schuri Heimerl op. cit. 137

124

119

(Monogr. 25).—Anthemis alpina (alpina-sphalm) auct. non L. (1759);
Baung. Enum. pl. Transsilv. III (1816) 145.—Exs.: Fl. exs. Austro-
Hung. No. 982.

Perennial. Rhizomes branched; stems usually few, unicapitulate,
ascending, simple, leafy, slender, 5—20 cm high, below with scattered
and above with dense, rather long, straight, appressed colorless hairs.
Lower and middle leaves sessile, broadly lanceolate; twice or thrice
parted into lanceolate lobes terminating in cusp, glabrous above, with
scattered, appressed, rather long, colorless hairs and punctate-glandular
hairs beneath, 0.7-3.5 mm long and 4-15 mm wide; upper cauline
leaves like involucral bracts. Capitula solitary, 0.9-1.2 cm long and
1.4—2.3 cm in dia (including ligules). Involucre 3-seriate, imbricate;
involucral bracts broadly lanceolate, 3-5 mm long and 1.5-—2.5 mm
wide, green in middle, herbaceous, with weakly developed midrib,
with rather long, appressed, colorless, sparse hairs, broadly membranous
on margin, dark brown, glabrous. Receptacular scales like involucral
bracts, reduced from periphery to center. Outer florets 9-20, pistillate,
ligulate, 5.5—8.5 mm long, with glabrous tube; ligules white, obovate,
4.0-6.3 mm long and 2—5 mm wide, obtuse or with 2-3 roundish teeth
at apex; disk florets white, 5-toothed, tubular, bisexual, glabrous, 2.2-
2.5 mm long. Achenes oblanceolate, about 2.7 mm long, 1 mm wide.
Flowering June to August.

On peaks of the Carpathians among rocks and on stony slopes.—
European part: Upper Dniester. General distribution: Central Europe.
Described from several places in Romania. Type in Vienna.

Note. A. schuri Sch. Bip. and some species that are similar but
absent from our flora were initially put in the genus Anthemis L.
on the basis of their habit, leaf shape and solitary capitula. However,
it was shown that these species, unlike Anthemis L., have flat,
winged achenes, which justified referring them to the genus Achillea
L. Here, of course, they form a separate group in section Ptarmica
(DC.) Koch.

GENUS 1517. Handelia Heimer! ':?
Heimerl in Oest. Bot. Zeitschr. LXX (1922) 215.

Capitula small, many-flowered, subglobose, in dense or lax
corymbs. Involucre 3-seriate, not imbricate; bracts oblong, obtuse,

'Treatment by L.I. Tzvetkova.
2Named in honor of the Austrian botanist H. Handel-Mazzetti.

125

120

greenish, broadly white membranous on margin, slightly pubescent
outside. Receptacle strongly convex, finely alveolate; receptacular
scales membranous, oblong. Florets bisexual, yellow, tubular, 5-
toothed, turbinate, glandular-hairy. Anthers with acute appendages.
Stigma bifid, exserted, with oblong-clavate branches. Achenes small,
papillose, turbinate, with 5 indistinct ribs, apically truncate and
with truncate, somewhat thick, weakly developed border. Leaves
oblong, thrice pinnately cut; basal leaves large, petiolate; cauline
leaves sessile, smaller. Perennial or biennial herbs, with dense
white-tomentose pubescence in lower part of plant, with tap root.
Monocarpic.
A monotypic genus.

1. H. trichophylla (Schrenk) Heimer] in Oest. Bot. Zeitschr. LKX
(1922) 215.—Achillea trichophylla Schrenk, Enum. plant. nov. I (1841)
48; Ldb. Fl. Ross. II, 2, 538; O. and B. Fedtsch. Perech. Rast. Turk.
IV, 182.—Exs.: Gerb. Fl. SSSR, No. 3261.

Perennial or triennial. Tap root, thickened in upper part. Stem one
or few, 10-100 cm high, 0.5—2 cm in dia, erect, ribbed, leafy, branched
above or only in inflorescence, at base densely white-tomentose-lanate,
more or less glabrous above. Leaves oblong, thrice pinnately cut,
segments narrow, about 0.3 mm wide, almost filiform; basal leaves
petiolate, densely pubescent, 5-30 cm long, 2—12 cm wide; cauline
leaves sessile, smaller, less densely pubescent, uppermost leaves
subglabrous. Capitula small, 3-5 mm long and in dia, in dense, less
often lax corymbs. Involucre 3-seriate, not imbricate; bracts oblong,
obtuse, greenish with broad white-membranous margin, 2-3 mm long
and 0.5-1.0 mm wide, weakly pubescent. Receptacular scales
membranous, glabrous, almost linear. Florets bisexual, yellow, tubular,
5-toothed, turbinate, about 2 mm long, 1 mm wide, glandular hairy.
Achenes brownish-yellow, 1.0-1.5 mm long, 0.3-0.6 mm wide.
Flowering May to August.

In mountains and foothills, on shallow, stony slopes.—Soviet
Central Asia: Lake Balkhash Region, Dzhungaria-Tarbagatai,
mountainous Turkmenia, Pamiro-Alai, Tien Shan. General Distribution:
Dzhungaria-Kashgaria. Described from Dzhungarian Alatau. Type in
Leningrad.

121
GENUS 1518. Santolina L. !:?

L. Sp. pl. (1753) 842.

Capitula on rather long, erect peduncles, one each at apices of
stem and lateral branches, but usually numerous, homogamous, with
numerous bisexual tubular florets. Involucre broadly cyathiform, 5-12
mm in dia and 3-6 mm long; involucral bracts coriaceous herbaceous,
imbricate in 3-4 irregular rows; outer bracts lanceolate, inner 1.5—2
times as long, oblong or linear-oblong, more or less membranous on
margin or not. Receptacle convex, usually almost subglobose, plump,
punctate-alveolate, densely squamose; scales rather hard, oblong or
linear, more or less carinate outside, obtuse or subacute. Corolla of
tubular disk florets yellow, 3-5 mm long, with 5 lanceolate recurved
teeth; corolla tube rather long, narrow, somewhat abruptly broadened
in upper 1/5-1/3, usually with 1-3 short processes at base, retrorse

126 and appressed to achene apex. Filaments rather long, glabrous; anthers
lacking distinct basal appendages, with oblong-lanceolate, subobtuse
apical appendage; pollen grains round, covered with spinules. Style
bifid; its branches narrow-linear, truncate. Achenes 1.5—2.5 mm long,
0.4-0.8 mm wide, with 3-5 fairly prominent longitudinal veins,
prismatic, narrowed toward base, lacking pappus (corona) at apex;
outer achenes often larger, with obtuse upper margin of achenes
somewhat raised on one side. Glabrous or somewhat hairy perennial
herbs and semishrubs with more or less erect or ascending (sometimes
prostrate), densely leafy stems and alternate, petiolate leaves with more
or less pinnatisect or pinnatilobate, sometimes almost entire lamina.

About 30 species of this genus are distributed in Southern Europe
and Northern Africa; some species are ornamental plants and often
grow wild.

Type of genus: Santolina chamaecyparissus L.

1. Plants grayish-tomentose; leaves 2-6 mm wide, with petiole slightly

expanded at base: i.::/.0..jA ANU wATAe 1. S. chamaecyparissus L.
+ Plants glabrous; leaves 0.8—3.0 mm wide, with petiole expanded at
aise’ GUE M. Raaliihs WAU Ret 2. S. viridis Willd.

1. S. chamaecyparissus L. Sp. pl. (1753) 842; DC. Prodr. VI, 34;
Stank. and Tal. Opred. Vyssh. Rast. Evrop. Chasti SSSR (1949) 623.—

'Treatment by N.N. Tzvelev.

*From Latin words sanctus—sacred, and linum—linseed; named for the medicinal
importance of the plant and its narrow leaves; may also be based on the ancient name
of Santones Region in France, where it is a very common plant.

127

122

S. incana Lam. FI. Fr. II (1778) 43.—Ic.: Pflanzenfam. IV, 5 (1890)
269; Hegi, Ill. Fl. VI,..2. (1929) 673.381; Javi and. Csap., Je; ii
Hung. (1933) 522.

Perennial. Plants 15-100 cm high, strongly aromatic, grayish-
tomentose from simple flexuous hairs; stems more or less woody at
base, usually strongly branched, erect or ascending, densely leafy.
Leaves 1-6 cm long, 2-6 mm wide, grayish from sparse tomentum, on
short (1/6—1/4 as long as lamina) petioles; petioles weakly broadened
at base; their lamina linear or narrow-linear, pinnately cut with numerous
lateral lobes, entire or 2—3-parted from base, linear or oblong, obtuse
or subobtuse, often not in same plane, forming pseudo-whorl. Capitula
solitary at apices of stem and lateral branches, but numerous on plant,
on long slender peduncles. Involucre 5-10 mm in dia, 3.5-5.5 mm
long; involucral bracts more or less tomentose outside. Florets tubular;
corolla 3-5 mm long, yellow. Achenes 1.5—2.0 mm long, 0.5—0.6 mm
wide, with 4, less often 3-5, prominent ribs, lacking pappus. Flowering
July to September.

Cultivated in gardens and parks as an ornamental and aromatic
plant, mostly in southern regions, at times found as an introduced or
escaped plant. General distribution: Western Mediterranean (in the
east up to Yugoslavia). Described from Southern Europe. Type in
London (Kew).

Economic Importance. Besides its ornamental (mostly as a bor-
der plant) importance, in its native country it is widely used as a
medicinal plant. It contains an insecticide.

2. S. viridis Willd. Sp. pl. III (1803) 1798; Poir. in Lam. Encycl.
VI (1804) 504; DC. Prodr. VI, 35.

Perennial. Plants 15-100 cm high, strongly aromatic, wholly
glabrous; stem more or less woody near base, usually strongly branched,
erect or ascending, densely leafy. Leaves 1.5-6.0 cm long, 0.8—3.0
mm wide, on shorter (1/8—1/4 as long as lamina) petioles; petioles
basally broadened; lamina narrow-linear, pinnately parted or pinnately
cut; lateral lobes numerous, entire or 2—3-parted from base, obtuse or
subacute, oblong-linear to obovate; often arranged not in a same plain,
forming pseudowhorl, very short on vegetative branches and usually
appressed to leaf axis. Capitula and achenes as in previous species.
Flowering June to September.

Cultivated as an ornamental and aromatic plant primarily in the
more southern regions; rarely escaped.—European part: Crimea (south-
ern part); Caucasus: Ciscaucasia. General distribution: Western Medi-
terranean. Described from Southern Europe. Type in Berlin.

128

123

Note. This species is often combined with the previous species as
a form or variety. However, it differs from the former species not only
in the absence of pubescence but also in leaf shape and, in my view,
it fully deserves the status of a separate species.

GENUS 1519. Otanthus Hoffmgg. and Link !:?

Hoffmgg. and Link, Fl. Port. II (1809) 36.—Diotis Desf. Fl. Alt.
II (1798) 260, non Schreb. 1791.—Neesia Spreng. Anleit. II, 2 (1818)
547, non Neesia Blume, 1835, nom. conserv.

Capitula (1)2—10(12), on short (1-14 mm long) thickened peduncles
in rather dense, terminal corymbs, homogamous with numerous bisexual
tubular florets. Involucre broadly poculiform, 6-10 mm in dia and
4.5-6.0 mm long, involucral bracts coriaceous-herbaceous, more or
less lacking membranous border, imbricate in 3-4 irregular rows; outer
bracts ovate, inner oblong, 1.5-2 times as long. Receptacle convex,
hemispherical, punctate-alveolate, squamose; scales rather stiff, oblong,
more or less carinate, usually tomentose above on outside. Florets all
tubular; corolla yellow, 3.0-4.5 mm long, with 5 lanceolate-deltoid
divergent teeth; corolla tube greatly swollen over larger lower part and
flattened dorsally, carinate along sides, covering ovary apex and
continued along its sides as 2 long (only slightly shorter than mature
achene) auriculate spurs. Filaments rather long, glabrous; anthers lacking
distinct basal appendages, but, with oblong-lanceolate subobtuse apical
appendages; pollen grains round, covered with spinules. Style bifid;
branches narrow-linear, truncate. Achenes 2-3 mm long, slightly
arcuate, with 5 (less often 4) prominent longitudinal veins, without
pappus (corona) but remaining attached to lower swollen part of corky
corolla, upper part fragile. Perennial herbs covered with dense whitish
tomentum, more or less branched, erect or ascending, densely leafy
stems, and alternate sessile leaves with oblong or linear-oblong, almost
entire lamina.

A monotypic genus.

Note. The single species of the genus, hardly, however, remaining
homogenous over its vast range, shows a definite close relationship
with the genus Santolina L., owing to its adaptation to the peculiar
growth conditions on coastal sands.

'Treatment by N.N. Tzvelev.

*From the Greek words ous (otos)—ear, and anthos—flower; so named because
of the unique structure of the corolla, whose tube is prolonged as two long ear-shaped
processes along both sides of the achene.

1

129

124

1. O. maritimus (L.) Hoffmgg. and Link, op. cit. 365.—Filago
maritima L. Sp. pl. (1753) 927.—Athanasia maritima (L.) L. Sp. pl.
(1763) 1182.—Santolina maritima (L.) Crantz, Inst. I (1766) 287;
Willd. Sp. pl. III, 1799.—S. tomentosa Lam. FI. Fr. II (1778) 41, non
Pers. 1807.—Diotis candidissima Desf. Fl. Alt. II (1798) 261; DC.
Prodr. VI, 34.—D. maritima (L.) Sm. Engl. Fl. HI (1825) 403; Boiss.
Fl. or. III, 253; Kolak. Fl. Abkh. IV, 229; Grossh. Opred. Rast. Kavk.
457.—Neesia maritima (L.) Spreng. ex Steud. Nom. ed. 2, II (1841)
188.—Ic.: Rchb. Ic. Fl. Germ. XVI (1854) t. 998; Pflanzenfam. IV, 5,
269; Bonnier, Fl. Compl. France, Suisse et Belg. V, t. 298.

Perennial. Plants 10-30 cm high, usually strongly branched at base,
covered with dense whitish tomentum; stem erect or ascending, densely
and uniformly leafy. Leaves 7-18 mm long and 2.5-6.0 mm wide,
sessile, with slightly broadened base; lamina oblong or linear-oblong,
somewhat thick, without distinct veins, subobtuse, obtusely denticulate
(teeth usually hidden in dense tomentum). Capitula in rather dense
apical corymbs of (1)2—10(12) capitula. Involucre 6-10 mm in dia,
4.5-6.0 mm long, more or less tomentose on outside. Florets all tubular;
corolla tubular yellow, strongly swollen in lower part, with 2 long,
downward directed auricles, firmly decurrent on sides of ovary (and
mature achene); achenes 2-3 mm long, adherent with lower swollen
part of corolla, lacking pappus. Flowering July to September.

On coastal sands. —Caucasus: Western Transcaucasia (Abkhazia).
General distribution: Atlantic Europe, Mediterranean, Balkans.
Described from Southern Europe. Type in London (Kew).

Note. The locality of this species in Abkhazia is the only place in
the Black Sea basin, and apparently the plant was introduced here.

Economic Importance. May be used as an ornamental plant.

Subtribe 2. CHRYSANTHEMINAE O. Hoffm. in Pflanzenfam.
IV, 5 (1889-1894) 273.—Trib. Senecionideae Less. subtrib. VI.
Chrysanthemeae Less. 2. Chrysanthemeae Less. Synops. Comp. (1832)
250 and subtrib. VII. Artemisieae Less. 2. Cotuleae Less. and 4.
Artemisieae Less. op. cit. 260, 263.—Receptacle glabrous or hairy,
but lacking scales.

130

125
GENUS 1520. Argyranthemum Webb. Ex Sch. Bip. ':”

Sch. Bip. in Webb. and Berth. Phyt. Canar. II (1842) 258.—Chry-
santhemum sect. Magarsa DC. Prodr. VI (1837) 65.—C. sect.
argyranthemum (Webb. ex Sch. Bip.) Benth. and Hook. f. Gen. pl. II
(1876) 426, p. p.; Hoffm. in Pflanzenfam. IV, 5, 278 p. p.

Capitula numerous (up to 50 or more), solitary at apices of leafy
branches, heterogamous, with 10-25 pistillate, ligulate peripheral florets
in one row, and numerous bisexual tubular disk florets. Involucre
patelliform, 10-18 mm in dia, and 4—6 mm long; involucral bracts
coriaceous-herbaceous, imbricate, in 3—4 irregular rows; outer bracts
lanceolate-ovate, subobtuse, with rather wide, light colored,
membranous border; inner bracts oblong, 2—2.5 times as long as outer,
with large, broadly oval, entire or more or less discontinuously toothed
membranous appendages, at apex, decurrent on margins of bracts.
Receptacle strongly convex, obtusely conical, plump, glabrous, weakly
punctate-tuberculate with scarcely visible alveolation. Corolla of ligulate
florets white, with strongly flattened tube but lacking auricles, 1.5—2.5
mm long and linear or oblong-linear limb, 8-15 mm long; corolla of
tubular disk florets yellow, 2.5—3.5 mm long, with tube narrowed in
lower part, rather abruptly broadened in upper part, with 5 deflexed
triangular teeth, 1/6-1/5 as long as tube. Filaments thicker in upper
part; anthers lacking distinct basal appendages, but with lanceolate-
ovate, subobtuse apical appendages; pollen grains round, spiny. Style
bifid; branches linear, truncate. Achenes heterogenous, glabrous, in
peripheral florets 3.0-4.5 mm long and 2.5-3.5 mm wide, with three
prominent, broad winged ribs (ventral and lateral) and scarcely visible
veins inbetween, strongly truncate in upper part, with base of corolla
enclosed in shallow depression, in tubular florets 1.8—2.5 mm long and
0.8—1.2 mm wide, more or less flattened laterally, with 5—8 longitudinal
ribs, of which one ventral rib (in outer achenes winged) prominent, in
others weakly developed, with truncate (longer on ventral side) corona
at apex up to 0.4 mm long, usually shallow and obtusely lobed
depending on number of ribs. Glabrous or subglabrous (with short
simple hairs), strongly branched semishrubs with branches woody to a
considerable length and alternate leaves with more or less divided or
lobed laminas.

The genus comprises 12-15 species occurring almost exclusively
on the Canary Islands; one of them is cultivated as an ornamental
plant.

'Treatment by N.N. Tzvelev.
2From the Greek words argyros—silver, and anthemos—flower; so named on the
basis of the silvery membranous appendages on the inner involucral bracts.

13

—_

126

Type of genus: Argyranthemum frutescens (L.) Sch. Bip.

Note. It shows a especially close relationship with the genus Chry-
santhemum L. and in relation to it appears more “ancient,” i.e., closer
to the ancestor of both genera.

*A. frutescens (L.) Sch. Bip. op. cit. 264, t. 91.—Chrysanthemum
frutescens L. Sp. pl. (1753) 877.—Pyrethrum frutescens (L.) Willd.
Sp. pl. III (1803) 2150; DC. Prodr. VI, 65.—Ic.: Sch. Bip. |. c.; Hegi,
Til Fle VI, 2:(1927) 595:

Perennial. Glabrous or subglabrous semishrub, up to 1 m high;
stems strongly branched above base, woody to considerable length,
erect. Leaves with inconspicuous punctate-glandular hairs, up to 10—
12 cm long, 4—5 cm wide, on rather long, narrow-winged petiole, basally
thickened and often with 1-2 very small lateral lobes on each side;
lamina ovate to oblong, once or twice pinnately parted up to very
narrow-winged axis, with 2—5 distant segments of different size and
form on each side; apical lobes linear or lanceolate, 0.6—4.0 mm wide,
with rather long cartilaginous cusp; uppermost leaves strongly reduced,
partly entire. Capitula numerous, solitary on leafy branches, sometimes
in corymbs, on rather long (to 15 cm long) peduncles. Involucre 10—
18 mm in dia, 4-5 mm long; inner involucral bracts with large, light-
colored (silvery) membranous apical process. Ligulate florets white,
with 8-15 mm long limb; tubular florets yellow. Achenes heterogenous;
in ligulate florets with 3 winged ribs, in tubular florets with 1 prominent
(often winged) and 4—7 weak ribs; corona up to 0.4 mm long, truncate.
Flowering June to October.

Cultivated as an ornamental plant (in gardens and flower beds in
the south, as a potted plant in the north). Native to Canary Islands.
Described from the Canary Islands. Type in London.

Economic Importance. A common ornamental plant. There are
many varieties that differ in the size of the capitulum, shape of the
leaves, and general morphology. It is distinguished by a particularly
long flowering period, according to Kiselev (Tsvetovodstvo, ed. II,
1953, p. 702), flowering outdoors almost throughout the summer until
late autumn, and indoors often throughout the winter.

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127
GENUS 1521. Chrysanthemum L.':?

L. Sp. pl. (1753) 887 p. p.—Ismelia Cass. in Dict. Sc. nat. XLI
(1826) 40.—Glebionis Cass. ibid. 41.—Pinardia auct. non Cass.: Less.
Syn. Compos. (1832) 255 p. p.—Xanthophtalmum Sch. Bip. Tanacet.
(1844) 17.—Chrysanthemum sect. Pinardia (Cass.) Boiss. Fl. or. III
(1875) 336 p. p.; Benth. and Hook, f. Gen. pl. II, 425 p. p.; Hoffm.
in Pflanzenfam. IV, 5 (1889) 277 p. p.

Capitula solitary or 2-10 on each stem and its leafy branches,
heterogamous, with 10—30 (in cultivated varieties sometimes more than
30) pistillate ligulate peripheral florets in one row, and numerous
bisexual tubular disk florets. Involucre patelliform or broadly
poculiform, 1.3-3.0 cm in dia, 6-8 mm long; involucral bracts
coriaceous-herbaceous, imbricate in 3—5 irregular rows; outer bracts
ovate, subobtuse, with narrower membranous border; inner bracts
oblong, 1.5—3 times as long as outer, with large, broadly oval, entire
or more or less discontinuously toothed apical process membranous
and decurrent on margins of involucral bracts. Receptacle strongly
convex, almost hemispherical, plump, glabrous, weakly punctate-
tuberculate, with inconspicuous alveolation. Corolla of ligulate florets
yellow (sometimes white in cultivated varieties), with strongly flattened
but wingless or more or less wingless, 2.5-5 mm long tube and oblong
or linear 8-25 mm long limb; corolla of tubular florets yellow or red,
3-5 mm long, tube narrowed in lower part and rather abruptly broadened
above, with 5 deflexed triangular teeth 1/7-1/5 as long as tube.
Filaments in upper part slightly thickened; anthers without distinct
basal appendages, with oblong-ovoid, obtuse, or subobtuse apical
appendage; pollen grains round, spiny. Style bifid; branches linear,
truncate. Achenes heterogenous, glabrous, 2.0-4.5 mm long, 1-4 mm
wide, those of ligulate florets with 3 or 2 prominent, winged,
longitudinal ribs and somewhat inconspicuous, 2-6 ribs in-between,
often slightly truncate at apex, of tubular florets with 6-12 more or
less uniform ribs, of which usually 1 ventral rib (often winged)
prominent or 1-2 prominent ribs (dorsal and ventral) and rest almost
indistinct; pappus absent; secretory canal and mucilaginous cells also.
Glabrous or slightly hairy (with sparse simple hairs) annual plants with
tap root, erect, usually weakly branched stem and alternate leaves with
more or less pinnatisect or lobate lamina.

'Treatment by N.N. Tzvelev.
7From the Greek words chrysos—golden, and anthemos—flowers, named for the
golden-yellow color of the ligulate florets in many species of this genus.

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128

Four out of five species of this genus, mainly distributed in the
countries of the Mediterranean, are widely cultivated in various parts
of the world and are often found as introduced or escaped plants.

Type of genus: Chrysanthemum coronarium L.

Note. In many botanical publications of the past as well as the
present, a whole series of general belonging to the tribe Anthemideae
(Chrysanthemum L., Leucanthemum Mill., Pyrethrum Zinn., Tanacetum
L., Brachanthemum DC. and many others) is combined under the
generic name “Chrysanthemum L.” However, with such a broad concept
of the genus, the borders between not only the genera Chrysanthemum
L. sensu lato and Matricaria L. (the combined genus is called
““Matricaria L.”’) but also the genera Chrysanthemum L. sensu lato and
Artemisia L. are lost. Thus, successive “consolidation” of genera
inevitably would lead to combining all genera of the tribe Anthemideae
with a glabrous or hairy receptacle into a single genus and also
contrasting them to the aggregate genus with scaly receptacles, but this
single character separating the genera would appear to be very artificial.
Therefore, we believe, the only correct way is to divide the species
included in these genera into a number of natural smaller genera;
however, this should be done, of course, on the basis not of a single
character but the entire set of characters, mainly the characters of the
reproductive organs. Greater attention should be paid especially to the
anatomical characters of the achene and to the structure of the pollen
grains, which have been used to a very limited extent in the present
study because of the shortage of time.

The genus Chrysanthemum L. sensu stricto occupies a well
delineated position in the tribe Anthemideae, showing a close
relationship only with the genus Argyranthemum Webb. ex Sch. Bip.,
and is much more distant from the genus Dendranthema (DC.) Des
Moul. The three sections into which the genus is divided are very
natural and well separated; however, in spite of the significant differ-
ences in the achene structure, their close affinity is beyond any doubt.
Section [smelia is closest to the genus Argyranthemum on the basis of
achene structure, and may be considered the most “primitive” in com-
parison with the two other sections.

Economic Importance. The three species of the genus are culti-
vated in many countries as ornamental plants, and one (C. roxburghii)
is cultivated in the countries of Southeast Asia as a vegetable.

1. | Achenes of peripheral ligulate florets with 2 prominent, winged,
lateral longitudinal ribs; leaves entire, irregularly coarsely toothed
to pinnatilobate (Sect. Xanthophtalmum) ............ 4. C. segetum L.

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129

+ Achenes of peripheral ligulate florets with 3 winged longitudinal
ribs (ventral and lateral); leaves once or twice pinnately parted
eee PAARARGR A RRA. ARR ARIAL IR REA 2

2. Achenes with 2-3 broad winged ribs; intermediate ribs indistinct or
almost so; corolla of tubular florets dark red (Sect. [smelia) .........
en RN NIA. Ee 1. C. carinatum Schousb.

+ Achenes with distinct but less prominent intermediate ribs; corolla

of tubular florets variously colored (Sect. Chrysanthemum) ........ 3.
3. | Leaves thrice pinnately parted or divided ....... 2. C. coronarium L.
+ Leaves pinnately parted, with irregularly coarsely toothed, less often
spatiale lOUES IA QTR AUN ee 3. C. roxburghii Cass.

Section 1. Ismelia (Cass.) DC. Prodr. VI (1837) 65.—Ismelia Cass.
1. c-—Achenes of peripheral ligulate florets with 3 broad-winged ribs,
especially ventral, projecting far above base of floret as acute winged
process; intermediate ribs indistinct or only almost so dorsally; achenes
of tubular disk florets strongly compressed laterally, with 2 prominent
ribs (dorsal and especially ventral); other ribs indistinct or only scarcely
visible. Corolla of tubular florets dark red.

Type of section: Chrysanthemum carinatum Schousb.

1. C. carinatum Schousb. Vextr. Marokko (1800) 198, t. 6; DC.
Prodr. VI, 65.—Jsmelia versicolor Cass. Dict. Sc. Nat. XLI (1826)
41.—Ic.: Schousb. 1. c.; Hegi, Ill. Fl. VI, 2, fig. 309d-h.

Annual. Glabrous or subglabrous plants, 20-70 cm high, with short
tap root; stem erect, simple or more or less branched above. Leaves
lacking punctate glandular hairs; basal leaves withering early; middle
and lower cauline leaves to 8—10 cm long, on more or less long petioles,
but always with small auriculate lobes close to their base; lamina
obovate to oblong, 2- (less often 1-) pinnately parted up to very narrow-
winged axis, with 3-8 lobes on each side; apical lobes ovate to linear,
1-4 mm wide, cartilaginous-acute. Capitula solitary or 2-10 on leafy
lateral branches, usually not in corymbs and blooming at different
times; peduncles 10-15 cm long. Involucre 1.5—2.8 cm in dia, 6-8 mm
long, glabrous; inner involucral bracts 1.5—2.0 times as long as outer,
with large light colored or brownish, scarious apical process widely
decurrent on margins of bracts. Corolla of ligulate florets usually 2 or
3-colored, yellow, with whitish or reddish tinge, darker toward base,
less often concolorous, yellow, with 2.5—4.0 mm long tube and 15-25
mm long limb; corolla of tubular florets dark red, 3.0-4.5 mm long.
Achenes 3.0-4.5 mm long and 2.5—4.0 mm wide. Flowering July to
October.

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130

Cultivated as an ornamental plant; in the south found at times as
an introduced or escaped plant. General distribution: Northwestern
Africa. Described from Morocco. Type in Copenhagen.

Economic Importance. A few varieties differing in flower color,
capitulum size, and morphology are cultivated. Apparently, it forms
hybrids with other species.

Section 2. Chrysanthemum.—C. sect. Glebionis (Cass.) DC. Prodr.
VI (1837) 64.—C. sect. Euchrysanthemum Clarke, Compos. ind. (1876)
146.—Glebionis Cass. 1. c.—Achenes of peripheral ligulate florets with
3 prominent, narrow-winged ribs (ventral and lateral), with 1-2 much
less prominent ribs between each pair; achenes of tubular florets weakly
compressed laterally, with 1-2 strong prominent longitudinal ribs
(ventral and dorsal) and 5-8 thin ribs. Corolla of tubular florets yellow
of various shades.

Type of section: type of genus.

2. C. coronarium L. Sp. p. (1753) 890; DC. Prodr. VI, 64; Boiss.
Fi. or. III, 336; Trautv. in Tr. Peterb. Bot. Sada, VIII, 2, 458; Schmalh.
Fl. II, 68; Kom. and Alis. Opred. Rast. Dal’nevost. Kr. II, 1028.—
Matricaria coronaria (L.) Desr. in Lam. Encycl. III (1792) 737.—
Pinardia coronaria (L.) Less. Syn. Compos. (1832) 255.—Ic.: Bonnier,
Fl. Compl. France, Suisse et Belg. V (1921) t. 295; Hegi, Ill. Fl., VI,
2, fig. 306.

Annual. Glabrous or subglabrous plants, 25—70 cm high, with short
tap root; stem erect, simple or more or less branched above. Leaves
lacking punctate glandular hairs; basal leaves withering early; middle
and lower cauline leaves to 8-10 cm long, sessile, oblong or oblong-
obovate, twice pinnately parted up to narrow-winged axis, with 4-10
lobes on each side, usually broadest in the upper third, gradually
narrowed to narrow-winged base, with more or less developed auricles,
apical lobe ovate to linear, cartilaginous-acute; upper leaves smaller
and less divided. Capitula solitary or in groups of 2-8 on leafy lateral
branches, usually not in corymbs; peduncles to 15-20 cm long. Involucre
1.5-3.0 cm in dia, 6-8 mm long, glabrous or subglabrous; inner
involucral bracts 2—3 times as long as outer, with large light colored
or brownish, scarious, apical appendage decurrent on margins of bracts.
Corolla of ligulate florets yellow of various shades, sometimes (under
cultivation) white, with 3-5 mm-long tube and 15—25 mm-long limb;
corolla of tubular florets yellow, 3.5-5.0 mm long. Achenes 2.8—4.0
mm long, 2.0-3.5 mm wide. Flowering July to October.

Cultivated as an ornamental plant; at times found as introduced or
escaped plant. General distribution: Mediterranean. Described on the

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basis of garden specimens originating from the Island of Crete (or
Sicily). Type in London.

Economic Importance. There are a large number of varieties of
this species known in cultivation since the 16th century. They differ
mainly in flower color, capitulum size, shape of leaves, and time of
flowering. There are “double” varieties with tubular florets entirely or
partly modified into ligulate florets.

3. C. roxburghii Cass. in Dict. Sc. Nat. XLI (1826) 41; Desf. Cat.
Hort. Par. ed. III (1829) 170; DC. Prodr. VI, 64; Kom. and Allis.
Opred. Rast. Dal’nevost. Kr. II, 1028.—C. coronarium auct. non L.:
Kom. in Tr. Peterb. Bot. Sada, XX V (1907) 644.—Matricaria oleracea
Buch.—Ham. in Wal. Catal. pl. East Ind. Mus. (1828) 111, nom. nud.—
Pinardia roxburghii (Cass.) Less. in Ind. III Sem. Hort. Petrop. (1836)
20.—Glebionis roxburghii Cass. ex DC. 1. c. in syn.

Annual. Glabrous or subglabrous plants, 20-60 cm high, with short
tap root; stem erect, simple or more or less branched above, usually
thicker and succulant than previous species. Leaves lacking punctate
glandular hairs; middle and lower cauline leaves 8—10 cm long, sessile,
usually with broad amplexicaul base, oblong, pinnatipartite up to
broadly winged axis; lobes of first order oblong to linear-oblong, usually
irregularly coarsely toothed, cartilaginous-subacute; upper leaves
smaller, often with entire lobes. Capitula solitary or 2-10 on each
leafy lateral branch, usually not in corymbs; peduncles to 15-20 cm
long. Involucre 1.4—2.2 cm in dia, 6-7 mm long, glabrous or
subglabrous; inner involucral bracts 2—3 times as long as outer, with
large, scarious apical appendage broadly decurrent on bract margins.
Corolla of ligulate florets yellow, with 3-S mm long tube and 8-20
mm long limb; corolla of tubular florets yellow, 3.0-4.8 mm long.
Achenes 3-4 mm long, 1.8—2.5 mm wide. Flowering July to September.

On waste ground, by roadsides, as an introduced weed (or escaped)
plant; at times cultivated as an ornamental or vegetable.—Far East:
Ussuri. General distribution: Southeast Asia; often introduced in other
countries. Described from garden-grown specimens originating from
India and China. Type in Paris.

Economic Importance. Cultivated as a vegetable in the countries
of Southeast Asia (especially in China and India); the young stems
before flowering are edible. There are varieties with strongly reduced
ligulate florets (var. breviradiatum DC. op. cit. 64) and totally lacking
ligulate florets (var. discoideum DC. op. cit. 64), which have not been
noted so far in the flora of the USSR. Since ancient times, they have
been propagated in many botanical gardens; therefore, this species can be
found, together with other annual chrysanthemums, as an ornamental

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or introduced weedy plant in other regions of the Soviet Union. Its
origin is not entirely clear; apparently, the species originated as a
result of the long-term cultivation of C. coronarium (with which it is
often combined) in the more humid and warm climate of Southeast
Asia.

Section 3. Xanthophtalmum (Sch. Bip.) Tzvel. comb. nova.—
Xanthophtalmum Sch. Bip. 1. c.—Achenes of peripheral ligulate florets
with 2 prominent, narrow-winged, lateral ribs and 3-6 approximate,
inconspicuous ribs on each side in between; achenes of tubular florets
almost cylindrical, with (9)10(12) almost similar, more or less uniformly
disposed, longitudinal ribs. Corolla of tubular florets yellow.

A monotypic section.

4. C. segetum L. Sp. pl. (1753) 889; DC. Prodr. VI, 64; Ldb. FI.
Ross. II, 559; Boiss. Fl. or. HI, 336; Schmalh. Fl. II, 67; Klok. in
Vizn. Rosl. URSR, 537.—C. laciniatum Gilib. Fl. lithuan. (1781) 218.—
Matricaria segetum (L.) Schrank, Baier, Fl. II (1789) 406; Desr. in
Lam. Encycl. meth. III, 735.—Pyrethrum segetum (L.) Moench, Meth.
(1794) 597.—Leucanthemum segetum (L.) Stank. in Stank. and Tal.
Opred. Vyssh. Rast. Evr. Chasti SSSR (1949) 630.—Ic.: Rchb. Ic. Fl.
Germ. XVI (1853) t. 986; Fedtsch. in Fler. Fl. Evrop. Ross. 974, Fig.
973; Bonnier, Fl. Compl. France, Suisse et Belg. V, t. 295; Hegi, Ill.
Fils WI,-29:fig. o3hSs

Annual. Glabrous or subglabrous plant, 20—60 cm high, with short
tap root; stem simple, less often somewhat branched, erect. Leaves
lacking punctate-glandular hairs; middle and lower cauline leaves 8-
10 cm long; lowermost leaves, petiolately narrowed at base, sessile,
amplexicaul, oblong, usually entire, irregularly coarse-toothed, less often
shallow-pinnatilobate; upper leaves smaller. Capitula solitary or 2-10
on each leafy lateral branch, usually not in corymbs; peduncles to 10—
15 cm long. Involucre 1.3—2.0 cm in dia, 5.5-7.5 mm long, glabrous,
inner involucral bracts 2—2.5 times as long as outer, with large, scarious,
apical appendage broadly decurrent on bract margins. Corolla of ligulate
florets golden-yellow, with 3-4 mm long tube and 8-15 mm long limb;
corolla of tubular florets yellow, 3.0-4.5 mm long. Achenes 2.0—2.5
mm long, 1.0-1.5 mm wide. Flowering July to September.

On waste ground, by roadsides, in cultivated fields, as an introduced
or escaped plant; often cultivated as an ornamental plant.—European
part: Baltic Region, Ladoga-IImen, Lithuania, Upper Dnieper, Middle
Dnieper, Volga-Don, Bessarabia, Black Sea Region, Lower Don;
Caucasus: Reported from Transcaucasia. General distribution: Europe
(except Arctic Region), Turkey, Northern Africa; often introduced. in
other countries. Described from Northern Europe. Type in London.

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Economic Importance. Often cultivated as an ornamental plant in
many varieties, differing mostly in the color of the ligulate florets and
the size of the capitula.

GENUS 1522. Leucanthemella Tzvel. Nom. Nov.!:?

Decaneurum. Sch. Bip. in Walp. Repert. II (1843) 918 and Tanacet.
(1844) 44, non DC. 1833.

Capitula solitary or 2-8 on apices stem and its leafy lateral
branches, heterogamous, with 10-30 pistillate, sometimes sterile (as a
result of partial reduction of pistil), 1-seriate peripheral ligulate florets,
and numerous bisexual tubular disk florets. Involucre patelliform, 10-
20 mm in dia, about 4-5 mm long; involucral bracts herbaceous,
imbricate, in 2—3 irregular rows; outer bracts lanceolate or broad-
linear, subobtuse, with narrower, brownish, scarious border; inner bracts
linear or lanceolate-linear, obtuse, usually not more than 1.5 times,
less often almost 2 times as long as outer, with rather wide, brownish,
sometimes dark-brown, scarious border, enlarged at apex as more or
less lacerate-toothed appendage along margin. Receptacle somewhat
strongly convex (to almost hemispherical), plump, glabrous, very weakly
punctate-tuberculate, with distinct alveolation. Corolla of ligulate florets
white, with strongly flattened but wingless or narrow-winged, tube
1.3—2.0 mm long, and elongate or oblong-linear, limb 10-25 mm long,
corolla of tubular disk florets yellow, 2-3 mm long, with tube narrower
in lower half, rather strongly and abruptly broadened in upper half,
with 5 weakly curved deltoid teeth, a fifth to a fourth as long as tube.
Filaments slightly thicker in upper part; anthers with scarcely visible
basal appendage and oblong-ovoid or ovoid, obtuse apical appendage;
pollen grains spherical, spiny. Style 2-parted; branches linear, truncate.
All achenes similar (but not developing in ligulate ray florets), glabrous,
2-3 mm long and about 0.8 mm wide, almost cylindrical, narrowed
toward base, with (8)10(12) prominent, almost evenly disposed
longitudinal ribs, extended at apex of achenes as subobtuse, thickened,
up to 0.3 mm long, teeth appearing like corona; mucilaginous cells and
secretory canals absent in seed coat. Perennial herbs with long branched
rhizome, lacking short nonflowering vegetative branches, with erect,
usually weakly branched stem and alternate, entire or 3-5 parted, sessile
leaves.

'Treatment by N.N. Tzvelev.
2Diminutive of Leucanthemum Mill.

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134

The two species of this genus sporadically distributed in the coun-
tries of Southeastern Europe and East Asia, are also found in the
USSR.

Type of genus: Leucanthemella serotina (L.) Tzvel.

Note. The species of this genus are similar to those of the genus
Leucanthemum Mill. in the external morphology of the achenes and
the habit of the plant, however, they are entirely different in the
anatomical structure of the achenes, which is very similar to that in
Pyrethrum Zinn. species. Occupying in a sense an intermediate position
between these two genera, Leucanthemella species have sterile ligulate
florets not typical of Pyrethrum and Leucanthemum. Considering also
the unusual ecology (Leucanthemella species are marsh plants) and
some habit peculiarities of these species, I prefer following C.H. Schultz
(Bipontinus, |. c.) and include them in a separate genus to which I give
a new name. The isolated range of the species of this genus and the
extremely sporadic distribution of each within its own range also point
to the isolation and relictual nature of this genus.

1. Leaves entire, oblong-lanceolate, coarsely toothed ............. cesses
MLS Ss AAA I, ss eae 1. L. serotina (L.) Tzvel.
+ Leaves linear, entire or 3—5-parted, with narrow-linear entire lobes
SVEN eTeR Tea siirere le Bere Crt at og Pa 2. L. linearis (Matsum.) Tzvel.

1. L. serotina (L.) Tzvel. comb. nova.—Chrysanthemum serotinum
L. Sp. pl. (1735) 888; Schmalh. Fl. II, 68.—C. uliginosum (W. and
K.) Pers. Syn. pl. II (1807) 480.—Matricaria serotina (L.) Desr. in
Lam. Encycl. III (1792) 730.—Pyrethrum serotinum (L.) Willd. Sp.
pl. III (1803) 2151; DC. Prodr. VI, 57.—P. uliginosum W. and K. in
Willd. op. cit. 2152; DC. op. cit. 57; Klok. in Vizn. Rosl. URSR,
539.—Tanacetum serotinum (L.) Sch. Bip. Tanacet. (1844) 35, nom.
altern.— Decaneurum serotinum (L.) Sch. Bip. ibid. 45.—
Leucanthemum serotinum (L.) Stank. in Stank. and Tal. Opred. Vyssh.
Rast. Evrop. Ch. SSSR (1949) 630.—Ic.: Rchb. Ic. Fl. Germ. XVI
(1853) t. 992; Hegi, Ill. Fl. VI, 2, f. 309 i, k; Jav. and Csap. Ic. Fl.
Hung. 522.—Exs.: Fl. exs. Austro-Hung. No. 2672; Fl. exs. reip.
Bohem. et Slov. No. 751; GRF No. 1367.

Perennial. Plants 30-150 cm high, with long, strongly branched
rhizome, more or less covered with short bifid and simple hairs, usually
almost glabrous; stems from base usually rather numerous, less often
solitary, erect, sparsely covered with short curly hairs, densely leafy,
simple or with few (1-8) upward directed branches above middle.
Leaves on both sides covered with punctate glandular hairs and short
curly hairs (to subglabrous), green; basal and lower cauline leaves

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135

withering before flowering; middle cauline leaves sessile, oblong-
lanceolate, to 10 cm long, cuneately narrowed toward base and here
with 1-2 pairs of linear lateral lobules (auricles), acuminate, coarsely
serrate-dentate, with acute long teeth; uppermost leaves strongly
reduced, often entire. Capitula solitary or 2—8 at apex of each leafy
lateral branch, usually not in corymbs; peduncles long, with very short
curly hairs; involucre 10-20 mm in dia and 4—5 mm long, glabrous or
subglabrous; inner involucral bracts usually not more than 1.5 times,
less often almost 2 times as long as outer, with rather wide brownish
membranous border. Corolla of ligulate florets white, with 1.5-2.0
mm-long tube and 15-25 mm-long limb; corolla of tubular florets
yellow, 2-3 mm long. Achenes 2-3 mm long, about 0.8 mm wide, with
(8)10(12) subobtuse, up to 0.3 mm-long teeth at apex; ovary of ligulate
florets lacking distinct teeth. Flowering August to September.
Marshes, marshy meadows.—European part: Middle Dnieper (Uzh
River valley in Zhitomir Region). General distribution: Central Europe
(Poland, Czechoslovakia, Hungary, Romania). Described from garden-
grown specimens originating from Southeastern Europe (Linnaeus
wrongly reported North America as its native country). Type in London.
Economic Importance. Sometimes cultivated as an ornamental plant.

2. L. linearis (Matsum.) Tzvel. comb. nova.—Chrysanthemum
lineare Matsumura in Bot. Mag. Tokyo, XIII (1899) 83, t. VI; Kom.
in Tr. Peterb. Bot. Sada XXV, 1 (1907) 643.—C. lineare var.
manshuricum Kom. ibid. 822, t. X; Kom. and Alis. Opred. Rast.
Dal’nevost. Kr. II, 1028.—Ic.: Matsum. op. cit.; Kom. op. cit.

Perennial. Plants 25-90 cm high, with long, strongly branched
rhizome, more or less sparsely pubescent with bifurcate and simple
hairs; stems many, less often solitary, erect, sparsely pubescent to
subglabrous, densely leafy, simple or branched above. Leaves green
above, lacking punctate glandular hairs and more or less glabrous, but
strongly scabrous from spinulose tubercles on both sides and along
margin, dull green beneath, sparsely pubescent, and also with numerous
fine punctate glandular hairs; basal and lower cauline leaves withering
before flowering; middle cauline leaves to 8-10 cm long, sessile, usually
with | pair of very small linear lobes at base, entire, toothless, narrow-
linear (to 3 mm, less often 5 mm wide) or 2-5 parted to narrow-
winged rachis, with few, entire, narrow-linear lobes; uppermost leaves
reduced, usually entire. Capitula solitary or 2-8 at apices of leafy
lateral branches, usually not in corymbs; peduncles rather long, covered
with very short appressed hairs. Involucre 10-15 mm in dia, 4—-S mm
long, sparsely pubescent to almost glabrous; inner involucral bracts
1.5 times, less often 2 times, as long as outer, with brownish or dark

14

—

136

brown membranous border. Corolla of ligulate florets white, with about
1.5 mm-long tube and 10-20 mm-long limb; corolla of tubular florets
yellow, 2.4-3.0 mm long. Achenes 2.4—3.0 mm long and about 0.8 mm
across, apically with 8-12 subobtuse, to 0.3 mm long teeth. Flowering
August to September.

Marshes, marshy meadows.—Far East: Ussuri. General
distribution: China (Manchuria), Korea, Japan (northern part).
Described from Japan. Type in Tokyo.

Note. Our specimens possibly differ from typical Japanese speci-
mens, forming a special geographic race (“var. manshurica Kom.’);
however, in the absence of material for comparison from Japan, it
would be premature to consider it as a separate species.

GENUS 1523. Leucanthemum Mill. !:2

Mill. Gard. Dict. ed. IV (1754); DC. Prodr. VI, 45 p. p.—Chry-
santhemum sect. Leucanthemum (Mill.) B. Fedtsch. Rast. Turk. (1915)
737.—C. sect. Pyrethrum sub-sect. Leucanthemum (Mill.) Kitam. in
Acta Phyt. et Geobot. IV (1935) 37 p. p.

Capitula solitary, less often 2-5 at apices of stem and its leafy
branches, heterogamous, with 12-40 pistillate, ligulate, 1-seriate
(sometimes many-rowed under cultivation) peripheral florets and large
number of bisexual tubular disk florets. Involucre patelliform 10—22
mm in dia, 3-5 mm long; involucral bracts herbaceous, imbricate, in
3-4 irregular rows; outer bracts lanceolate, subobtuse, with narrow
membranous border, inner linear or lanceolate-linear, obtuse, 1.5-2
times as long as outer; with more or less appendicularly broadened
membranous border at apex. Receptacle slightly convex, plump, lacking
scales, weakly punctate-tuberculate, with distinct alveolation. Corolla
of ligulate florets white, with strongly flattened, winged, 1-3 mm-long
tube and linear or oblong, 12-25 mm-long limb; corolla of disk florets
tubular, yellow, 2-3 mm long, with tube narrower and dorsally flattened
in lower half, more or less strongly and abruptly broadened in upper
part with 5-deltoid, weakly curved teeth, 1/5—1/4 as long as tube.
Filaments thicker in upper part, anthers lacking basal appendage, but
with lanceolate-ovoid subobtuse apical appendage; pollen grains
spherical, spiny. Style bifid; branches linear, truncate. Achenes
heterogenous, glabrous, 1.8—3.2 mm long and about 0.7—0.8 mm wide,

'Treatment by N.N. Tzvelev.
*From the Greek words: leukos—white, and anthemos—flower; so named be-
cause of the white color of the ligulate florets.

137

almost terete, narrowed toward base, with (8)10(12) prominent and
almost evenly disposed longitudinal ribs, much lighter in color than
rest of achene; achenes of ligulate florets distinctly flattened dorsally,
more strongly curved, with ribs more approximate ventrally, lacking
apical corona or frequently with unilateral, to 0.8 mm long, more or
less irregularly toothed corona on edge; achenes of tubular florets
lacking apical corona or corona of up to 0.3 mm long subobtuse teeth
being continuations of ribs; testa with regular rows of large mucilaginous
cells dorsally on ribs, and with longitudinal secretory canals between
ribs. Perennial herbs with more or less long rhizome, reduced vegetative
branches, erect, simple, less often weakly branched stems and alternate
leaves with entire, less often pinnately lobed lamina.

About 20 species of the genus are distributed mainly in the

mountainous regions of Southern and Central Europe; only one of
these species has a wider range, including all of Europe and the north-
ern part of the Asian continent (except the extreme north); three spe-
cies are found in the USSR.

Type of genus: Leucanthemum vulgare Lam.
Note. The genus occupies a quite isolated position, well separated

from other genera of the tribe Anthemideae, including the closest genera
Pyrethrum Zinn. and Leucanthemella Tzvel., in the unusual anatomical
structure of the achenes, which was first noted by J. Briquet and F.
Cavillier (in E. Burnat, Flora des Alpes Maritimes, VI, 1, 1916, p.

80).
142 Economic Importance. Some species of the genus are ornamental
plants.

1. Lamina of basal leaves more or less rotund; achenes of ligulate
florets with well developed, irregularly toothed, 0.5—-0.8 mm long
corona. ‘alongtimatpin. Carpathianseqns..Aaie, niche cain
ORO Tae lscc. eed 1. L. rotundifolium (W. and K.) DC.

+ Lamina of basal leaves oblong or oval; achenes of ligulate florets
lacking corona or with unilateral corona up to 0.4 mm long ... 2.

2. Involucral bracts with blackish-brown membranous border; basal
leaves with broadly oval lamina. Carpathians ..........00......008
DOI, SIE Pe ee civeuiiesavcnsbadretdhee 3. L. subalpinum (Simonk.) Tzvel.

+ Involucral bracts with light-colored or brownish membranous border;

basal leaves usually with oblong or oval lamina ...................6
OAS A SORA AL hoc DRT RRR ee ede. 2. L. vulgare Lam.

Series 1. Rotundifolia Tzvel.—Achenes of ligulate florets with

well developed. 0.5—0.8 mm-long corona, irregularly toothed on margin;
achenes of tubular florets with corona of up to 0.3 mm-long subobtuse

teeth.

143

138

1. L. rotundifolium (W. and K.) DC. Prodr. VI (1837) 46; Klok.
in Vizn. Rosl. URSR, 538.—Chrysanthemum rotundifolium W. and K.
in Willd. Sp. pl. III (1803) 2144 and Ic. pl. rar. hung. III (1812) 262,
t. 236.—Tanacetum waldsteinii Sch. Bip. Tanacet. (1844) 35, nom.
nov.—Pyrethrum rotundifolium W. and K. ex M. Pop. Ocherk Rast. i.
Fl. Karpat (1949) 248.—Ic.: W. and K. op. cit.; Rcehb. Ic. Fl. Germ.
XVI (1853) t. 989; Jav. and Csap. Ic. Fl. Hung. (1933) 523 and (1934)
t. 38.—Exs.: Herb. norm. n. s. No. 1585; Fl. exs. Austro-Hung. No.
3787; Fl. exs. reip. Bohem. et Slov. Nos. 81, 949; Fl. Cechoslov. exs.
No. 92; Fl. Polon. exs. No. 743; Pl. Polon. exs. No. 172.

Perennial. Plants 20-70 cm high, with rather long branched
rhizome, glabrous or subglabrous (with sparse, very short simple hairs);
stems solitary or few, erect, densely leafy, simple, less often with few
lateral branches. Leaves lacking punctate glandular hairs; basal leaves
8-10 cm long, petiolate, petioles almost as long as lamina and
broadened at base, lamina rotund or broadly obovate, uniformly crenate
or toothed; cauline leaves cuneately narrowed in short petiole or sessile,
lacking auricles, oblong or oval, uniformly serrate-dentate (sometimes
biserrate-bidentate), and with acute teeth at apex; uppermost leaves
reduced, sometimes entire. Capitula solitary, less often 2-5; peduncles
rather long. Involucre 13-20 mm in dia, 3-4 mm long, glabrous or
subglabrous; involucral bracts with blackish-brown membranous border,
very narrow in outer bracts, broader in inner, appendicularly broadened
at apex. Corolla of ligulate florets white, with 1.0-1.8 mm-long tube
and 12-25 mm-long limb; corolla of tubular florets yellow, 2.2—3.0
mm long. Achenes excluding corona 1.8—2.8 mm long, about 0.9 mm
wide; corona in achenes of ligulate florets 0.5—0.8 mm long, to 0.3 mm
long in tubular florets. Flowering June to August.

Meadows, scrubs, forest glades; in middle and upper mountain
zones.—European part: Upper Dniester (Carpathians). General distri-
bution: Central Europe (Carpathians). Described from the Carpathians.
Type in Vienna or Budapest.

Series 2. Vulgaria Tzvel.—Achenes lacking corona, or only ligu-
late florets with up to 0.4 mm-long unilateral corona.

2. L. vulgare Lam. FI. Franc. II (1778) 137; DC. Prodr. VI, 46;
Ldb. Fl. Ross. II, 542; Boiss. Fl. or. III, 335; O. and B. Fedtsch.
Perech. Rast. Turk. IV, 183; Kom. and Alis. Opred. Rast. Dal’nevost.
Kr. II, 1031; Grossh. Fl. Kavk. IV, 129 and Opred. Rast. Kavk. (1949)
459; Kryl. Fl. Zap. Sib. XI, 2739.—L. ircutianum DC. Prodr. VI (1837)
47; Ldb. Fl. Ross. II, 543; Turez. Fl. baic.-dahur. II 43.—L.
leucanthemum (L.) Rydb. in N. Am. Fl. XXXIV (1916) 235.—L.

144

139

georgicum D. Sosn. in Zap. Nauchno-Prikl. Otd. Tifl. Bot. Sada, VI
(1929) 14 (description in Georgian).—L. vulgare var. ircutianum (DC.)
Kryl. Fl. Zap. Sib. XI (1949) 2739.—Chrysanthemum leucanthemum
L. Sp. pl. (1753) 888; Schmalh. Fl. II, 68.—C. dentatum Gilib. FI.
lithuan. III (1781) 219.—C. ircutianum Turcz. in Bull. Soc. Nat. Mosc.
XI (1838) 94, nom. nud.—C. leucanthemum var. ircutianum (DC.)
Kryl. Fl. Alt. (1904) 618.—C. leucanthemum ssp. ircutianum (DC.)
Sukacz. in Izv. Ross. Akad. Nauk (1918) 965.—C. vulgare (Lam.)
Parsa, Fl. Iran. III (1949) 264, non Bernh. 1800.—Matricaria
leucanthemum (L.) Desr. in Lam. Encycl. meth. III (1792) 731.—
Tanacetum leucanthemum (L.) Sch. Bip. Tanacet. (1844) 35.—
Pyrethrum leucanthemum (L.) Franch. Fl. Cher. et Loir. (1885) 307.—
Ic.: Rchb. Ic. Fl. Germ. XVI (1853) t. 988; Majevski, Fl. Ed. 3, Fig.
83; Syreistsch. Ill. Fl. Mosk. Gub. III, 266; Fedtsch. and Fler. FI.
Evrop. Ross. Fig. 974; Monteverde, Bot. Atl: Tabl. 36; Hegi Il, Fl.
VI, 2, t. 265, fig. 1 and fig. 325; Zemlinskii, Lekarstv. Rast. SSSR, Ed.
2, 426; Mikhail. in Fl. Beloruss. V (1959) Plate 31.—Exs.: Fl. Gall.
et Germ. exs. No. 1236; FI. Ital. exs. No. 978; Fl. Stir. exs. No. 147;
Fl. Pol. exs. No. 181; Pl. Pol. exs. No. 170; Pl. Lit. exs. No. 19; Pl.
Finl. exs. No. 389; Pl. Bulg. exs. No. 95; GRF No. 271.

Perennial. Plant 15-80 cm high, with inclined or horizontal, often
reduced rhizome, glabrous or more or less hairy, hairs simple, very
short or longer, curly; stems solitary or few, erect, densely leafy, simple,
less often sparingly branched. Leaves lacking punctate glandular hairs;
basal leaves 10-15 cm long, long (often exceeding lamina) petiolate,
with linear or oval cuneately narrowed lamina, usually obtusely toothed
or crenate, less often shallow-lobed; lower cauline leaves short petiolate,
others sessile, cuneately narrowed toward more or less broadened (often
auriculate) base, oblong or linear-oblong, obtuse or subacute, more or
less toothed, with subobtuse or acute teeth, sometimes more or less
pinnatilobate at base; uppermost leaves reduced, sometimes entire.
Capitula solitary, less often 2—5, on rather long peduncles. Involucre
13-22 mm in dia, 3.5—5.0 mm long, glabrous or subglabrous; involucral
bracts with light colored or brownish membranous _ border,
appendicularly broadened at apex of inner bracts. Corolla of ligulate
florets white, with 2-3 mm-long tube and 10-25 mm-long limb; corolla
of tubular florets yellow, 2-3 mm long. Achenes 1.8-3.2 mm long
and about 0.8 mm wide, lacking corona, or ligulate florets with
somewhat reduced, to 0.4 mm long, unilateral corona. Flowering May
to August.

Meadows, scrubs, forest glades, sometimes as weed in fields and
kitchen gardens; to upper mountain zone.—European part: All re-
gions, except extreme north; Caucasus: All regions except Talysh;

145

140

Western Siberia: All regions except north; Eastern Siberia: Yenisei
(southern part), Angara-Sayans, Dauria (western part), Lena-Kolyma
(as an introduced plant); Far East: Ussuri and Kuril Islands (as an
introduced plant); Soviet Central Asia: Aralo-Caspian and Lake
Balkhash regions (northern part, rare). General distribution: Western
Europe, Mongolia, China (northwestern part); as an introduced or
escaped plant also in North America, Japan, New Zealand. Described
from Europe. Type in London.

Note. Despite its wide range this species is, apparently, replaced
only in the mountainous regions of Central and Southern Europe by
closely related ecogeographic races. Specimens with a more or less
developed corona on the achenes of the ligulate florets are found equally
often throughout its range and, in my opinion, have no taxonomic
importance, although they were described as a separate species (L.
ircutianum DC. 1. c.) and a variety (Chrysanthemum leucanthemum
var. auriculatum Peterm. Deutschl. Fl. 1849, p. 302). Almost throughout
its range but usually on drier places, specimens with considerably more
dense and stiff pubescence throughout (Chrysanthemum leucanthemum
var. hispidum Boenn. Prodr. fl. Mon. 1824, p. 257) and specimens with
more or less pinnatilobate cauline leaves near the base are found, which
more than once have been described as new species, varieties and forms,
but also hardly have any taxonomic significance. Specimens with capitula
lacking ligulate florets have been reported from Western Europe
(Chrysanthemum leucanthemum var. discoideum Koch, Syn. ed. 2, 1844,
p. 416); so far, these have not been found in the Soviet Union.

Economic Importance. Cultivated as a frost-resistant ornamental
plant. A “double” form with tubular florets entirely or partially modified
into ligulate florets is also found. According to S.E. Zemlinsky (op.
cit., pp. 426-427), it has long been used in folk medicine.

Another close ecogeographic race from the Pyrenees is often
cultivated as an ornamental plant.—L. maximum (Ramond) DC. Prodr.
VI, 46.—Chrysanthemum maximum Ramond. in Bull. Soc. Philom. I
(1800) 140. It is distinguished by the larger size of the piant, leaves
and capitula, serrate-dentate leaves with acute teeth, sometimes
becoming lobed, and a slightly broader, lighter colored, membranous
border on the involucral bracts.

3. L. subalpinum (Simonk.) Tzvel. comb. nova.—L. raciborskii
M. Pop. and Chrsh. in M. Popov, Ocherk Rast. i Fl. Karpat (1949)
248; Klok. in Vizn. Rosl. URSR, 539.—Chrysanthemum leucanthemum
var. subalpina Schur in Verh. und Mitth. Siebenb. Ver. X (1859) 137.—
Tanacetum subalpinum (Schur) Simonkai, Enum. Fi. Transsilv. (1886)
313.—Ic.: M. Pop. and Chrsh. op. cit..249.—Exs.: Fl. Pol. exs. No.
171 (sub L. vulgare ssp. triviale var. alpicola Gremli).

146

141

Perennial. Plants 10-30 cm high, with inclined or horizontal
rhizome, glabrous or weakly pubescent; stems solitary or numerous,
erect, simple, less often with 1-2 branches. Leaves lacking punctate
glandular hairs; basal leaves up to 5-6 cm long, long-petiolate, with
oval or broadly oval, cuneately narrowed lamina, obtusely toothed or
crenate; middle cauline leaves sessile, cuneately narrowed toward more
or less broadened base, oblong or linear-oblong, more or less toothed.
Capitula solitary, less often 2-3 on a single stem. Involucre 10-18 mm
in dia, 3.5—4.5 mm long, usually glabrous; involucral bracts with
blackish-brown membranous border, broader in inner bracts. Corolla
of ligulate florets white, with 1.5—2.5 mm long tube and 8-14 mm long
limb; corolla of tubular florets yellow, 1.7—2.5 mm long. Achenes 1.5-
2.0 mm long and about 0.6-0.7 mm wide, always without corona.
Flowering June to July.

Stony slopes, grassy areas; in upper mountain zone.—European
part: Upper Dniester (Carpathians). General distribution: Central Eu-
rope (Carpathians). Described from Eastern Carpathians. Type in Sibiu
(Romania, former Hermanstadt).

Note. Morphologically, a very weakly separated, alpine, Carpathian
race, related to L. vulgare through plants with intermediate characters. One
must also mention the great similarity of some specimens of L. vulgare
from the north of the European Part of the USSR with this species.

GENUS 1524. Coleostephus Cass. !:?

Cass. in Dict. Sc. Nat. XLI (1826) 43.—Myconia Neek. ex Sch.
Bip. in Webb. and Berth. Phyt. Canar. II (1842) 245, non Lapeyr.
1813, nec Miconia Ruiz and Pav. 1794, nom. conserv.—Chrysanthemum
sect. coleostephus (Cass.) Boiss. Fl. or. III (1875) 335; Benth. and
Hook. f. Gen. pl. II, 425 p. p.; Hoffm. in Pflanzenfam IV, 5, 278.

Capitula solitary of 2-15 at each apex of main stem and its leafy
branches, heterogamous, with 15-40 pistillate but always sterile (as a
result of partial reduction of pistil), peripheral ligulate florets in one
row, and large number of bisexual tubular disk florets. Involucre
patelliform, 8-20 mm in dia, 4-6 mm long; involucral bracts
herbaceous, imbricate, in 2, less often more or less 3, irregular rows;
linear or oblong-linear, sometimes spatulately broadened at apex,
obtuse, with or without narrow membranous border; inner bracts not
more than 1.5 times as long as outer, usually all bracts equal. Receptacle

'Treatment by N.N. Tzvelev.
*From the Greek words: coleos—sheath, and stephos—crown. Named for the
strongly truncate sheath-like pappus of the achenes.

147

142

strongly convex, obtusely conical, plump, lacking scales, slightly punc-
tate-tuberculate. Corolla of ligulate florets yellow, with strongly flat-
tened but wingless, about 1.5 mm-long tube and 6-15 mm-long, ob-
long or linear limb; corolla of tubular disk florets yellow, 2-3 mm
long, with tube narrower in lower half, strongly and rather abruptly
broadened in upper half, with 5 weakly curved triangular teeth, 1/6-
1/5 as long as tube. Filaments apically thickened; anthers lacking basal
appendages, but with lanceolate-ovoid, subacute apical appendage;
pollen grains spherical, spiny. Style bifid; branches linear, truncate.
All achenes similar (but undeveloped in ligulate florets), glabrous,
1.8-2.0 mm long and about 0.7 mm wide, almost terete, narrowed
toward base, with 8-10 somewhat prominent longitudinal ribs and very
loose hull of achenes, pappus as 1.2—1.8 mm long corona, usually
obliquely truncate over more than a half its length dorsally, auriculate.
Corona of ligulate florets large, somewhat funnel-shaped at throat.

Glabrous or weakly pubescent (pubescence of simple curly hairs)
annuals with a tap root, erect, usually weakly branched stem and
alternate, entire, more or less toothed leaves.

Of the seven species of this genus, distributed primarily in the
countries of the western Mediterranean and the Canary Islands, one
species is a widely distributed Mediterranean weed.

Type of genus: Coleostephus myconis (L.) Cass.

Note. The name Myconia Necker, Elem. I (1790) 22, used by
some authors for this genus, is not the generic name, but a mononomial
name of the species and, therefore, cannot be accepted as having
priority.

1. L. myconis (L.) Cass. in Dict. Sc. Nat. XLI (1826) 43.—
Chrysanthemum myconis L. Sp. pl. ed. 2 (1763) 1254; Boiss. Fl. or.
III, 335.—Matricaria myconis (L.) Desr. in Lam. Encycl. meth. II
(1792) 736.—Pyrethrum myconis (L.) Moench, Suppl. (1802) 247;
DC. Prodr. VI, 61; Trautv. in Tr. Peterb. Bot. Sada, VIII, 2 (1883)
455; Schmalh. Fl. II, 70.—Myconia chrysanthemum Sch. Bip. in Webb
and Benth. Phyt. Canar. II (1842) 247.—M. myconis (L.) Briq. and
Cavil. in Burnat. Fl. Alp. Marit. VI, 1 (1916) 77.—Ic.: Rehb. Ic. FI.
Germ. XVI (1853) t. 986; Hegi, Ill. Fl. VI, 2 (1929) fig. 307.

Annual. Glabrous or weakly hairy, 10-40 cm high plant, with short
tap root and erect simple or sparingly branched stem. Leaves lacking
punctate glandular hairs, entire, more or less serrate-dentate; basal
leaves on rather long petioles, with broadly oval lamina cuneately
narrowed toward base, withering early; cauline leaves usually numerous,
sessile, to 6 cm long, oval or oblong, cuneately narrowed toward base.
Capitula solitary or 2-15, one each at apices of main stem and its leafy

148

143

branches, on rather long peduncles. Involucre 8-10 mm in dia and 4—
6 mm long; involucral bracts almost equal. Corolla of ligulate florets
yellow, with about 1.5 mm-long tube and 6—15 mm-long limb; corolla
of tubular florets 2-3 mm long. Achenes 1.8-2.0 mm long, with
obliquely truncate, auriculate, 1.2—-1.8 mm long apical corona. Flowering
May to July.

By roadsides, in crops as a weed.—European part: Crimea (re-
ported from the vicinity of Sevastopol, not found at present). General
distribution: Mediterranean Region, Balkans-Asia Minor, Central
Europe (as an introduced plant in Germany and Czechoslovakia).
Described from Southwestern Europe. Type in London.

GENUS 1525. Matricaria L.!>?

L. Sp. pl. (1763) 1256.

Achenes small, (0.5)0.8-1.0(2.0) mm long, 0.25-0.5 mm wide,
cylindrical, laterally compressed, dorsally convex, obliquely truncate
at apex, with 3—4 thin ribs on ventral side, somewhat distant, brown or
light brown, smooth, without or with scarcely visible coronate, toothed
pappus. Capitula heterogamous or often homogamous (lacking
peripheral ligulate florets); peripheral ligulate florets pistillate, with
short, white, decurved ligules; tubular disk florets 4-5, spatulate, yellow
or greenish, with erect tube, narrow or swollen from middle. Anthers
with deltoid acute apical appendages. Receptacle fistular, often conical.

Annual herbs always aromatic with twice pinnate or pinnate leaves.

Type of genus: M. recutita L.

1. Capitula with peripheral ligulate florets having white ligules ...... 2.
+ Capitula homogamous, with only tubular florets «0.0.00... 3
2. Stem erect, 10-30 cm high; corona 10-20 mm in diameter; plants

wholly. glabrous 220... edie luclingmanea kan 1. M. recutita L.

+ Stem ascending, 8-15 cm high; capitula 7-10 mm in dia, whole
plant more or less hairy; leaves less divided .............cscesscsseesesseteeneeeees
OH RARE SIRS EE 2. M. tzvelevii Pobed.
3. Florets greenish, with tube without constriction; involucral bracts
white and lustrous along margin; achenes with scarcely visible,
white;4oothed' pappus:or lacking i. UA. Mai Gena aah.
ONKOL ae 3. M. matricarioides (Less.) Porter ex Britton

'Treatment by E.G. Pobedimova.
*From the Latin word: mater—mother, named for the use of the plant in women’s
diseases.

149

144

+ Florets yellow, with tube swollen and constricted in middle; involu-
cral bracts brownish along margin, less often whitish; achenes lack-
ING | PAaPPUS ~c.28s. 54h Bel ee 4. M. aurea (L.) Boiss.

Section 1. Matricaria Sch. Bip. Ueb. die Tanacet. (1844) 24.—
Chamomilla DC. Prodr. VI (1837) 51, p. p.—Capitula heterogamous,
tubular florets 5-lobed; pappus absent, or less often as scarcely visible
toothed corona.

Type of section: type of genus.

1. M. recutita L. Sp. pl. (1753) 891.—M. chamomilla L. Sp. pl.
(1763) 1256; Georgi Beschr. d. Russ. R. IV, 3, 1253; MB. FI. taur.-
cauc. II, 327; Ldb. Fl. Alt. IV, 114; DC. Prodr. VI, 50; Ldb. Fl. Ross.
II, 515; Boiss. Fl. or. III, 323; O. and B. Fedtsch. Perech. Rast. Turk.
IV, 184; Grossh. Fl. Kavk. IV, 126.—Chrysanthemum chamomilla
Patze, Mey. Elkan. Fl. Prov. Preuss. (1850) 304; Schmalh. Fl. II, 67.—
Matricaria deflexa Gilib. Fl. Lithuan. III (1785) 221; Ej. Exerc. Phytol.
I (1792) 179.—Ic.: Majevski, Fl. (1954) 580; Fedtsch. and Fler. Evrop.
Ross. 973; Kom. Sbor, Sushka i Razved. Lek. Rast. ed. 3, Plate 35.—
Exs.: Pl. Finl. exs. No. 390 and No. 978 p. p.; Fl. Polon. exs. No. 744;
Fl. Aust.-Hung. No. 3788; Fl. exs. Reip. Boh.-Sloven. Nos. 946, 1276;
Fl. Stiriaca exs. No. 687; Fl. Gal. et Germ. exs. No. 1235.

Annual. Whole plant glabrous, stem 10-30 cm; high, branched
from base or mainly sulcate in inflorescence, leafy up to top. Leaves
oblong, 15(20-30)60 mm long, 5(10-15)18 mm wide, pinnatisect or
twice pinnately cut into narrow-linear lobes with short cusp, sessile,
slightly broadened at base. Inflorescence corymbose; capitula on rather
long peduncles, 1.2(3.0-5.0) 6.5 cm long, heterogamous. Receptacle
oblong-conical, fistular. Involucral bracts usually 1-seriate, green,
obtuse, whitish-membranous along margin, lacking teeth. Ligules of
ligulate florets recurved, white; disk florets yellow, with 5-toothed
long tube. Achenes very small, 0.8-1.0 mm long, 0.25 mm wide,
concave, laterally compressed, obliquely truncate at apex, smooth, with
3 thin, white, similar ribs, extending ventrally, dorsally smooth, brown;
pappus absent, less often as scarcely visible toothed corona. Flowering
May to September.

In fields, near residential buildings, in kitchen gardens, weedy
places.—European part: All regions; Caucasus: Ciscaucasia, Dagestan;
Western Siberia: Ob River Area, Upper Tobol, Altai; Eastern Siberia:
Yenisei, Lena-Kolyma, Angara-Sayans, Dauria; Far East: Okhotsk, Uda
River Area, Ussuri (very rarely); Soviet Central Asia: Aralo-Caspian
Region, Lake Balkhash Region, Tien Shan. General distribution:
Scandinavia, Central Europe, Atlantic Europe, western Mediterranean

150

145

Region, Balkans-Asia Minor, North America. Described from Western
Europe. Type in London.

Economic Importance. The flowers contain 0.5—0.12% essential
oil, anthemic acid, and glucoside. The oil contains azulene, 2 sesquit-
erpenes and sequiterpenic alcohol. Used in medicine internally as a
diaphoretic and anticonvulsive preparation, and externally as an emol-
lient in the form of a poultice, rinse, etc. (Entsikl. slovar’ lekarstv.
e’firnomasl., i yadovitykh rast. 1951)*. In the literature (Grossh. Rast.
bogat. kavk., **2nd ed., 1952, pp. 222 and 253), it is reported as a
plant containing small quantities of vitamins A and C.

2. M. tzvelevii Pobed. in Addenda, XXV, 871.

Annual. Roots short, very hard toward autumn; stem slender,
repeatedly branched from base, 8-15 cm high, covered with short
scattered pubescence of setose straight hairs, later with pubescence
only below capitula. Leaves oblong, 15-20 mm long, 3-5 mm wide,
imparipinnate with 3-4 pairs of simple, somewhat thick, terete, white-
mucronate leaflets sparsely short-setose or glabrous, scattered over
stem and lateral branches; upper leaves with 1-2 pairs of leaflets, or
upper leaves entirely linear. Peduncles 2.0—-5.5 cm long, slender, with
one capitulum each. Capitula 7-10 mm in dia. Receptacle narrow-
conical, fistular, covered with fine white protuberances at points of
attachment of florets. Involucre usually 1-seriate; involucral bracts
broadly elliptical, with wide, white-membranous, often erose margin.
Peripheral florets ligulate, pistillate, with short, wide, white ligules,
1.5-3.0 mm long, 1.5 mm wide, three-toothed at apex, middle tooth
shorter than laterals recurved by flowering; disk florets tubular, with
5 more or less deltoid, yellow lobes and greenish tube, constricted in
middle. Anthers with very small lanceolate apical appendages. Style of
peripheral florets strongly exserted from tube, bifid at apex; style of
disk florets more or less included in staminal tube. Achenes 1.5 mm
long, 0.5 mm wide, terete, laterally compressed, obliquely truncate at
apex, with 5 thin ribs, lacking pappus. Flowering August.

Found on rubbly mountain slopes.—European Part: Crimea.
Endemic. Described from southern Crimea (Bolvan Mountain in Sudak
District). Type in Leningrad.

Section 2. Anactidea DC. Prodr. VI (1837) 50.—Capitula only
with tubular florets; corolla 4-lobed; pappus absent or with very short
entire corona.

*Encyclopedic Dictionary of Medicinal, Essential Oil and Poisonous Plants—
Translator.
**Plant Wealth of the Caucasus—Translator.

15

—

146
Type of section: Matricaria matricarioides (Less.) Brit.

3. M. matricarioides (Less.) Porter ex Britton in Mem. Torrey
Bot. Club. V (1884) 341; Majevski, Fl. (1954) 580.—Artemisia
matricarioides Less. in Linnaea, VI (1831) 210 (excl. syn.).—
Matricaria discoidea DC. Prodr. VI (1837) 50; Ldb. FI. Ross. II,
544.—Chrysanthemum suaveolens Aschers. ex Schmalh. FI. II (1897)
68.—Matricaria suaveolens (Pursch) Bush. Fl. Nordwest. Tiefebene
(1894) 496; Grossh. Fl. Kavk. IV, 126.—Cotula matricarioides Bong.
Obs. veget. Sitcha (1832) 147.—Tanacetum suaveolens Hook. FI. bor.
am. I (1833) 327, tab. 110.—Ic.: Syreistsch. Ill. Fl. Mosk. Gub. III
(1910) 270; Kom. and Alis. Opred. Rast. Dal’nevost. Kr. II, Plate
306.—Exs.: Fl. pol. exs. No. 180; GRF No. 464a, b; Pl. Finl. exs. No.
979.

Annual. Root thick, with numerous slender branches. Stem solitary
or few, erect, 5-30 cm high, mostly branched in upper half, fistular,
glabrous, sometimes pubescent only below capitula. Leaves oblong,
30(40-50)60 cm long, 5(15—18)20 mm wide, twice pinnate, divided
into linear, short, cuspidate segments, basally broadened and slightly
amplexicaul, glabrous. Inflorescence corymbose, at apices of main stem
and branches; peduncles short, 5-10-15 mm long, with single capitulum.
Capitula 7-15 mm in dia, with ovoid-conical, glabrous receptacle.
Involucral bracts 3-seriate, elliptical, obtuse, with wide, white, lustrous,
membranous margin. Florets homogamous, tubular, greenish yellow,
4-lobed, tube not extended. Achenes oblong, 1.25-—1.5—2.0 mm long,
0.5 mm wide, slightly curved at apex, weakly truncate obliquely,
dorsally smooth, brownish, ventrally with 3 thin ribs, laterally reddish,
especially above, pappus absent or with scarcely visible toothed corona.
Flowering July to October.

In fields and meadows, roadsides, in gardens, near houses, as a
weed and introduced plant, growing wild along river banks, in river
valleys and along seacoasts.—Arctic: Chukotka; European part: All
regions; Caucasus: Eastern and Western Transcaucasia; Western
Siberia: Upper Tobol, Irtysh; Eastern Siberia: Sakhalin, Kuril Islands
(wild), Lena-Kolyma, Angara-Sayans, Dauria, Far East: All regions
(wild); Soviet Central Asia: Lake Balkhash Region. General
distribution: Scandinavia, Central Europe, eastern Mediterranean
Region, North America, [?New] Zealand. Described from Unalaska.

Note 1. It grows naturally in Northeast Asia and Northwest America.
In Europe, it is a weed which has spread widely from cultivation in gar-
dens, where it was introduced in the middle of the last century.

Note 2. The plant from Lake Khanka (upper reaches of Odarka
River), collected by Chersky on July 21, 1911, is distinguished by

152

147

having very small capitula, a narrow conical receptacle, and small
leaves. It differs sharply from the Far Eastern specimens and possibly
represents an ecological form.

Economic Importance. Used as a medicinal plant like M. recutita.

4. M. aurea (L.) Boiss. Fl. or. III (1875) 324; Grossh. Fl. Kavk.
IV, 126.—M. raddeana Winkl. in Tr. Peterb. Bot. Sada, XI (1890)
122; O. and B. Fedtsch. Perech. Rast. Turk. IV, 184.—Cotula aurea
L. Sp. pl. (1763) 1257; DC. Prodr. VI, 78.—Anacyclus aureus Lam.
Ill. (1797) t. 700, non L.—Ic.: Lam. ibid.: Fiori and Paol. Ic. FI. Ital.
418; Winkl. op. cit. Plate 3, Fig. 2.—Exs.: GRF No. 3262 (sub M.
raddeana Winkl.).

Annual. Root slender, weak, with numerous branches. Stems 4—20
cm high, prostrate or ascending, solitary or numerous, branched,
glabrous, sometimes sparsely pubescent below inflorescence, sulcate.
Leaves oblong 7(15—20)50 mm long, 3(4—7)10 mm wide, pinnate, with
simple thin segments or deeply divided into thin, subacute lobes, sessile,
semiamplexicaul, basally broadened, glabrous. Capitula on more or
less long or short leafless or sparsely leafy, glabrous or sparsely
pubescent peduncles, small, 4(5-6)8 mm in dia. Receptacle ovoid.
Involucral bracts elliptical, obtuse, green, with wide, especially at apex,
brown and whitish membranous margin. All florets tubular, yellow, 4-
lobed, with tube constricted in middle, lower or upper part of tube
swollen. Achenes small, 0.5 mm long and 0.25 mm wide, oblong,
sharply obliquely truncate at apex, laterally compressed, brown, smooth,
with 3-4 white, thin, sharp ribs running ventrally, lacking pappus, in
isolated cases with long pappus. Flowering March to May. (Plate VII,
Fig. 3).

On limestone deposits, sands, saline lands, flood-plain meadows
with stony soil, sometimes as weed.—Caucasus: Eastern Transcaucasia,
Dagestan; Soviet Central Asia: Kara-Kum, mountainous Turkmenia,
Pamiro-Alai. General distribution: Atlantic Europe, Eastern and West-
ern Mediterranean Regions, Balkans-Asia Minor, Iran.

Note. Apparently, in describing M. raddeana Winkler was not
aware of M. aurea, because he compares his new species only with M.
matricarioides (M. discoidea, by his nomenclature), from which M.
raddeana, of course, is well distinguished. It must be noted that the
stems of M. raddeana plants are more erect than in M. aurea and are
not prostrate, the achenes are slightly less curved with broader ribs.
These small characters vary and cannot be used to distinguish them.

155

148
GENUS 1526. Microcephala Pobed.!:?

Pobed. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XXI (1960)
356.

Achenes 1.0—-1.5 mm long, 0.25-0.5 mm wide, terete laterally
compressed, dorsally slightly convex, with one thin rib, obliquely
truncate at apex, with 3—5 thin ribs on ventral side, brownish, more or
less densely covered with small white scales or papillae, sometimes
also pubescent, with large, a half as long as achene or slightly more,
membranous, sharply lobated, inequilateral pappus. Capitula
heterogamous; peripheral ligulate florets pistillate, with short, wide,
white ligules, uncate at end of anthesis; tubular disk florets usually
with short tube, inflated in lower half and enclosed by long pappus.
Anther appendages ovoid, almost half as long as anther. Receptacle
conical or hemispherical.

Small annual herbs of desert and semidesert areas with pectinate-
pinnate or divided leaves, and villous pubescence on both sides.

Type of genus: M. lamellata (Bge.) Pobed.

1. Capitula lacking outer ligulate florets, very rarely with them ..............
BO Ee a ., eth 3. M. subglobosa (Krasch.) Pobed.
Capitulawith:outer ligulate) floretsia3 ak A: BR 2:
2. Pappus of achenes 10-lobed, inequilateral, dorsally longer; capitula
small, 5-15 mm in dia. Plants 3-15 cm high, sparingly branched,
with moreorless/pubescentileavesi... Bieicekes. eh eee
SPs BRA Tg ori ae terest eke 1. M. lamellata (Bge.) Pobed.
+ Pappus of achenes 6-7 lobed or entire, ventrally longer; capitula
(8)15-17(22) mm in dia. Plants (10)15—20(30) cm high, weakly
pubescentio/Lanwenl ania. 2. M. turcomanica (Winkl.) Pobed.

+

1. M. lamellata (Bge.) Pobed. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 357.—Matricaria lamellata Bge. in Mém.
Sav. étr. Pétersb. VII (1854) 335; Trautv. in Tr. Peterb. Bot. Sada, I,
274; IX, 456; O. and B. Fedtsch. Perech. Rast. Turk. IV, 185.—M.
lamellata var. discoidea Winkl. in Tr. Peterb. Bot. Sada, XI (1889)
121.—Chamaemelum lamellatum Boiss. Fl. or. III] (1875) 326.—
Tripleurospermum lamellatum (Bge.) K. Rech. in Oest. Bot. Zeitschr.
XCVII (1950) 233 in obs.—Ic.: Paulsen, Stud. Veg. Pamir. (1912) 62;
Bot. Tidsskr. XXXII (1912) 53.—Exs.: H.F.A.M. No. 192; GRF No.
1873.

'Treatment by E.G. Pobedimova.
"From Greek words: micros—small, and cephalon—head.

153

149

KS

*

RS

Plate V.
1 —Tripleurospermum subnivale Pobed., habit of plant, involucral bracts,
bisexual floret, dorsal and ventral view of achene; 2 — T. szovitzii (DC.) Pobed.,
habit of plant, involucral bracts, bisexual floret, dorsal and ventral view of
achene: 3 —T. elongatum (F. and M.) Bornm., habit of plant, involucral bracts,
bisexual florets, dorsal and ventral view of achene; 4— Microcephala sub-
globosa (Krasch.) Pobed., habit of plant, involucral bracts, bisexual floret,

achene.

156

150

Annual. Plants grayish, villous with white hairs. Stem (3.5)8—10(15)
cm high, weakly branched almost from base, villous. Leaves oblong,
(10)15-—25(30) mm long, (3)5—10(12) mm wide, pectinate-pinnate or
divided into linear or lanceolate, short, and apiculate segments, villous
on both sides, petiolate; uppermost leaves usually entire, linear. Capitula
solitary, 5-15 mm in dia, on long, densely white-pubescent peduncles.
Receptacle conical, acute, narrowed, finely tuberculate. Involucre
patelliform; involucral bracts oblong-linear, slightly broadened toward
apex, inner narrowed toward base, almost transparent, with wide, white
membranous border, densely white-tomentose-lanate, ciliate. Capitula
heterogamous, with smell of chamomile; peripheral florets pistillate,
with white, wide, small ligule, 2.0—2.5-3.0 mm long, 2 mm wide,
usually decurved, especially after anthesis, unequally 3-toothed at apex,
tubular florets 5-parted, lemon-yellow, 2.5-3.0—-3.5 mm long, tube
strongly constricted in middle, swollen lower half usually enclosed by
pappus. Anther appendages ovoid, almost a half as long as anther.
Achenes laterally compressed, dorsally curved, 1.0-1.5 mm long, 0.25—
0.5 mm wide, with 4—6 ribs, densely white-squamulose; pappus 0.5
mm long, inequilateral, dorsally longer, 10-lobed, lobes acuminate,
dentate-serrate; seeds not fused with pericarp, easily separating on
soaking, fusiform, brown, 1 mm long. Flowering April to November.

In saline and sandy deserts and semideserts, on takyrs, in steppes,
and as weed in crops.—Soviet Central Asia: Aralo-Caspian, Lake
Balkhash Region, Kyzyl-Kum, Kara-Kum Amu-Darya, Syr-Darya,
Pamiro-Alai, Tien Shan. Described from southern Kyzyl-Kum (coll.
Agatma). Type in Paris.

2. M. turcomanica (Winkl.) Pobed. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XXI (1961) 357.—Matricaria turcomanica (Winkl.)
Pobed. Fl. Turkm. VII (1960) 95.—M. lamellata var. turcomanica
Winkl. in Tr. Peterb. Bot. Sada, XI (1889) 121; O. and B. Fedtsch.
Perech. Rast. Turk. IV, 185.

Annual. Plants green, weakly pubescent. Stem (10)15—20(30) cm
high, strongly branched from base, sparsely or densely pubescent.
Leaves oblong, (17)20—25(60) mm long, (8)10—15(35) mm wide,
gradually reduced upward, pinnate, with more or less oblong, 5—10
mm long linear segments, in turn usually divided, subacute, green,
very weakly pubescent or grayish-lanate. Capitula solitary, (8)15—17(22)
mm in dia, on long, weakly pubescent peduncles thickened and densely
pubescent above. Receptacle ovoid, acute, slightly narrowed at apex,
brownish. Involucre patelliform; involucral bracts oblong-linear, 2.5—
5.0 mm long, 1.5—2.0 mm wide, slightly broader above, green, weakly
carinate, with wide, white-membranous border, glabrous or subglabrous,

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151

ciliate. Capitula heterogamous, with smell of chamomile; peripheral
florets pistillate, with 4-6 mm-long and 2-4 mm-wide white ligules,
always decurved after anthesis, usually spreading at anthesis, unequally
3-toothed at apex; tubular florets lemon-yellow, 2-3 mm long, strongly
constricted in middle, swollen lower half of tube enclosed by pappus.
Anther appendages ovoid, almost a half as long as anther. Achenes
weakly compressed laterally, with 4 ribs on ventral side and usually 5
on dorsal, 1.0—-1.5 mm long, 0.25—0.5 mm wide, broader part densely,
white-squamulose; pappus 0.5 mm long, inequilateral, ventrally longer
with 6-7 acuminate lobes. Flowering April to May.

In white haloxylon forests, sandy deserts.—Soviet Central Asia:
Kara-Kum, Pamiro-Alai. General distribution: Iran. Described from
southern Turkmenia (between Ashkhabad and Merv). Type in
Leningrad.

3. M. subglobosa (Krasch.) Pobed. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XXI (1961) 358.—Matricaria subglobosa Krasch.
in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 3 (1936) 345.—M. lamellata
var. discoidea Rgl. in sched.—Pyrethrum discoideum Winkl. in sched.
non Ldb.

Annual. Stem 3—15 cm high, simple or branched almost from base,
with slender, erect branches, sparsely white-lanate. Leaves succulent,
oblong, (8)10—15(30) cm long, (2)5—7(10) cm wide, pinnate, with short
linear or lanceolate acute segments, sparsely grayish-lanate, partly green,
petiolate; uppermost leaves linear, entire. Capitula solitary, 4-10 mm
in dia, on long, sparsely, less often densely pubescent peduncles,
thickened and more strongly pubescent above. Receptacle hemispherical,
less often slightly oblong, obtuse, finely tuberculate. Involucre
patelliform; involucral bracts oblong-linear, narrow, slightly recurved,
broader at apex, green with more or less narrow membranous border,
weakly pubescent, All florets only tubular, very rarely white; ligulate
(f. radiata Krasch.), lemon-yellow, with smell of chamomile, slightly
constricted in middle, lower half of tube not inflated and not enclosed
entirely by pappus. Anther appendages ovate, acute, almost a half as
long as anther. Achenes slightly convex on outside, with 3-4 scarcely
visible ribs on ventral side, 1.0-1.5 mm long, 0.25-0.5 mm wide,
densely white-squamose in broader part; pappus 0.5 mm long,
inequilateral, with longer margin pointing ventrally, not deeply lobed,
lobes acuminate. Flowering April to June. (Plate V, Fig. 4).

In clayey or sandy deserts, alkaline meadows, haloxylon forests on
sands.—Soviet Central Asia: Lake Balkhash Region. Described from
Kazakhstan (Alekseevka on the Chu River). Type in Leningrad.

158

152

Note. This northeastern race of M. lamellata (Bge.) Pobed. is
distinguished by the absence (with rare exceptions) of ligulate florets,
a hemispherical or slightly elongated, rounded receptacle, a uniformly
broad corolla tube, a pappus with the elongated part turned to the
ventral and not the dorsal side, as well as weaker pubescence on the
plant as a whole.

GENUS 1527. Tripleurospermum Sch. Bip.!:?

Schult. Bip. Ueb. die Tanacet. (1844) 31.—Chamaemelum Vis.
Oss. Matric. an. 1, 2 (1844) 24, non Mill. 1754.

Achenes large, 1.25-3.0 mm long, 0.25-1.0 mm in dia above,
cylindrical-triangular, dorsoventrally compressed, dorsally convex, with
2 red resinous glandular hairs at apex, carinate-ribbed laterally and
ventrally, with 3 large, cartilaginous, corniculate, whitish ribs (less
often with fourth and fifth thin, scarcely visible ribs, on dorsal side),
brownish-black or light brown, usually rugose, less often smooth, with
coronate, short and entire, or long, membranous, lobed; pappus, usually
weakly or strongly prominent laterally. Capitula many-flowered,
heterogamous peripheral florets pistillate, 1-seriate, less often absent;
disk florets bisexual, tubular, 5-toothed, with ovate-deltoid yellow teeth,
usually with resinous reddish-brown dots at apex. Anthers with ovate-
deltoid or globose apical appendages.

Annual or biennial herbs with large branches and twice or thrice
pinnate leaves with linear, lanceolate or ovate, deeply divided segments.
Species of this genus grow only in the Northern Hemisphere where,
together with exotic species, they number about 30.

Type of genus: 7. inodorum (L.) Sch. Bip.

It is difficult to determine the exact number of species of this
genus in the absence of a detailed study of the entire genus, because
often Tripleurospermum species have been included in the genera
Pyrethrum, Chrysanthemum, Matricaria, Chamaemelum, etc.

However, the genus Tripleurospermum is morphologically quite
well defined and easily recognized by the structure of the achenes, but
the study of its species only on the basis of herbarium material is
extremely difficult. With reason, An.A. Fedorov writes, “... often you
see the most incorrect identifications made by both beginning botanists
and well known specialists, who evidently have fallen victim to equally

'Treatment by E.G. Pobedimova.
"From Greek words: treis—three, pleura—rib, and sperma—seed; named for the
3-ribbed achenes.

159

153

gross mistakes in identifying species of this genus” /Bot. Mat. Gerb.
Bot. Inst. Akad. Nauk SSSR, 11 (1949) 165]. For an exact identification
of herbarium specimens of this genus, it is essential that material be
collected not only in full flower but also with fruits. The study of the
pappus (a good taxonomic character for distinguishing species of the
genus Tripleurospermum) on ovaries of still young florets does not
give a correct idea, because the pappus changes as the achene matures.
The shape of the receptacle, a character no less important in the genus,
is difficult to determine without damaging the herbarium specimen.
Having strong superficial similarity, the species of this genus often
appear, in the nature of the receptacle, florets, and achenes, not only
different but even as members of different genera. Only a careful study
of the species in nature based on a large quantity of material in different
developmental stages will enable taxonomists to form a correct concept
of their number and geographic distribution. Since I could study only
a few Tripleurospermum species in nature, I believe that even my
study is not without potential error.

1. Plants perennial with slender woody rhizome ..............::seseeeeeeee 2
+? Piasanaualior biennial ........:00....5000.: Me GL. 12.
2... Reine with. single capitulum ........2ig2m in@er Seh........ 3:
+ Stem branched, with few capitula (less often simple). ................ 8.
3. _Involucral bracts with dark brown and black membranous margin

Ce 252005) A sont snnennnn ee a EAD Bee zee IEEE wepicnn tn 4.
+ Involucral bracts with narrow, y, brown, membranous margin or green,

with light colored membranous Marin ...........::cesceeeseeeseeeeseeeeeeees 6.

4. Capitula small, 15-20 mm in dia; involucral bracts with narrow,
brown membranous margin; pappus long, a half as long as achene
or slightly longer, dark brown all over or only at apex. Plants small,
3. S=1 7 remy Me etic i...2l5 3. T. grossheimii (An. Fed.) Pobed.

+ Capitula large, 20-45 mm in dia; involucral bracts with broad,
blackish-brown, membranous margin; pappus shorter, a third to a
half as long as achene, with dark brown margin of lobes. Plants up
I POI cP EN 3b 38 50 LL oc ceccensins See nate e RDM t ane cemowsvasessanses 5.

5. Capitula with large broad involucre, 15-20 mm in dia; involucral
bracts black-bordered; outer florets white, 25—35 in,capitulum, with
short and broad, 13—15 mm long and 5-6 mm wide ligule ...................
ole MEER Melb Sai!) ces bh. SUT, ede ag 2. T. subnivale Pobed.

+ Capitula with smaller and narrower involucre, 10-15 mm in dia;
involucral bracts brown-bordered, outer florets white, 20(30—35)38
in capitulum, with narrower, 9-15 mm long and 2-5 mm wide,
Retest). .siihnelg..Jisou eae: 1. T. caucasicum (Willd.) Hayek.

160

154

10.

11.

Involucral bracts with narrow, brown, membranous margin; leaves
covering lower half of stem, hard, somewhat thick, twice or thrice-
pinnately cut into narrow, linear somewhat long secondary lobes
Lapa tall be ocVebiSdabl LAUER AUS gSdSe. MREC DEE ADEE EE AED: PENCE ch
Involucral bracts green, with light-colored margin; leaves only at
base, pinnately cut, with segments palmate or pinnately divided
into lanceolate, short and broad secondary ODES ................ccsseeeeeeeeees
spiel £8, Rte eel Aare Se, 13. T. colchicum (Mand.) Pobed.
Stem 15-30 cm high; capitula large, 2.5—-4 cm in dia, leaf lobes
oblong; achenes with single thin rib on dorsal side. Plants of alpine
meadows xs. 542... 210!) BLO es Ay 4. T. szovitzii (DC.) Pobed.
Stem 15 cm high; capitula 2—3 cm in dia; leaf lobes reduced; achenes
dorsally not ribbed. Plants of stony slopes in lower mountain zone
Serer r ete em Mane EI Men Peau t ere REM Myler L 14. T. tchichatchevii Boiss.

Involucral bracts with dark brown MarQiN ............ccccccesscceeseeeenees 9.
Involucral bracts green or with scarious margin, sometimes with
VELY, NALrOW ;DrOWM MALGIN ... seven scceaebteetd ike. d Casedee wel. asek cbee Ak ki

Receptacle conical; involucral bracts with narrow dark brown margin
veuestedesiasses aise MORO aU. L IE 7. T. ambiguum (Ldb.) Fr. and Sav.
Receptacle hemispherical; involucral bracts with more or less wide,
dark Drown, Dar OIM,.......Sfeceeseitee ML tEEES, ERUoRS eC EM, PRES fe segs 10.
Leaves narrow, divided in long and narrow lobes; peduncles long;
involucral bracts narrow; ligules of peripheral florets 12-15 mm
long, 2-3 mm wide ....... 5. T. transcaucasicum (Manden.) Pobed.
Leaves broader, divided in short and broad lobes; peduncles shorter;
involucral bracts broader; ligules of peripheral florets broader, 12—
LSiimmelong,.s=9ianm ‘widens. rca. 2M 01 .. eat. eae eats cence
1S, BRON eee awe, ann, as 6. T. karjaginii (Mand. and Sof.) Pobed.
Leaves ovate-oblong, broad, 80-90 mm long, 50-90 mm wide,
pinnately cut or twice-pinnate; achenes smooth; pappus white, a
third as long as achene ... 12. T. rupestre (Somm. and Lev.) Pobed.
Leaves oblong, thrice-pinnate; achenes rugose, pappus white, longer
on ventral side, a third as long as aChene ..........s.sssssssssseesessseeseseseeeeeeees
Secsicntisi tases mob Nact ALAN Gaeta 11. T. elongatum (F. and M.) Bornm.

. Achenes with long, white, two-lobed pappus; middle rib columnar,

covering almost entire ventral side. Plants 10-30 cm high ................
wonton sStAGhOULs SUMS, SLC, iNeed Estee 9) 22. T. parviflorum (Willd.) Pobed.
Achenes lacking pappus, or pappus less than a fourth as long as

achene, 2f2:2.,Aeelon Ds. PaNCerMatT, Auak PL pees, Ea, eee ceca 13.
Achenes light colored, smooth, with 3 very thin, white ribs ...... 14.
Acheénés darkibrownyrupose’ ...veiurain. ew, . pea a... jo

Achenes without pappus, with 2 small glandular hairs on dorsal
side; margin of outer involucral bracts broadly white-membranous;
capitula usually lacking peripheral florets (if with peripheral florets,

then ligules very short, usually exserted from involucre) ..............0.

MR PRY, SOROR R R A 19. T. disciforme (C.A.M.) Sch. Bip.

+ Achenes with short, white, membranous, lobed pappus, with 2 large
glandular hairs on dorsal side; margin of outer involucral bracts
narrow and pale-brown, membranous; capitulum with large
peripheral florets (two times as long as involucre) ...............04

2 ale pe mene Aahisie dle a4 A 21. T. sevanense (Mand.) Pobed.

15" Pappus white; membranous; 1ODed tn. MRI). acess
BOAT EL ASN, SOR 20. T. decipiens (Fisch. and Mey.) Bornm.

+ Pappus as small, cartilaginous, toothed border. ..............:ceseee 16.
16. Involucral bracts with more or less wide, brown or blackish-brown
BOPAO NEE, SALE e Sco Seitcensssovges eeecnach tet onehotatat eelieeeenat tes tmevetactacanores exe ry.

+ Involucral bracts with whitish, light brown or brown, but then very
PPATTOW TRIALS YASS eT iresecser seen eetc csc Ptb an tcteteetan cece 18.

17. | Green part of involucral bracts deltoid; leaves bipinnate, oblong;
stem erect, sparingly branched, sometimes with more branches .......

Cea eed ARES FERC A 9. T. phaeocephalum (Rupr.) Pobed.

+ Green part of involucral bracts oblong or narrowed in middle, border
narrower than green part, blackish-brown in lower part, light brown

at apex; leaves bipinnate, obovate; stem strongly branched .....

Bi Mkt ON ALEC Be aia eer 10. T. subpolare Pobed.

18. Involucral bracts with very narrow, brown marin ...............006 19.
+ Involucral bracts with white, lustrous or light brown border apically
BPI TIS, i 08 NINN otk An LU oes hess Pisdetedecvedehas sooth 20.

19. Receptacle hemispherical; leaves succulent, thrice pinnately cut into
short thick pinnules. Plants of the Baltic Coast, 10-40 cm high ...
WERE IERIE Gt Ma 16. T. maritimum (L.) Koch

+ Receptacle conical; leaves twice or thrice pinnately cut into narrow
filiform pinnules. Plants of the Pacific Coast, 10-50 cm high ........

Ee OE Rt 8. T. tetragonospermum (F. Schm.) Pobed.

20. = Involucral bracts with white lustrous margin ..............csseseeees 21.
+ Involucral bracts with light brown, apically broadened border, or
with narrow white margin; outer bracts often pale green, lacking
membranous Mafrin ..............08 15. T. inodorum (L.) Sch. Bip.

21. Capitula 10-15 mm in dia; stem 8-30 cm high, branched from base
EAN ON I RL RTI RR 17. T. limosum (Maxim.) Pobed.

+ Capitula 18 mm in dia; stem 40 cm high, branched only in upper
HAP ER RO ERROR 18. T. breviradiatum (Ldb.) Pobed.

161 Subgenus 1. Chionogeton Pobed. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XXI (1960) 344.—Achenes with more or less long
pappus, a third to a half as long as achene, usually three-lobed; capitula
solitary, very large, less often few; involucral bracts often with more
or less wide, black or white, membranous margin; stem usually simple,

162

156

less often with few short branches. Perennial herbs with woody, slen-
der, branched rhizome, usually growing in alpine zone in high-moun-
tain moist meadows and near patches with snow.

Note. Besides our species, T. melanophyllum (Babel.) Pobed. from
northern Iran also belongs to this subgenus.

Type of subgenus: Tripleurospermum caucasicum (Willd.) Hayek.

Series 1. Caucasica Pobed. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1960) 344.—Stem simple; margin of involucral bracts
wide, membranous, blackish or dark brown; pappus long, almost a half
as long as achene, entirely brown or with brown lobes.

1. T. caucasicum (Willd.) Hayek in Denkschr. Akad. Wiss. Wien,
Math.-Nath. Kl. XCIX (1924) 204.—Triplerospermum caucasicum
Owerin in Zap. Kavk. Obshch. Sel’sk. Khoz. God 20, Nos. 4, 5, 6
(1874) 138, 141 and 150.—Pyrethrum caucasicum Willd. Sp. pl. Il
(1800) 2156; MB. FI. taur.-cauc. II, 322, III, 577; DC. Prodr. VI, 56;
C.A.M. Verz. 75; Ldb. Fl. Ross. II, 548.—Anthemis caucasica Adam,
in Weber and Mohr Beitr. I (1805) 71.—Chrysanthemum alpinum MB.
Casp. (1800) 211, nom. nud.; Georgi Beschr. d. Russ. R. HI, 5, 1250.—
C. caucasicum Pers. Syn. II (1807) 462; Wehrh. Gartenstauden, II,
1101.—Tanacetum caucasicum Sch. Bip. Ueb. die Tanac. (1844) 60.—
Chamaemelum caucasicum Boiss. Diagn. Ser. I, XI (1849) 20; Boiss.
Fl. or. III, 331 p. p.; Grossh. Fl. Kavk. IV, 128.—Ic.: Meissner in
Gartenwelt, XXXIV (1930) 385.—Exs.: Woron. Pl. abch. exs. Nos.
583, 598; GRF No. 3246.

Perennial. Rhizome woody, dark brown, usually branched near
tip; stems few, 20-40 cm high, erect, simple, less often branched,
glabrous, pubescent only below capitula, sulcate throughout, usually
leafy in lower half, leafless in upper third. Leaves oblong, bipinnate
with narrow, oblong, linear, acute or subobtuse segments, glabrous;
basal leaves and those on vegetative branches 30(40-50)70 mm long,
10(15)—20 mm wide, petiolate, (petioles 15-30 mm long); cauline leaves
10(20)—25 mm long, 6-10 mm wide, sessile, with rachis broadened at
base, bearing short linear segments. Capitula solitary, 30-45 mm in
dia with peripheral florets. Receptacle hemispherical. Involucre 10-15
mm in dia; bracts oblong, obtuse, broadly brown membranous along
margin, glabrous. Peripheral florets 20(30—35)38 in capitulum; ligules
white, 9-15 mm long, 2-5 mm wide, obtuse, with 3 shallow teeth;
tubular florets yellow, with short infundibuliform tube and deltoid,
acute or broadly deltoid, subobtuse lobes, sometimes apically dark
brown. Stigma deeply divided. Anther appendages conical with uncate
apex. Achenes oblong-linear, smooth, 2-3 mm long, 0.75—1.0 mm wide,

165

157

with 3 thick white ribs, reddish glands on outside, inconspicuous;
pappus a third to half as long as achene, deeply three-lobed ventrally,
with long deeply toothed border on outside equaling lobes; pappus
brown entirely or only at apex. Flowering June to September.

On moraines, along borders of glaciers, near patches of snow in
alpine zone.—Caucasus: Ciscaucasia, Eastern, Western and Southern
Transcaucasia, Dagestan. General distribution: Armenia-Kurdistan, Balkans-
Asia Minor (Bulgaria)? Described from Caucasus. Type in London?

Note. One specimen of this species with the label “Caucasus,
Mussin-Puschkin,” apparently from the authentic material, is preserved
in the herbarium of the Botanical Institute, Academy of Sciences of
the USSR. In describing this species, Willdenow included “Chrysan-
themum caucasicum com. Mussin-Puschkin” among its synonyms.
Unfortunately, this label is incomplete, without mention of the date of
collection and locality. In the literature, this species has been reported
repeatedly for Bulgaria [Velenowsky, Fl. Bulg. (1891) 272; Lipsky,
Fl. Kavk. (1899) 348; Hayek op. cit. 204], which is extremely doubt-
ful. I did not examine the material from Bulgaria. Probably, there are
some close species morphologically resembling it in Bulgaria.

2. T. subnivale Pobed. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XX (1960) 490.—Chamaemelum caucasicum var. pumilum
Boiss. Fl. or. III (1875) 331.—C. melanolepis An. Fedor. in Bot. Mat.
Gerb. Bot. Inst. Akad. Nauk SSSR, XI (1949) 165, non Boiss.—C.
caucasicum auct. fl. Cauc. non Willd.; Grossh. Fl. Kavk. IV, 128.

Perennial. Rhizome slender, woody, dark brown, usually branched,
bearing nonflowering and 1-2 or few flowering shoots; stems 10-25
cm high, ascending, simple usually glabrous in middle, covered with
scattered, short appressed hairs below and in upper part, particularly
dense below capitulum, leafy up to apex, less often leafless in upper
third. Leaves oblong, 15(25-40)35 mm long, 4(10-11)15 mm wide,
pinnately cut into short thick segments, in turn thrice palmately cut,
less often pinnate, cuspidate, sparsely pubescent, less often glabrous;
basal leaves and those on nonflowering branches petiolate; cauline
leaves smaller and sessile, broader at base, with simple linear segments.
Capitula solitary, large, 30-45 mm in dia including ligulate ray florets;
receptacle hemispherical. Involucre 15—20 mm in dia, hemispherical;
involucral bracts oblong, obtuse, with wide (broader than middle green
part of bract, especially at apex) blackish-brown membranous erose
border. Peripheral ligulate florets 25-35; ligules white, 10(12—14)20
mm long, 3.0(4.0-5.0)5.5 mm wide, slightly narrowed at apex, obtuse,
with 3 shallow teeth; tubular florets golden-yellow, with tube gradually
broadening toward apex and deltoid or broadly deltoid, subobtuse

166

158

recurved lobes at full bloom, sometimes very dark brown at apex.
Stigma deeply divided. Anthers with oblong-conical apical appendage.
Achenes 2.0—2.5 mm long, 1.0—1.5 mm wide, compressed, with 3 thick
whitish ribs on ventral side, light brown, smooth (immature), outside
with 2 small, red, resin glands at apex; pappus membranous, 1.5-2.0
mm long, more than a half as long as achene (immature), three-lobed,
with 3 equal lobes ventrally and flat, dark brown lobe of same length
on dorsal side. Flowering July to September. (Plate V, Fig. 1.)

In alpine meadows, to 2,500-3,500 m.—Caucasus: Ciscaucasia,
Dagestan, Eastern and Southern Transcaucasia. Described from
Azerbaidzhan (Shakh-Dag Mountains). Type in Leningrad.

Note. Tripleurospermum melanophyllum (Nabel.) Pobed., described
by Nabelek [Publicat. de la Facul. des Sci. de Univ. Masaryk (1925)17]
from Kurdistan, is the closest species, which differs from the Caucasian
species by the conical receptacle, smaller capitula, shape of the pappus
and other minor characters. The ranges of both species are separate,
which even more strongly confirms their independent status.

3. T. grossheimii (An. Fed.) Pobed. comb. nova.—Chamaemelum
grossheimii An. Fed. in Bot. Zhurn., 34, 3 (1949) 290.—Ic.: An. Fed.
ibid.

Perennial. Rhizome woody, branched, brown; stems simple, erect
or weakly ascending, 3.6-17 cm high, slender, longitudinally striate,
glabrous or sparsely pubescent, leafy only in lower half. Leaves
gradually reduced toward stem apex, oblong, pinnately cut, with linear-
lanceolate segments, acute, 2-, 3- or 5-fid, glabrous or sparsely
pubescent, especially at base; basal leaves 20-35 mm long, 6-10 mm
wide, long-petiolate; cauline leaves with 2-3 pairs of linear segments,
sessile, broader at base, amplexicaul, with linear simple segments.
Capitula 15-20 mm in dia, including ligulate florets; receptacle
hemispherical. Outer involucral bracts narrow, triangular, middle and
inner oblong, with blackish-brown, narrow membranous border. Ligules
of peripheral ray florets oblong, broad, 8-10 mm long. 3.0-3.5 mm
wide, with 3 obtuse, more or less large apical teeth, often recurved at
flowering; disk florets yellow, with tube slightly swollen in upper part.
Anthers with oblong-conical apical appendages. Achenes 2.5-3.0 mm
long, 0.5—1.0 mm wide, with 3 thin ribs on ventral side, sometimes
fourth very fine rib on outside, dorsally smooth, eglandular above or
with 2 very small red glandular hairs; pappus large, a half as long as
achene or slightly longer, three-lobed ventrally, middle lobe free almost
from base, dorsally compressed, fimbriate, brown at apex. Flowering
August.

167

159

In alpine meadows.—Caucasus: Possibly, growing in Southern
Transcaucasia. Described from Great Ararat Mountain. Type in
Leningrad.

4. T. szowitzii (DC.) Pobed. comb. nova.—Pyrethrum caucasicum
B. szowitzii DC. Prodr. VI (1837) 56; Ldb. Fl. Ross. II, 548.—
Chamaemelum szowitsii Boiss. FI. or. III (1875) 332; Grossh. Fl. Kavk.
PVy 128;

Perennial. Rhizome slender, woody, very dark brown, bearing few
nonflowering and 1-2 flowering shoots; stems 15-30 cm high, erect,
simple, unicapitulate, glabrous, leafy in lower half, upper half leafless
or with strongly reduced leaves, more or less densely pubescent below
capitulum. Leaves hard, thick; lower cauline leaves and those on
nonflowering shoots oblong, 30-50 mm long, 15-20 mm wide, twice
or thrice pinnately cut into narrow linear, more or less long cuspidate
lobes; cauline leaves gradually reduced, all leaves sessile. Capitula
large, 25-40 mm in dia including ligulate florets; receptacle
hemispherical. Outer involucral bracts narrow triangular, weakly
pubescent, especially along margin, inner obovate, oblong, obtuse,
glabrous; all bracts green, narrowly brown-membranous along margin,
with wide, brownish discontinuous border at apex. Peripheral ligulate
florets 17-15 mm long, 5-7 mm wide, tubular, with gradually broadened
tube. Anthers with oblong-triangular appendages. Achenes with long
white pappus a third as long as achene, three-lobed ventrally, lobes
connate to half, with short toothed border on dorsal side. Flowering
May to July. (Plate V, Fig. 2.)

Humid meadows in alpine zone.—Caucasus: Eastern, Western, and
Southern Transcaucasia. Described from Transcaucasia (Guriel). Type
in Leningrad.

Note. In the original description of this species, the size of the
pappus given is in error. De Candolle reported the pappus as short,
which was repeated also by Boissier (FI. or. III, 332), whereas, judg-
ing from the type material, the pappus of T. szovitzii is large, a third
as long as the achene.

Series 2. Transcaucasica Pobed. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XXI (1961) 345.—Stem with the branches shorter
than or equal to it.

5. T. transcaucasicum (Manden.) Pobed. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XXI (1961) 346.—Chamaemelum
transcaucasicum Manden. in Zap. po Sist. i Geogr. Rast. Tibl. Bot.
Inst. XXI (1959) 64.

160

163

MZ
7

,
Mf

Plate VI.
1 — Tripleurospermum limosum (Maxim.) Pobed. habit of plant, involucral
bracts, receptacle, peripheral pistillate floret, bisexual disk floret, achene in
dorsal and ventral view; 2— T. disciforme (C.A.M.) Sch. Bip., habit of the
plant, involucral bracts, bisexual disk floret, achene in dorsal and ventral view;
3 — Cotula coronopifolia L., habit of plant, involucral bracts, receptacle,
peripheral pistillate floret, bisexual disk floret, achenes of pistillate and bisexual

disk florets.

168

161

Perennial. Rhizome woody, slender; stems 20-60 cm high, few,
branched in upper half, branches slightly exceeding main stem, sparsely
covered with long white hairs, more dense at nodes and below
inflorescence, leafy throughout. Leaves on sterile shoots and basal
leaves obovate-oblong, 30-85 mm long, 4-15 mm wide, twice pinnately
cut into long linear cuspidate pinnules, with more or less long petioles,
broader at base and with short pinnate secondary lobes, sparsely
pubescent at base; cauline leaves distant, gradually reduced upward,
sessile. Capitula 20-30 mm in dia including peripheral florets, on
leafless peduncles, 20-50 mm long, ligules narrow-lanceolate, 12-15
mm long, 2-3 mm wide; receptacle hemispherical. Outer involucral
bracts lanceolate, scatteredly pubescent, subobtuse; middle bracts
oblong, obtuse, with narrow, brown, hairy border; inner bracts obovate,
oblong, membranous at apex, with very dark brown border. Achenes
2 mm long, obpyramidal, very dark brown, rugulose and with small
tubercles, glandular-hairy at apex on dorsal side, with narrow, acute,
whitish ribs, middle rib on ventral side with deep depressions along
sides; pappus white, short, a fourth as long as achene. Flowering June
to August.

In high-mountain, wet meadows.—Caucasus: Eastern and Southern
Transcaucasia. Described from Akhalkalaka. Type in Tbilisi.

Note. The specimens of this species collected from Tskhara-
Tskharo Mountain by A.L. Pastukhov in 1911 and from Goderdz Pass
by D.I. Litwinow in 1913 are noteworthy for the heavy pubescence of
the leaves, stems, and outer involucral bracts.

6. T. karjaginii (Mand. and Sof.) Pobed. comb. nova.—
Chamaemelum karjaginii Mand. and Sof. in Dokl. Akad. Nauk Azerb.
SSR, XV, 3 (1959) 249.—C. caucasicum auct. cauc. non Willd.—C.
kotschyi auct. cauc. non Boiss.

Perennial. Rhizome slender, branched, woody, brown; stems 1-3,
15-30 cm high, erect, with few short branches, not exceeding main
stem, only in upper half, glabrous, pubescent below capitula, leafy up
to apex. Lower leaves and those on nonflowering shoots ovate-oblong,
35-40 mm long, 8-18 mm wide, twice-pinnately cut into thin linear
secondary lobes, simple pinnate at base and on petiole, with short
segments, cauline leaves oblong, less divided gradually reduced upward;
all leaves glabrous. Capitula solitary at apices of stem and branches,
3.0-3.5 cm in dia; receptacle hemispherical. Outer involucral bracts
oblong-deltoid, middle oblong, inner obovate; all bracts with wide,
almost blackish-membranous border. Outer ligulate florets 20, 12-15
mm long, 3-5 mm wide. Achenes unknown; pappus in tubular florets
three-lobed, brown, like that of 7. caucasicum. Flowering August.

169

162

On mountain slopes.—Caucasus: Southern Transcaucasia. De-
scribed from Armenia (Daralagyoz). Type in Tbilisi; isotype in
Leningrad.

Note. A doubtful species, as the characters mentioned by the au-
thors to distinguish it from T. transcaucasicum usually vary: broader
leaves, shorter peduncles, broader and rich brown-bordered involucral
bracts, broader ligules of ray florets, etc. There is an isotype preserved
in the herbarium of the Botanical Institute, Academy of Sciences of the
USSR, from which it is difficult to judge the stability of these characters.

Subgenus 2. Tripleurospermum.—Leucoglossa DC. sect. generis
Pyrethri DC. Prodr. VI (1837) 53, p. max. p.—Cynoclines DC. sect.
gen. Pyrethri DC. ibid. 58, p. min. p.—Achenes with short coronate
pappus, less often pappus long, two-lobed. Capitula numerous;
involucral bracts with more or less narrow light brown or wide white,
lustrous, less often with blackish-brown margin. Stem usually branched.
Annual herbs, rarely perennial, widely distributed along seacoasts,
plains, often weeds, ruderals, rarely in mountains.

Type of subgenus: T. inodorum (L.) Sch. Bip.

Section 1. Phaeocephala Pobed. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XXI (1960) 346.—Leucoglossa sect. gen. Pyrethri
DC. Prodr. VI (1837) 58, pro min. p. Achenes with short, coronate,
entire or toothed pappus; involucral bracts with wide, membranous,
very dark brown or blackish margin. Annual, less often biennial or
perennial herbs.

7. T. ambiguum (Ldb.) Fr. and Sav. Enum. I (1875) 236 (tantum
nomen, excl. pl. extrem.).—Pyrethrum ambiguum Ldb. FI. Alt. IV
(1833) 118; DC. Prodr. VI, 54; Ldb. Fl. Ross. II, 547.—Matricaria
ambigua Kryl. Fl. Alt. III (1904) 625; O. and B. Fedtsch. Perech.
Rast. Turk. IV, 185; Schischk. in Kryl. Fl. Zap. Sib. XI, 2737.—
Chamaemelum ambiguum (Ldb.) Boiss. Diagn. Ser. I, XI (1849) 20.—
Exs.: Pl. alt. exs. No. 87.

Perennial. Rhizome creeping, branched near apex, with few
floriferous shoots and nonflowering branches; stems erect, 15-60 cm
high, simple, sometimes weakly or more or less strongly branched near
apex (Soviet Central Asian plants), sulcate, glabrous, sparsely pubescent
above, especially below capitula, less often with long, curly, intertwined
hairs. Leaves oblong-oblanceolate, spatulate, or oblong, (3.0)3.5—
4.0(10.0) cm long, (0.5)0.8—1.2(2.5) cm wide, twice pinnately cut into
narrow filiform lobes, terminating in short cusp; basal leaves petiolate,
cauline sessile, slightly broadened at base, gradually reduced upward.

170

163

Capitula solitary at apices of main stem and branches, less often in the
fork of main stem, 3-5 cm in dia with peripheral ligulate florets;
receptacle conical. Involucre flat-hemispherical; involucral bracts 2-
3-seriate, outer lanceolate, subacute, inner oblong; all bracts with
narrow, dark brown, undulate membranous border, broadened at apex,
glabrous or sparsely hairy. Peripheral ligulate florets with horizontally
divergent ligules, 8-9 mm long, 3-4 mm wide; disk florets golden
yellow, often with red thickening at tips of lobes. Achenes 2 mm long,
very dark brown, rugose-tuberculate, with 3 thick whitish ribs,
sometimes also with 1-2 fine ribs dorsally, with 2 small, red, glandular
hairs above on dorsal side; pappus membranous, slightly longer than
in T. tetragonospermum, broadly crenate. Flowering April to September.
(Plate VII, Fig. 1.)

In alpine and subalpine meadows, alpine tundra near patches of
snow, high-mountain meadows and deciduous forests, at 2,000—3,500
m.—Western Siberia: Altai; Soviet Central Asia: Dzhungaria-
Tarbagatai, Tien Shan (only in northern part). General distribution:
Iran (northern part), Dzhungaria-Kashgaria, Mongolia. Described from
Altai. Type in Leningrad.

Note. It has been reported that the plant collected in the Tien Shan
in Mukhurdai Pass by Roborovsky had pink ligules of the ray florets;
in other respects the plant is indistinguishable from T. ambiguum.

This high-mountain alpine species is still being confused with the
Far Eastern coastal species T. tetragonospermum (F. Schun.) Pobed.,
from which it is well distinguished by a woody rhizome (and not an
annual root), longer and narrower leaves, less leafy stems, a broader,
dark brown (not narrow brownish) border of the involucral bracts,
involucral bracts always with scattered pubescence along the margin
(and not glabrous), unbranched or weakly branched (especially in alpine
plants) stems, short-pubescent peduncles, especially on thickened part
below the inflorescence where sometimes dense, curly, white hairs are
found, a somewhat longer and bluntly broad-toothed pappus (and not
entire). We have not included the Dauria Region in the distribution of
this species, although a specimen of T. ambiguum from Baikal, collected
by Maak as No. 20 and sent in 1858, is preserved in the herbarium of
the Botanical Institute, Academy of Sciences of the USSR, which raises
no doubt about the correctness of the identification. However, to-date
this species has not been recollected from Lake Baikal. It must be
noted that this species is also not found in the Sayans. The plants
collected by B.K. Schischkin from the Kegen District of Kazakhstan in
1935 and earlier, in 1930, by P.A. Smirnov from the Narym Range
(village of Katon-Karagai), are distinguished by elongated leaves with
short broad segments, pubescence throughout or only along the rachis,

17

—

164

large capitula, 4-5 cm in dia including ray florets, a broader membra-
nous border of the involucral bracts, and a longer, membranous pap-
pus.

8. T. tetragonospermum (F. Schm.) Pobed. in Bot. Mat. Gerb.
Bot. Inst. Akad. Nauk SSSR, XXI (1961) 346.—Chamaemelum
tetragonospermum F. Schm. Fl. Sach. (1868) 148.—Matricaria
ambigua auct. non Ldb. (cum Pyrethrum ambiguum): Miyaba FI. Kuril.
(1890) 242; Kom. in Tr. SPb. Bot. Sada, XXV (1907) 640; Kom. and
Alis. Opred. Rast. Dal’nevost. Kr. II, 1028.—Tripleurospermum
ambiguum Fr. and Sav. Enum. Pl. Jap. I (1875) 236.—Matricaria
tetragonosperma (Fr. Schm.) Hara and Kitam. in Mem. Coll. Sci. Kyoto
Imp. Univ. ser. B, 15 (Comp. Jap., 2) (1940) 336.— Tripleurospermum
inodorum var. ambiguum Reich. ex Herder.—Ic.: Iinuma Somoku
Dzusetsu, Iconograf. Pl. Nippon, XVII (1912) t. 20; Miyoshi Atlas
Jap. Veget. t. 65.

Biennial or annual. Root vertical, curved, branched. Stems 7-50
cm high, more or less erect, thick, sulcate, glabrous, branched, especially
above; branches long, ascending, densely leafy. Leaves oblanceolate-
oblong or oblong, 7-11 cm long, 1.5—3 cm wide, twice or thrice
pinnately cut into narrow filiform lobes, apical lobe 3-10 mm long,
0.3-0.6 mm wide, with short cusp, glabrous or sparsely pubescent;
lower leaves sessile, middle cauline leaves short-petiolate; all leaves
broadly amplexicaul; peduncles sulcate, thickened below capitula, often
sparsely pubescent. Capitula few, at apices of main stem and branches,
3.5-4.0 cm in dia. Involucre flat-hemispherical; involucral bracts usually
4-seriate, imbricate; outer bracts ovate-oblong, obtuse, sometimes
basally with somewhat long teeth, middle narrow-oblong, obtuse, inner
lanceolate-oblong, obtuse; all bracts membranous, with prominent
darker midrib on dorsal side, narrow light brown along margin.
Receptacle spherical-conical, about 9 mm long. Peripheral ligulate
florets 1-seriate, white, horizontal; ligules 15-18 mm long, 3.0—4.5
mm wide; disk florets golden yellow, with tube abruptly broadened at
apex. Anthers basally obtuse, filaments thickened at apex. Achenes 2—
3 mm long, with 3 or 4 ribs, rugose-tuberculate, with 2 roundish
oleiferous glandular hairs above on convex dorsal side; pappus
membranous, short, more or less entire. Flowering June to October.

Along seacoasts.—Far East: Kamchatka (rarely), Anadyr (very
rarely), Ussuri, Sakhalin, Kuril Islands. General distribution: Japan,
China. Described from Sakhalin. Type in Leningrad.

Note. This species is found mainly on the islands of the Pacific
Ocean; on the mainland, it usually is in the Pacific Coastal Region.
Three plants collected from Kamchatka and one from the Penzha Inlet

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apparently were introduced there by chance, since this species is com-
pletely absent from the Okhotsk coast of the continent.

9. T. phaeocephalum (Rupr.) Pobed. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk, XXI (1961) 347.—Chrysanthemum grandiflorum Hook.
in Parry’s Sec. Voy. (1825) 398, non Brouss. (1804).—Pyrethrum
inodorum var. nanum Hook. FI. Bor. Am. 1 (1833) 320.—Matricaria
inodora var. nana Ldb. FI. Ross. II, 2 (1844) 547.—M. inodora 9.
monocephalum Sch. Bip. Ueb. die Tanac. (1844) 30.—M. inodora f.
phaeocephala Rupr. Fl. Samojed. (1845) 42.—M. grandiflora Brit. in
Mem. of the Torrey Bot. Club. 5 (1893-1894) 340; Tolm. in Tr.
Polyarn. Kom., 13, 43; Perfil’ev, Flora Severnogo Kraya, 3, 352.—M.
phaeocephala (Rupr.) Stefansson FI. Isl. ed. 2 (1924) 223.

Perennial or biennial. Root short, thick with numerous fibrous
rootlets. Stems 1-few, 2—3 from root, up to 30 cm high, erect, simple
or weakly branched near apex, sulcate, glabrous. Leaves sparsely
covering stem, oblong, (3.0)3.5—4.0(9.0) cm long, (0.8)1.0—2.0(3.0)
cm wide, usually glabrous, simple pinnate with 2—3-lobed flat lobes,
0.2-1.5 mm wide, gradually narrowed toward apex, rachis flat, 1.5-
2.0 mm wide. Peduncle thickened below capitula, glabrous; capitula
solitary at stem apex or, few in case of branching, large, 3-6 cm in
dia. Involucre hemispherical, 2—3-seriate; involucral bracts with
triangular green part and wide border, broader than green part of bract,
especially at apex, blackish-brown, undulate, entire or discontinuous.
Receptacle hemispherical, slightly elongated. Ligulate florets 12—20
mm long, 3-5 mm wide, white, 3-toothed; disk florets with wide short
tube, slightly broader at apex, golden-yellow, opened florets sometimes
with anther tube exserted from florets. Achenes 1-2-3 mm long, with
more or less white, thick, obtuse ribs, light brown, sometimes almost
white, weakly rugose outside, with 2 fairly large, red, oleiferous
glandular hairs; pappus white, membranous, 0.5 mm long, 4—5-lobed,
usually horizontally spreading. Flowering June to August.

Along Arctic coasts, sometimes as weed.—Arctic: Arctic Europe,
Novaya Zemlya, Arctic Siberia, Chukotka, Anadyr; European part:
Karelia-Lapland (only northern part). General distribution: Arctic North
America. Described from the Kolguev Island. Type in Leningrad.

Note. In European specimens of this species a highly prominent
anther tube can be seen in some tubular florets during full bloom in
almost all capitula; in American specimens, the anthers are hardly
visible or only a third exserted; the pappus in American plants is
slightly shorter than in European plants. Siberian specimens of this
species differ from American and European specimens by having brown
(and not almost black) involucral bracts with a lighter-colored margin.

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166

These characters are hardly of taxonomic importance, as they are
extremely variable.

10. T. subpolare Pobed. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XXI (1961) 347.—Matricaria ambigua auct. non Ldb. (cum
Pyrethro ambiguo).—M. grandiflora auct. non Britt.: Perf. Fl. Sev.
Kraya 3 (1936) 352.—Ic.: Bot. Mat., XXI (1961) 348.

Annual or biennial. Root short, branched. Stems few, 25-50 cm
high, erect or ascending, branched only in upper half, weakly sulcate,
glabrous, usually reddish-violet, sometimes basally dark violet. Leaves
dense, obovate, (4.0)4.5—7.0(10.0) cm long, (1.0)1.5-—2.5(3.5) cm wide,
glabrous, twice or thrice pinnate, particularly strongly divided in upper
part into narrow filiform lobes; rachis slightly thickened and cristate at
base. Peduncle 1.5-6.0 cm long, thickened below capitula, glabrous.
Capitula more or less in corymbs, at apices of main stem and branches,
sometimes horizontal, 3-5 cm in dia. Involucre hemispherical, 2-3-
seriate; outer involucral bracts with dark brown, almost black border
below, narrower than green part of bract, with very wide, brown or
light brown, entire or discontinuous border at apex; green part of bracts
oblong, with almost parallel margin or sometimes narrowed more or
less in middle. Receptacle hemispherical. Ligulate florets 10-18 mm
long, 3-5 mm wide, white, 3—4-toothed; disk florets bright yellow,
with broad tube, gradually broadening upward, lobes usually with
longitudinal reddish veins at tips. Anther tube exserted; anthers roundish
at base, apically obtuse, with short pointed appendages. Achenes 2-3
mm long, with light colored or brown obtuse ribs, rugose outside, with
2 small roundish, red, oleiferous glandular hairs; pappus white,
membranous, 4—5-lobed, more than 0.25 mm long.

Along seacoasts and ruderal places.—European part: Karelia-
Lapland. Endemic. Described from the Solovetsky Islands (B. Zayatsky).
Type in Leningrad.

Section 2. Chlorocephala Pobed. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XXXI (1961) 349.—Sect. Leucoglossa DC. Prodr.
VI (1837) 53, p. min. p.; Ldb. Fl. Ross. II (1844) 547, p. min. p.—
Involucral bracts green, with narrow, white or brown membranous
margin; capitula small, 1-3 cm in dia; pappus short, with weakly lobed.
Perenniai herbs, with leaves in lower half of stem.

Type of section: T. elongatum (F. and M.) Bornm.

11. T. elongatum (F. and M.) Bornm. in Fedde, Repert, Beih.
XXXIX (1944) 337.—Pyrethrum elongatum F. and M. ex DC. Prodr.
VI (1837) 56.—Matricaria heterocarpa C. Koch in Linnaea, XXIV

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167

(1851) 333.—Chamaemelum elongatum Boiss. Diagn. ser. I, 11 (1849)
20.—C. heterocarpum Boiss. Fl. or. III (1875) 329.

Perennial. Rhizome slender, woody; stems numerous, 30-40 cm
high, branched, leafless above. Leaves oblong, thrice pinnatisect in
linear, filiform, acute, entire lobes. Capitula solitary at apices of stems
and branches, 1.5—2.0 cm in dia; receptacle hemispherical. Involucral
bracts obovate-oblong, with narrow, white or slightly brownish
membranous border, obtuse, glabrous. Ligulate florets white, 10-15
mm long, 2-3 mm wide, 3-toothed, at apex; tubular florets
infundibuliform, with broad triangular lobes. Anther appendages conical,
small. Stigma deeply divided. Achene 2 mm long, | mm wide, dark
brown, rugose outside, with 2 oleiferous glandular hairs at apex, and
3 thick white ribs on ventral side; pappus white, membranous, longer
on ventral side, 1/3 as long as mature achene. Flowering May. (Plate
V, Fig. 3.)

Stony places.—Caucasus: Eastern and Western Transcaucasia.
Described from Kartalinia (region between Tbilisi and Kutaisi). Type
in Leningrad.

12. T. rupestre (Somm. and Lev.) Pobed. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XXI (1961) 350.—Chamaemelum rupestre
Somm. and Lev. in Nuov. Giorn. Bot. Ital. ser. II, 2 (1895) 86; Grossh.
Opred. 458.

Perennial. Rhizome slender, woody, dark brown, with short, densely
leafy vegetative branches; stem 30 cm high, branched from base,
arcuately ascending, blackish-violet, elongate, with solitary capitula,
glabrous. Leaves of nonflowering branches and basal leaves 8-9 cm
long, 9 cm wide, on 5 cm long petioles, basally toothed, with one
larger tooth; all leaves ovate-oblong, once or twice-pinnately cut into
narrow linear lobes (apical lobes shorter), cuspidate; cauline leaves
much smaller, less divided, sessile. Capitula small, 1 cm in dia, with
white peripheral florets; receptacle hemispherical. Involucre almost
glabrous; outer involucral bracts triangular, brown, inner oblong,
membranous along margin, obtuse, weakly emarginate. Ligules of
peripheral florets elliptical, 5-6 mm long, almost as long as involucre.
Achenes 2.0—2.5 mm long, oblong, compressed, smooth, with 3 thick
ribs on ventral side, dorsally lacking ribs; pappus 1/3 as long as achene.
Flowering July to August.

On rocks.—Caucasus: Western and Southern Transcaucasia.
Described from Lower Adzharia (Shvakhevi). Type in Florence.

13. T. colchicum (Mand.) Pobed. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XXI (1961) 350.—Chamaemelum colchicum Mand.

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168

in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954) 377.—
C. tehihatchevii auct. cauc. non Boiss.: Grossh. Opred. 458.—Ic.: Fl.
Gruzii, VIII, 345.

Perennial. Rhizome slender, branched, woody, dark brown, with
nonflowering and few flowering shoots; stems 20-30 cm high, leafless
or leafy only in upper half, sulcate, sparsely hairy, more or less densely
below capitula. Basal leaves and those on nonflowering shoots ovate,
30-40 cm long, 10-15 mm wide, pinnatisect, with segments palmately
or pinnately divided into lanceolate, short, broad lobes, appressed-
hairy, long-petiolate; cauline leaves small, reduced upward along stem,
oblong, pinnate, with simple linear lobes, sparsely pubescent, sessile.
Capitula solitary on stems, 1.5—2.5 cm in dia; receptacle hemispherical.
Outer involucral bracts narrow-triangular, inner oblong or obovate; all
bracts light green, narrow, light-brown along margin, glabrous.
Peripheral ligulate florets 10-15 mm long, 3-4 mm wide, abruptly
narrowed toward base, with 3 small teeth at apex; tubular disk florets
with wide tube. Achenes 2 mm long, slightly rugose, with broad ribs
on ventral side, and narrow depression inbetween; corona white, trilobed
ventrally, lobes connate up to half, dorsally with more or less short
dentate border. Flowering June to July.

In passes, on rocks along riverbanks.—Caucasus: Western
Transcaucasia. Endemic. Described from the Klukhorsk District
(Nakharsk Pass). Type in Tbilisi.

14. T. tchichatchevii (Boiss.) Bornm. in Fedde, Beih. LXXXIX
(1944) 336.—Chamaemelum tchihatchewii Boiss. in Tchih. As. Min.
2 (1860) 256; Boiss. FI. or. III, 332.—Matricaria tchihatchewii (Boiss.)
Voss. in Vilmorin’s Bumeng. ed. 3, Sib. et Voss. (1896) 504; Hand.-
Mazz. in Ann. Nat. Hofmus. Wien 23, 194.—Pyrethrum tchichatcheffii
Hort. W. Robinson, Engl. Fl. Gard. (1883) t. 213.

Perennial. Rhizome slender, brown, woody; stems few, 15 cm high,
simple, ascending, leafy in lower half, sparsely pubescent, leafless and
glabrous in upper half, tetraquetrous, slender. Leaves ovate-oblong,
20-30 mm long, 10-12 mm wide; lower leaves twice pinnately cut into
short, narrow, cuspidate lobes, petiolate; upper leaves simple, pinnate,
sessile; all leaves canescent. Capitula 2-3 cm in dia; receptacle
hemispherical. Involucral bracts oblong-linear, obtuse, with very narrow,
pale, membranous border, weakly and sparsely pubescent. Ligules of
peripheral florets elliptical-oblong, subobtuse, 3-toothed. Achenes black,
slightly curved, dorsally not punctate, slightly rugose, with coronate
pappus, slightly lobed and white, a half as long as achene. Flowering
May to July.

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On forested seaside hills, dry stony slopes, and open woodlands.—
Possibly, growing in Western Transcaucasia. General distribution:
Balkans-Asia Minor. Described from Samsun (in the Pontic Province
of Asia Minor). Type in Paris?

Section 3. Tripleurospermum.—Sect. Rhytidospermum Sch. Bip.
Ueb. die Tanacet. (1844) 32.—Sect. Chamomilla gen. Matricariae DC.
Prodr. VI (1837) 51, pro min. p.—Achenes granulose-rugose; pappus
short, coronate, entire, finely toothed. Peripheral florets with more or
less long ligules; anther appendages ovoid-triangular. Annual herbs,
with twice or thrice pinnate leaves.

Type of section: T. inodorum (L.) Sch. Bip.

Series Inodora 1. Pobed. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 351.—Involucral bracts with narrow, white
or light brown border.

15. T. inodorum (L.) Sch. Bip. Ueb. die Tanacet. (1844) 32;
Lindemann, Prodr. Fl. Chersonen, 106; C. Koch Syn. 1026.—Matricaria
inodora L. Fl. Suec. ed. II (1755) 765; DC. Prodr. VI, 52; Ldb. FI.
Ross. II, 545; Kryl. Fl. Zap. Sib. XI, 2735.—M. chamomilla L. Sp. pl.
(1753) 891.—Chrysanthemum inodorum L. Sp. pl. (1763) 1253;
Schmalh. Fl. II, 68.—Chamomilla inodora Gilib. Exerc. phytol. I (1792)
179.—Pyrethrum inodorum Moench, Meth. (1794) 597.—P.
parviflorum Willd. Sp. pl. Ill, 3 (1803) 2158.—Rhytidospermum
inodorum Sch. Bip. II, Canar. III, 2 (1836) 277.—Chamomilla inodora
C. Koch in Linnaea, XVII (1843) 45; Ej. Syn. 45.—Chamaemelum
inodorum (L.) Vis. Fl. Dalm. II (1847) 85; Boiss. Fl. or. II, 327;
Grossh. Fl. Kavk. IV, 127.—Ic.: Fedtsch. and Fler. Fl. Evrop. Ross.
Fig. 971; Syreistsch. Ill. Fl. Mosk. Gub. III, 270; Hegi Ill. Fl. VI, 2,
fig. 303 and 304.—Exs.: GRF No. 824; Fl. exs. Aust.-Hung. No. 3789;
Fl. Gall. et Germ. exs. No. 2864; Pl. Finl. exs. No. 978 p. p.

Annual or biennial. Root slender, fusiform. Stem erect or ascending,
30-75 cm high, cylindrical, fistular, with numerous light colored and
green furrows, glabrous, usually branched in upper half. Leaves ovate-
oblong or oblong, thrice pinnately cut into narrow filiform long lobes,
of which apical lobe 4-10 mm long and 0.25—1.0 mm wide, with short
cusp, glabrous, sessile or on usually short petioles thickened at base;
peduncles long, unicapitulate, thickened below capitula, sometimes
weakly pubescent. Capitula few or numerous, at apices of main stem
and branches, 1.5-4.0 cm in dia. Involucral bracts oblong or
oblanceolate, slightly broader at base, more or less narrow-membranous
along margin, white or light brown; outer bracts sometimes lanceolate,

170

pale green, without membranous margin. Receptacle hemispherical or
broadly conical, glabrous, weakly punctate. Peripheral florets ligulate,
white, horizontal, ligules 50-20 mm long; disk florets golden-yellow,
tube gradually broadening in upper half, often (especially in southern
form) lobes with bright red glandular hairs at apex. Achenes 2.0-2.5
mm long, dark-brown, flattened, rugose, with 3 white narrow ribs
ventrally and along sides and with 2 large, roundish, oleiferous, red
glandular hairs above on convex dorsal side; pappus short, coriaceous,
almost entire. Flowering June to September.

In fields, meadows, cultivated lands, along irrigation channels,
roadsides, riverbanks, on saline land and mountain slopes.—European
part: Dvina-Pechora, Baltic Region, Ladoga-IlImen, Upper Volga,
Volga-Kama, Upper Dnieper, Middle Dnieper, Volga-Don, Trans- Volga
Region, Lower Volga, Upper Dniester, Bessarabia, Black Sea Region,
Lower Don, Caucasus: Ciscaucasia, Dagestan, Eastern Transcaucasia;
Western Siberia: Upper Tobol, Irtysh; Far East: Ussuri (introduced);
Soviet Central Asia: Aralo-Caspian Region (rarely), Lake Balkhash
Region, Syr-Darya. General distribution: Central Europe, Atlantic
Europe, Balkans-Asia Minor. Described from Sweden. Type in London.

Note. M.I. Nazarov found a hybrid in the herbarium between T.
inodorum and Matricaria chamemilla L., which was morphologically
similar to the latter species but had achenes like those of T. inodorum.
However, the presence of 4, sometimes 5, ribs on the achene is a
character typical of Matricaria chamomilla L.

Schultz-Bipontin (Sch. Bip. Ueber die Tanacet. 1844, 33) reports
a hybrid between T. inodorum and Anthemis tinctoria L., which was
described as a separate species—Anthemis hybrida Hampe.

The two specimens of this species from Central Asia preserved in
the herbarium of the Botanical Institute, 1) from the village of
Ulutavskoe (vicinity of Kzyl-Orda, collected by S.Yu. Lipschitz on
July 14, 1929, No. 436; and 2) from the village of Mugan (near the
village of Petropavlovka, Turgai Region, April, 1905), differ from
typical T. inodorum by having strong stem pubescence, especially below
the capitula. Besides, the first specimen also has pubescent leaves.

The plants of this species in the Trans-Volga Region and Lower
Volga on saline soils and along mountain slopes of the Caucasus differ
by having smaller capitula with shorter ligules, sometimes decurved at
flowering, a conical receptacle (not always persistent), outer involucral
bracts always narrowly lanceolate, pale green and with reddish veins
present at the apices of the lobes of the disk florets. However, all these
characters are also found in isolated cases in typical specimens of T.
inodorum.

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16. T. maritimum (L.) Koch. Syn. ed. II, p. 3 (1845) 1016.—T.
inodorum maritimum Sch. Bip. Ueb. die Tanacet. (1844) 33.—
Chamaemelum maritimum Boiss. Diagn. ser. I, XI (1849) 21; Willk.
in Bot. Zeitschr. 33, 22 Jahrg. (1846) 249.—Matricaria maritima L.
Sp. pl. (1753) 891; Georgi Beschr. d. Russ. Reich. IV, III, 1252; Lucé,
Fl. Osil, 282.—Matricaria inodora B. maritima Wahlenb. Fl. Suec.
(1826) 533; Ldb. Fl. Ross. II, 2, 546.—Chamaemelum inodorum var.
B. maritima Vis. Fl. Dalmat. II (1847) 85.—Chrysanthemum inodorum
L. Sp. pl. (1763) 1253.—Pyrethrum maritimum Smith, Fl. Brit. II (1800)
901; DC. Prodr. VI, 53.—Matricaria inodora* maritima Lindem. Bild.
ur Nordens FI. III (1926) tabl. 535 B.—Ic.: Lindem. |.c.—Exs.: FI.
Gall et Germ. exs. de C. Billot No. 1503.

Annual or biennial. Root branched below in numerous slender
fibrous roots. Stems solitary, 10-15-40 cm high, branched from base
or only above, glabrous, reddish, especially at base, leafy up to tip.
Leaves oblong or obovate, (2.5)3.0—5.0(9.0) cm long, (0.8)1.0—2.0(2.5)
cm wide, thrice pinnately cut into short succulent lobes, acute or
subobtuse, glabrous; lower leaves petiolate, middle and upper sessile.
Capitula 2.5—3.5 cm in dia; receptacle hemispherical. Outer involucral
bracts lanceolate, acute; inner narrow-oblong, obtuse; all bracts green,
with very narrow, light brown, membranous border. Peripheral ligulate
florets with more or less broad, short ligules. Mature achenes dark
brown, 1.5—2.0 mm long, 0.8—1.0 mm wide, with 3 light brown, thick,
obtuse ribs, of which central occupying almost entire ventral side,
dorsally rugose, with 2 red oleiferous glandular hairs above; pappus
with 2 lobes on each side of achene and short border on dorsal and
ventral sides. Flowering July to September.

Along sandy seacoasts.—Ladoga-Ilmen (introduced). General dis-
tribution: Scandinavia, Atlantic Europe. Described from seacoasts of
Northern Europe. Type in London.

Note. I am citing this West European coastal species, reported
from Leningrad city, as introduced. Earlier, when the seacoasts were
not as developed, this species could have grown more widely along
seacoasts.

Schmalhauzen (FI. II, 68) reported the species from the Baltic Sea
area (Finland, Estonian SSR—Tallin).

17. T. limosum (Maxim.) Pobed. comb. nova.—Chamaemelum
limosum Maxim. Prim. Fl. Amur. (1859) 156.—T. inodorum B. limosum
Rgl. ex Herder in Bull. Soc. Nat. Mosc. XL (1867) 43.—Matricaria
limosa (Maxim.) Kudo in Contr. Know. Fl. North Saghal. (1923) 58,
exsl. specim.—Matricaria inodora auct. non L.: Forbes and Hemsl. in
Journ. Linn. Soc. XXIII (1888) 439; Kom. Fl. Man’chzh. III, 639.—

178

181

Lee

Matricaria maritima ssp. limosa (Maxim.) Kitamura, Compos. Japon.
II (1940) 335.

Annual or biennial. Root short, fibrous. Stems erect, 8-35 cm
high, simple or strongly branched from base, branches appressed to
stem, upward spreading, erect, long, near apex branched again, sulcate,
glabrous, densely leafy. Basal leaves withering after anthesis; middle
cauline leaves oblong, 5.5—9.5 cm long and 2.5-—2.7 cm wide, thrice-
pinnate, with narrow, filiform, obtuse lobes, 0.5 mm wide, glabrous on
both sides, simple pinnate at base, amplexicaul; upper leaves strongly
reduced. Peduncles 1.5—3.0 cm long, below capitulum glabrous,
thickened and often pubescent. Capitula numerous, solitary at apices
of main stem and branches, 10-15 mm in dia. Involucre hemispherical,
3-seriate; bracts more or less equal, outer linear-lanceolate, inner
oblong, obtuse, with single brownish green stripe on light green
or whitish dorsal surface, and broad (sometimes broader than
middle greenish part), white, lustrous or slightly brownish, membranous
border. Receptacle broadly ovoid-conical. Peripheral florets with short
broad ligules, 4.5 mm long, 1.5—2.0 mm wide, less often 8-10 mm
long and 1.5-2.0 mm wide; disk florets bright yellow, with reddish
veins at tips of lobes. Anther tube sometimes strongly exserted; anthers
basally obtuse, with oblong apical appendages. Achenes with 3 whitish
broad ribs, narrow, 2 mm long, 1 mm_ wide, dorsally brown, rugose,
with 2 large, red, oleiferous glandular hairs at apex; pappus lobed,
membranous, 0.6 mm long. Flowering June to August (Plate VI,
Fig. 1.)

Along open sunny banks of rivers and lakes in marshes.—Far
East: Zeya-Bureya, Uda River Area, Ussuri. General distribution: Japan,
China. Described from the vicinity of Nikolaevsk-on-Amur. Type in
Leningrad.

18. T. breviradiatum (Ldb.) Pobed. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XXI (1961) 352.—Pyrethrum breviradiatum Ldb.
in Mém. de |’ Acad. des sc. Pétersb. V (1815) 577.—Matricaria inodora
€. breviradiata Ldb. Fl. Ross. II (1844) 547.

Annual. Stem 40 cm high, branching only in upper half. Capitula
18 mm in dia; peripheral ligulate florets almost a half as long as disk,
sometimes absent; receptacle more or less hemispherical. In other
respects it does not differ from the previous species.

Eastern Siberia: Lena-Kolyma. Endemic. Described from lower
reaches of the Kolyma River. Type in Leningrad.

Note. Besides the type specimens of Ledebour, there are no other
specimens of this species in the Herbarium of the Botanical Institute,
Academy of Sciences of the USSR, Leningrad. It has not been collected
since Ledebour’s time. Its status as a separate species is doubtful.

173

179 tate VII.
1—Tripleurospermum ambiguum (Ldb.) Fr. and Sav., habit of plant; involucral
bracts, bisexual floret, dorsal and ventral view of achene: 2—T. sevanense
(Mand.) Pobed., habit of plant, receptacle, bisexual floret, dorsal and ventral
view of achene; 3—Matricaria aurea (L.) Boiss., habit of plant, bisexual floret,
and achene with and without pappus.

182

174

Section 4. Leiospermum Sch. Bip. Ueb. die Tanacet. (1844) 33.—
Pseudo-chamomilla DC. Prodr. VI (1837) 51, p. p.; Ldb. Fl. Ross. II,
544 p. p.—Achenes smooth; pappus absent or scarcely visible along
margin; peripheral florets not ligulate or short-ligulate, with short style;
disk florets five-toothed. Annual herbs.

Type of section: T. disciforme (C.A.M.) Sch. Bip.

19. T. disciforme (C.A.M.) Sch. Bip. Ueb. die Tanacet. (1844)
34.—Chrysanthemum disciforme C.A.M. Verzeichn. (1831) 75.—
Matricaria disciformis DC. Prodr. VI (1837) 51; Ldb. Fl. Ross. II,
544.—M. corymbifera DC. Prodr. VII (1838) 297.—Chamaemelum
disciforme (C.A.M.) Vis. Fl. Dalmat. II (1847) 85; Boiss. Diagn. I, 11,
21; Boiss. Fl. or. III, 328; Grossh. Fl. Kavk. IV, 127.— Matricaria
sive, C. nanocephalum Rupr. in Bull. Phys.—Math. Ac. Sci. Pétersb.
XXIV (1856) 231.—C. disciforme B. quadrilobum Boiss. FI. or. II
(1875) 328.—C. disciforme f. radiata Sosn. ex Grossh.; Fl. Kavk. IV,
127.— Exs.: Pl. or. exs. No. 298.

Biennial. Plants glabrous; stem erect, 40-100 cm high, sulcate-
striate, branched near apex. Leaves oblong, twice or thrice pinnate,
(3.0)4.5-6.0(17.0) cm long, (0.8)1.5—2.0(3.5) cm wide, with narrow
linear segments; lower leaves petiolate, middle and upper sessile,
broadened at base; upper leaves strongly reduced. Inflorescence
corymbose at apices of stem and branches, with 3—4 capitula on 3.5—
10.0 cm long leafless peduncles; capitula 8-12 mm in dia; involucral
bracts obtuse, with wide, white, membranous, lustrous border; receptacle
hemispherical. Capitula usually with only tubular disk florets, latter 5-
toothed, 3.0-3.5 mm long, with oblong, 2.0-2.5 mm-long tube and
short 0.25-0.5 mm long lobes, red, resinous, glandular hair at tip of
each lobe, less often capitula also with short peripheral ligulate florets
shorter than involucre. Anthers narrow, oblong, with deltoid apical
appendages slightly exserted from throat of floret. Stigma branches
exserted from anther tube. Achenes smooth, 1.25—1.5 mm long, 0.25-
0.5 mm wide, with 3 narrow white ribs on ventral side in upper border
part, 2 red oleiferous glandular hairs at apex on dorsal side. Flowering
May to September. (Plate VI, Fig. 2.)

In wet mixed herb meadows along riverbanks, ravines, in dry
riverbeds, and as weed in cultivated fields and by roadsides—Caucasus:
Talysh, Southern Transcaucasia; Soviet Central Asia: Kyzyl-Kum
(southeastern), Pamiro-Alai Region, Tien Shan, mountainous
Turkmenia. General distribution: Armenia-Kurdistan, Iran. Described
from Talysh. Type in Leningrad.

Note. The involucral bracts in this species vary in shape and width
of the membranous border; the outer bracts are sometimes entirely

175

without the membranous border and with an acute tip. Frequently,
ligulate ray florets with short (shorter than the involucre) white ligules
occur with tubular florets in the capitula, but in all other characters
they are indistinguishable from the typical plants. From the Zeravshan
Valley (Oal Ravine). O. Fedtschenko collected T. disciforme with
achenes having a quite long, scaly, lobed, pappus, but smooth and not
rugose as in 7. decipiens.

20. T. decipiens (Fisch. and Mey.) Bornm. in Fedde, Repert, XLIX
(1940) 240.—Pyrethrum decipiens Fisch. and Mey. Ind. Sem. Hort.
Petrop. II (1835) 48.—Chamaemelum decipiens (Fisch. and Mey.)
Boiss. Fl. or. II (1875) 328.

Annual or biennial. Stems erect, 40-100 cm high, cylindrical,
sulcate, glabrous, branched above. Leaves larger in lower part of stem,
oblong, 6-9 cm long, 0.8-2.0 cm wide, twice or thrice ternate, with
short cuspidate segments, broadened toward base, once or twice pinnate;
leaves in upper part of stem gradually reduced, sessile. Inflorescence
corymbose, compound, with numerous capitula on 5—15 cm long leafless
peduncles, thickened below capitula. Capitula 7-10 mm in dia;
involucral bracts obtuse, with wide, white, membranous border;
receptacle hemispherical; capitula with only tubular florets. Anthers
narrow, oblong, with deltoid apical appendages, slightly exserted.
Stigma branches exserted from anther tube. Achenes rugose, with 3
white ribs on ventral side and 2 red oleiferous glandular hairs at apex
on dorsal side; pappus membranous, lobed, very short. Flowering June.

In wet alpine and subalpine meadows, and cultivated fields.—
General distribution: Northern Iran, Asia Minor. Described from
northern Iran (Hoi). Type in Leningrad.

Note. This species has not yet been collected in the USSR, but is
found close to its border in northern Iran. Possibly it has been over-
looked in the Caucasus, as it is quite similar to T. disciforme. How-
ever, the species are so different in the structure of their achenes that
they cannot be confused. Besides in the achenes, T. decipiens differs
from T. disciforme by having a longer receptacle and often a more
compound corymb.

21. T. sevanense (Mand.) Pobed. in Bot. Mat. Gerb. Bot. Inst.

Akad. Nauk SSSR, XXI (1960) 353.—Chamaemelum sevanense Mand.

in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954) 376.—

C. trichophyllum auct. cauc. non Boiss.; Grossh. Fl. Kavk. IV, 128.
Annual. Plants glabrous; stems erect, 30-45 cm high; sulcate-striate,
branched from base or only above. Leaves oblong; lower leaves 4.0—

183 4.5 cm long, 0.8-10.0 cm wide, long-petiolate, basally broadened and

176

with short, segments, twice or thrice pinnate, with filiform thin cuspi-
date segments, gradually reduced upward. Capitula few, solitary, on
4-10 cm long peduncles, 1-2 cm in dia, with ligulate florets. Involu-
cral bracts few; outer bracts ovate, obtuse, pale green, with narrow
pale brown border, inner bracts obovate, very broadly, especially at
apex, white-membranous. Outer florets with 3-toothed white ligules;
tubular disk florets with orange glandular hairs at tips of lobes. Achenes
1.5 mm long, 0.5 mm wide, broadened at apex, smooth, with thin
filiform ribs, large, red, oleiferous glandular hairs, pappus white,
membranous, lobes less than a fourth as long as achene. Flowering
July. (Plate VII, Fig. 2.)

In ravines, wet meadows.—Caucasus: Southern Transcaucasia.
Described from Armenia (near Bavadyan-Dar). Type in Tbilisi; isotype
in Leningrad.

Note. It differs from T. disciforme by having outer involucral bracts
with a narrow pale brown border, ray florets present in the capitulum
with long white ligules that are twice as long as the involucre, and
achenes, with two large, red, oleiferous glandular hairs on the dorsal
side, and a white-membranous, lobed, short pappus. It also differs
from T. decipiens, by having smooth and not rugose achene.

Section 5. Gastrosulum.—Sect. Eugastrosulum (Sch. Bip.)
Pobed.—Eugastrosulum in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XXI (1961) 354.—Sect. Eugastrosulum generis Gastrosuli Sch.
Bip. Ueb. die Tanacet. (1844) 29.—Achenes with two white lateral
ribs and a third suberous, triangular rib, more prominent ventrally,
separated from lateral ribs by narrow furrows; achenes dorsally brown,
with glandular hairs at apex; pappus membranous, bilobed, long, almost
half as long as a achene, dorsally longer.

Type of section: Tripleurospermum parviflorum (Willd.) Pobed.

22. T. parviflorum (Willd.) Pobed. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XXI (1961) 354.—T. praecos (MB.) Bornm. in -
Beich. Bot. Centralbl. LX (1940) 192.—Pyrethrum parviflorum Willd.
Sp. pl. III, 3, (1804) 2158.—P. proecox M.B. FI. taur.-cauc. II (1808)
324.—Gastrosulum praecox Sch. Bip. Ueb. die Tanacet. (1844) 29.—
Matricaria inodora y. praecox Ldb. Fl. Ross. II (1844) 546.—M.
praecox DC. Prodr. VI (1837) 52.—Chamomilla praecox C. Koch in
Linnaea, XVII (1843) 45.—Chrysanthemum monspeliense Georgi,
Beschr. d. Russ. R. IV, 3 (1800) 1251?—Chamaemelum praecox Vis.
in Jorn. Bot. Ital. 2 (1844) 34; Boiss, Fl. or. III (1875) 326; Stapf.
Erg. Polak. exp. (1885) 58; Grossh. Fl. Kavk. IV, 127.—Ic.: Fl. Turkm.
VII, 97.

184

Bit

Annual. Stems solitary or few, simple or weakly branched, 10-30
cm high, sulcate, weakly pubescent, especially below capitula. Leaves
oblong, 3-5 cm long, 0.5—1.0 cm wide; basal and lower cauline leaves
bipinnate, long-petiolate; upper cauline leaves smaller, pinnate, sessile;
all leaves deeply divided into linear, long, subulate segments; rachis
broadened at base, and with pinnate, short, distant segments. Capitula
solitary on leafless peduncles, 3-10 cm long, weakly pubescent, slightly
thickened below capitula; receptacle conical, acute or obtuse, elongate, -
less often almost hemispherical, punctate. Outer involucral bracts
oblong, inner obovate, wide; all bracts 2-3 mm long, more or less
broadly white- or light brown-membranous along margin. Florets in
capitulum heterogamous; peripheral florets pistillate, with white, short,
broad ligules, 3—6 mm long, 2-3 mm wide, apically bifid, or 3-toothed,
with short middle tooth; tubular disk florets lemon-yellow, 1.5—3.0
mm long, with 5 broadly deltoid lobes. Anthers with very minute, 0.25
mm long, narrow-triangular appendage, exserted. Style branches with
red stripe in center and yellow border at ends. Achenes on maturity
1.0-1.25 mm long, 0.5 mm wide, mucilaginous, ventrally with wide
spongy column, separated by narrow furrows from lateral winged ribs,
convex, dorsally smooth and brown; pappus long, a half as long as
achene, membranous, bilobed, with long ovoid lobes on ventral side
and broad coarsely toothed lobe dorsally. Flowering March to June.

In wormwood deserts and semideserts.—European part: Lower
Volga; Caucasus: Ciscaucasia, Eastern and Southern Transcaucasia;
Soviet Central Asia: Kara-Kum, mountainous Turkmenia. General
distribution: Eastern’ Mediterranean Region, Armenia-Kurdistan, Iran.
Described from unknown locality. Type unknown.

Note. The reports (DC. Prodr. VI, 52; Vis. in Giorn. botan. Ital.
II, 1, 35) of the occurrence of T. parviflorum in Crimea are erroneous.
Apparently, this error began with De Candolle (l.c.), who reported the
classic locality of this species as Crimea. The error of De Candolle or
Visiani is also repeated by I.P. Mandenova in Fl. Gruzii (VIII, 336).

It must be noted that specimens of this species from Armenia
usually have pubescent stems, leaves, and peduncles. On Meyer’s label
such plants are annotated as puberula.

185

178
GENUS 1528. Pyrethrum ':?

Zinn. Catal. Pl. Gotting (1757) 414; Scop. Fl. Carn. ed. 2, II
(1772) 148; Gaertn. De fruct. et sem. pl. II (1791) 430, non Medik,
1775.—Balsamita Mill. Gard. Dict. Abridg. ed. 4 (1754).—Gymnocline
Cass. in Bull. Soc. Philom (1816) 199 and in Dict. Sc. Nat. XX (1821)
119 p. p.—Richteria Kar. and Kir. in Bull. Soc. Nat. Mosc. XV (1842)
126.

Capitula solitary or more numerous, 2—40, less often to 100 (section
Gymnoclines on single stem, and then usually aggregated in corymbose
or corymbose-paniculate inflorescence, usually heterogamous, with 1-
50, outer pistillate ligulate florets, in one (less often in more or less
2) row, and numerous bisexual, tubular disk florets, very rarely (in P.
majus) homogamous, with bisexual tubular disk florets only. Involucre
patelliform, less often more or less poculiform, 4-25 mm in dia, 3-8
mm long; involucral bracts herbaceous or coriaceous-herbaceous,
imbricate in (2)3—4(5) irregular rows, often more or less membranous
along border, less often entirely without border; outer bracts lanceolate
or lanceolate-linear, inner oblong or oblong-linear, usually 1.5—2 times,
less often (section Brachyglossa) more or less 3 times as long as outer.
Receptacle usually strongly or weakly convex, less often almost flat,
solid, glabrous, more or less punctate-tuberculate, sometimes distinctly
alveolate. Corolla of ligulate florets white or pink of different shades,
with dorsally flattened but wingless or almost wingless 1.0-2.5 mm
long tube and oval, oblong, or linear, less often (section Gymnoclines)
reniform, 2-25 mm long limb; corolla of tubular disk florets yellow,
1.2—4.0 mm long, usually with shorter tube, in upper half slightly and
gradually broadened, less often rather strongly and abruptly broadened
(section Leucanthemopsis), usually more or less recurved triangular or
lanceolate-triangular teeth, 1/7—1/4 as long as tube. Filaments more or
less thickened in upper part; anthers lacking distinct basal appendages,
with lanceolate-ovoid or broadly lanceolate, usually tubular apical
appendages; pollen grains spherical, spiny. Style bifid; style branches
linear, truncate. Achenes glabrous, more or less prismatically cylindrical,
narrowed toward base, 1.0-3.5 mm long and 0.3-1.0 mm wide, with
5-10 (very rarely to 12) more or less protruding longitudinal ribs.
Ligulate florets usually with achenes more or less shifted ventrally;
pericarp usually more compactly enclosing kernel (loosely in section
Leucanthemopsis), lacking mucilaginous cells (less often present, but

‘Treatment by N.N. Tzvelev.
7From the Greek word pyrethron or pyrethros—the name of a composite used by
Dioscorides and Nicander, possibly belonging to this genus.

186

179

never in regular rows) and secretory canals; pappus 0.1—1.5 mm long,
as more or less toothed or lobed corona along margin, sometimes
divided to base in lobes of various form and size or truncate almost to
base dorsally, often smaller in ligulate florets than in tubular. Perennial
herbs (rarely somewhat woody at base), more or less covered with
simple or branched hairs (sometimes entirely glabrous), with erect or
basally ascending stem and alternate leaves, more or less divided, less
often (sections Balsamits and Balsemitopsis) with entire laminas.

The genus comprises about 100 species, distributed in Europe
(except Far North), North Africa, and in the greater part of nontropical
Asia (up to Lake Baikal in the east and the Himalayas in the south).
The Mediterranean Region, the Caucasus, and Northwest Asia are
particularly known for the number and diversity of species; the number
of species in the genus significantly decreases toward the north and
east of these regions. Fifty-six species are found in the USSR.

Type of genus: Pyrethrum corymbosum (L.) Willd.

Note. In recent times, this genus usually is united either with the
genus Tanacetum L. or with a whole series of other closely related
genera under Chrysanthemum L. sensu lato, with which one can hardly
agree (cf. notes to the genera Chrysanthemum and Tanacetum). In my
opinion, it shows the greatest affinity not so much with the genus
Tanacetum as with the genera Trichanthemis Rgl. and Schmalh. and
Cancrinia Kar. and Kir., which even O. Hoffmann (Die natiirl.
Pflanzenfam. Teil IV, Abteil 5, 1889, 282-283), who accepted the
genus Chrysanthemum in the broadest sense, considered as separate
genera. The species P. tianschanicum possibly might be included with
equal justification under either the genus Pyrethrum or the genus
Trichanthemis, but the species P. majus and the rarely encountered
eligulate specimens of the species in section Parthenium could easily
be referred formally to the genus Cancrinia. Besides, some species of
the genus (e.g., P. komarovii and P. tricholobum) are so similar to
species of Tripleurospermum Sch. Bip., that in the absence of mature
achenes they still occupy a somewhat intermediate position between
these genera and have been included in the genus Pyrethrum, to some
extent, only tentatively. The numerous sections of the genus
(unfortunately, their number cannot be reduced satisfactorily), despite
their apparent, relatively less distinctness by comparison with the
sections of closely related genera, still by no means always show a
clear affinity with each other, whereas their relationships with other
genera often are even more distinct. All this points to the need for a
more detailed revision of the genus making comprehensive use of not
only morphological but also anatomical characters, which have not
been used in the present description for lack of time.

180

Economic Importance. Many species of the genus (in particular,
P. parthenium, P. coccineum, etc.) are used as ornamentals; some
species (P. cinerariifolium, P. coccineum, P. roseum) produce the in-
secticidal substance pyrethrum, which is used in the manufacture of
different insecticides (pyrethrum powder, etc.). The species P. majus
and P. balsamita are vegetables.

1. Lamina entire, more or less toothed, oblong to broad oval, some-
times lyrate in cauline leaves, with large broadly oval apical lobe

and few :very:smallwlateral lobes, e::1:3.4. cies. .f Alec east hes D
+ Lamina moreloriess pittnatelyseut:)). <7.1.:2650) mde. bot. ccp hone 5.
2. Limb of corolla of ligulate florets 9-17 mm long. Southern Altai
(sect. Balsamitopsis) ........... 15. P. kelleri (Kryl. and Plotn.) Krasch.
+ Limb of corolla of ligulate florets 4-8 mm long, or ligulate florets
entirelysabsent:(sect. Balsamita)»..x is eiti2... 26m tee eee ee ee 3,

3. Ligulate florets absent. Cultivated plant, less often escaped .............
ey, te SSR UTES. 21, meee gree eee 5. P. majus (Desf.) Tzvel.
Ligulate florets presents Caticasus s «ch iats.i320....4.. detuseecen. tee ame 4.
Basal leaves with lamina more or less abruptly narrowed in petiole;
capitula (5)10—-50(70); involucral bracts greenish or brownish
WIA MERU. Le, OR OT, SPR ge tens ies 3. P. balsamita (L.) Willd.
+ Basal leaves with lamina gradually cuneately narrowed in petiole;
capitula (1)2—20(15); involucral bracts usually dark brownish-violet
ELON AMEE kt, Be oae ee Be 4. P. balsamitoides (Nabel.) Tzvel.
5. Lamina of basal and lower cauline leaves ovate or broadly oval,
pinnately lobed, with 2—5 oblong, entire, lobes on each side. Alpine
plant, 5-15 cm high. Carpathians (sect. Leucanthemopsis) ....
SOEUR MES he, Siete 2b a een. pees 27. P. alpinum (L.) Schrank.
Leaves (except uppermost) more or less pinnatisect ...............00. 6.
Plants 50-150 cm high, lacking short nonflowering shoots. Basal
leaves withering early; cauline leaves numerous, pinnately parted
(or pinnately cut), with entire, more or less toothed lobes. Capitula
(8)10-80(100), aggregated at stem apex in compound corymbs;
involucres 4-8 mm in dia. Caucasus (sect. Gymnoclines) ........... A
+ Plants usually shorter, usually with short non-flowering shoots and
leaves of various shapes. Capitula fewer, often solitary .............. 8.
7. Involucre 4—6 mm in dia; corolla limb of ligulate florets 2-4 mm
long win, ASE. Utd 1. P. macrophyllum (W. and K.) Willd.
+ Involucre 5—8 mm in dia; corolla limb of ligulate florets 8-11 mm
LOmptici!. act. tet. 2200 ALA po tath ogee 2. P. sorbifolium Boiss.
8. Capitula (1)4—25(40), in rather lax corymbose-panicules; involucre
6-11 mm in dia; corolla limb of ligulate florets 5-7 mm long;
lamina twice or thrice pinnately cut, with apical lobes up to 1 mm

+

DN +

188

So +

10.

i.

12.

is.

14.

{3.

16.

een stat ditterent type wisitestl. ive. Llanes 9.
Plants silver-gray from dense pubescence or strongly flattened,
appressed, bifid hairs. Found only in cultivation (sect. Cinerariifolia)

Spo 8) eee Oe OPE 16. P. cinerariifolium Trev.
Plants glabrous or more or less covered with simple or bifid, usually
ereepane montinttoned hairs’ 2.0.0... QUAL a 10.

Semishrubs, 5-25 cm high, with thick woody rhizome and long,
often more or less branched, woody bases of nonflowering and
flowering shoots; apical lobes of leaves linear-filiform, to 0.6 mm

WIGS ca ReE a ROMMLALIYO S220 5.552 023001. Liles de eas. I AL: Pd.
Herbs, less often more or less woody at base, but then apical lobes
of leaves broad, not linear-filifOrmM ..0....... eee eeeeneceeeeteeeneeeeees 14.

Corona of achenes 1.2—2.5 mm long, divided to or almost to base
into 6-10 scales or lobes. Western Tien Shan (sect. Tri-

eo) |) 42. P. tianschanicum Krasch.
Corona of achenes 0.2-1.0 mm long. Caucasus (sect. Xylo-
UNE Srcia oiacisassivsancssdcndnacecdaaaadeectbenlcaneeeeeenraes ¥2.
Plants dull green, weakly pubescent; corona of achenes 0.2-0.3
i ct! ae Se oe 10. P. heldreichianum Fenzl. ex Tchih.
Plants grayish, densely pubescent; corona of achenes 0.5—1.0 mm
og ae 2a" a a nd eee ere Py a 15.

Outer involucral bracts broadly lanceolate, equal to or two-thirds as
long as inner; membranous border of bracts brownish .................
Perr ah | inna oe 29. P. ordubadense Manden.
Outer involucral bracts lanceolate, a half to two-thirds as long as
inner; membranous border of bracts very dark brown .............
oo eee 1 LE 28. P. kotschyi Boiss.
Plants with densely leafy stem and not always with short,
nonflowering shoots, usually covered with short simple hairs; all
leaves rather long petiolate. Achenes 1.0-1.5 mm long. (sect.

Pirtlentatay 335.00. AN, ead SRO i EE 15.
Middle and upper cauline leaves sessile, less often short petiolate,
but then stem very weakly leafy and achenes larger ................. 22.

Receptacle weakly convex; lamina once or twice pinnately cut,
with lanceolate or broadly linear apical lobes ..............ccceseeseeeeees
RL REN, AR ik. 6. P. peucedanifolium (Sosn.) Manden.
Receptacle strongly convex; lamina with apical lobes oblong to
Pu Teo) 1 ne Sie BEE cee as 7 16.
Plants grayish from dense fine tomentum ..............:::cccesseeeseees if.
Plants green, weakly pubescent (almost glabrous) ................0 18.

182

189 fe

23s

190 26.

Lamina with 1st and 2nd order lobes usually entire, oblong-ovate
to almost semicircular, often obtuse-crenate or shallow-lobed along
margin; involucre 9-11 mm in diameter \.).:0c20 ha -sch-dee eect ceestee ee
ah sede sahsbed dates cu tes sbcSe CEA MOREEUCL « SMepaneaLD: SEMEL I 13. P. kubense Grossh.
Lamina with secondary lobes pinnately parted or pinnately lobed,
with oblong or ovate apical lobe and other segments; involucre 7—

TO 'mmi«indialy iw. ees nel oe BK 12. P. fruticulosum Biehl.
Corolla limb in ligulate florets (0)2.5—6.0(7.0) mm long ............ 19.
Corolla limb in ligulate florets (6)7—12(15)mm long ................. 20.

Plants yellowish-green, cultivated, sometimes escaped ................
a deka AIRMEN LAUSD, EERO, Ped eR 8. P. parthenium (L.) Sm.
Plants green, usually wild; lamina more strongly divided .................
ee AALS TNORREL, SELB co tesOeih ky. RRCACUIN BEN TN Ren 10. P. grossheimii Sosn.
Capitula solitary or 2—15(20); involucre 9-16 mm in dia; plant of

upper mountain zone ............ 9. P. glanduliferum Somm. and Lev.
Capitula (1)5—20(30); involucre 6-10 mm in dia. Lower mountain
POURS RT ESE GSR, 6 occcas le cete clan cenacesectons dR PERE EERE onEae ZL

Lamina with pinnae and pinnules, entire or weakly lobed .................
wsitchawng os vadigu cle uder uaslnslaaeiese en ante aeece Renata 7. P. parthenifolium Willd.
Lamina with lobes pinnately parted or pinnately lobed, with oblong
or ovate apical lobe and segments .............. 11. P. sevanense Sosn.
Receptacle weakly convex, almost flat. Plants usually grayish from
dense pubescence, less often (P. galae) green and more weakly
pubescent, but then leaves pinnately parted «0.0.0.0... cee eeeee eee 23.
Receptacle strongly convex, hemispherical or almost so, less often
(in P. silaifolium) weakly convex, but then plants green, weakly
pubescent, with twice or thrice pinnately cut leaves ................. 29.
Involucre 9-20 mm in dia, narrow-patelliform; achenes with 8—12
ribs; leaves twice pinnately cut. Caucasus. (sect. Richteriopsis.)
TEER AEA ESA EMM ST HEER | 4) nes Crys pe tn 8 Mab te 28k Ep ee Pa Sales 24.
Involucre 6-13 mm in dia, poculiform; achenes with 6-8 ribs; leaves
less divided. Soviet Central Asia. (sect. Brachyglossa.) ........... 26.
Outer involucral bracts broadly ovate, with rather wide membra-
nous: borden (F554... Ae, ee ee 37. P. aucherianum DC.
Outer involucral bracts narrow-lanceolate, acuminate, lacking or
almost lacking ‘membranous: border\<:%.:.-c:.2i0iss-.aysieeeee sabes ceee 25.
Pubescence appressed, grayish; all involucral bracts acuminate .....
AR ACM Lepr) PL te) .a 0) ani! Oo Ae 36. P. oxylepis (Bordz.) Tzvel.
Pubescence more lax, brown on drying; inner involucral bracts
SUNDGHULISE ss cevcceeaccsces te ne otetes amecce estas 35. P. sericeum (Adam) MB.
Outer involucral bracts a third to a half as long as inner. Plant
densely tomentose or with simple hairs. Trans Ili Alatau and Ketmen
TATE ih. coca pesceusin etstecceeiqgah reeds 41. P. semenovii (Herd.) Winkl.

191

at.

28.

ao.

30.

32.

33.

34.

35.

183

Outer involucral bracts a half to two-thirds as long as inner; Pubes-
cence of bifid hairs. Western Tadzhikistan ranges ............ i

Lamina irregularly pinnately parted, with entire, more or less lobate
lobes; plants dull green, more weakly pubescent ..............s:ccesseeeeeeees
SC. AAS SELON Ss RT, GI, TN lcd, Mh 39. P. galae M. Pop.
Lamina pinnately cut, with more or less pinnately parted or lobed
lobes; plants grayish from rather dense pubescence ................ 28.
Pubescence of plants appressed, tomentose, consisting of flexuous
Shinpic:and bifid Haig 0.) ives sascsccessosves 38. P. hissaricum Krasch.
Pubescence of plants densely tomentose, comprising more or less
erect, bifid and simple hairs ................ 40. P. mikeschinii Tzvel.
Apical lobes and segments of leaves broad, broadly ovate to
lanceolate, more than 1 mm wide; corona of achenes 0.5—1.5 mm

long, toothed or lobed to middle. (sect. Pyrethrum.) ..........:.0+ 30.
Apical lobes of leaves linear or linear-lanceolate, to 1 mm or more
wide, but then corona of achenes 0.1—0.3 mm long ................06 33.

Plant dull or grayish green from rather dense pubescence; capitula
solitary or 2—6 on single stem. Caucasus ...........:.sccceccsteeeeeeeteeeneeeeees

Beteeteen,, Weakly pubescent ....0eii..0s eocders.dhsedal bee Meanaaese 31.
Involucre and peduncles of capitula glabrous; Plants very weakly
pubescent with long simple hairs. Dzhungarian Alatau ................
ACURA OVA HOS AAR RL 33. P. corymbosiforme Tzvel.
Involucre and peduncles of capitula more or less pubescent; plants
primarily comprising bifid hairs ..0000..:35..00008. ss.cc05. Aes 32.
Involucral bracts with light colored or brownish membranous border;
receptacle 2—3 times as wide as long. A widely distributed species
GIR IATTOMIR HG RA IR 31. P. corymbosum (L.) Willd.
Involucral bracts with dark brown, usually wider membranous
border; receptacle usually 1.5 times as wide as long. Carpathians
URINE dh sdisivdh en ld AA RANA 32. P. clusii Fisch. ex Rchb.
Leaves with broader apical lobe and segments less often (in P.
tricholobum and P. komarovii) narrower, linear or linear-filiform;
corona 0.1—0.4 mm long, very rarely (in P. komarovii) 0.5—1.3 mm
long. Caucasus:.(sect: | Pyrethrellum.) \iiisicseisicatbsdtias Uaciekaelleosavs 34.
Leaves with narrower, linear or linear-lanceolate lobes; corona 0.4—
1.5 mm long, less often to 0.3 mm. Siberia and Soviet Central Asia.

(Geet RICKI CTIA: ) aositensssercsthth Koh! LN. A SLDSALY - O REM Bhsioes 43.
Apical lobes of leaves linear-filiform, up to 0.8 mm wide ....... 35.
Apical lobes and segments of leaves broader, almost semicircular
to.oblong.or broadly. lanceolate sasiiiioneltedis LENGE. A DMI wees 36.

Corona 0.5—1.3 mm long. Southern Transcaucasia .............ccseeee
wena 2). aed peel nomads. 26. P. komarovii Sosn.

192

184

38.

39.

40.

41.

42.

43.

Corona 0.2-0.3 mm long. Main Caucasus Range ...........::ccsccseeeseeeeee

scabies pnnep a hSoRee eee eae 25. P. tricholobum Sosn. ex Manden.
Ligulate florets pink of different shades «0.0.0.0... eeceeeeseeeeeees 37.
Ligulate flonets;white; widen eases. nendan tah. etic eee 38.

Lamina pinnately cut, with pinnae more or less dentate or pinnately
lobed;;caulineyleaves»tathenmnumerous ps -sietacete: helo eka.
ina teanasnes ROR Os Gores aaah oer syseml, J 19. P. roseum (Adam) MB.
Lamina twice-pinnately cut; cauline leaves fewer ...............000
Se ee eee ee 20. P. coccineum (Willd.) Worosch.
Plants 20-80 cm high, with rather densely leafy stem and weakly
developed short nonflowering shoots; lamina pinnately cut, with
segments more or less toothed or lobate ...............:cssccessreceereeees 39.
Plants 5—50 cm high, with weakly leafy stem and well developed,
short nonflowering shoots; lamina of basal and lower cauline leaves
pinnately cut, with pinnately parted segments .................:ccee0e 40.
Marshy plants, with oblong, rooting, nonflowering shoots at base.
Rangemof:the Lesser-Gaucasus 3. .ccen.-telvoxy no. dads tel. cae
winisd seni yi te spike ede Sonn SAA eR, Oe 17. P. punctatum (Desr.) Bordz.
Rupicolous and meadow plants with short nonflowering branches.
Abkhazian ASSR ss.c.:0cinsadieeelne cate 18. P. marionii Albov.
Involucre 12-17 mm in dia; achenes 2.4—3.0 mm long with very
short (about 0.1 mm long), finely toothed corona. Western part of
the Greater Caucasus (upper reaches of the Rioni River) .................
saemeeducnanokns 21. P. chamaemelifolium (Sosn. and Lev.) Sosn.
Involucre 8-13 mm in dia; achenes 1.8—-2.5 mm long with longer
(0.2-0.5 mm long), more or less toothed corona. Eastern part of the
GreatéemCaucdsus .::..4...n¢s tecoieiute. ge eels)... Rosana. 41.
Plants 10-40 cm high, not aromatic, usually covered with sparse
bifid and simple hairs; capitula on long (up to 25 cm long) peduncles;
receptacle weakly convex, almost flat ........ 22. P. silaifolium Stev.
Plants 5—18 cm high, strongly aromatic, usually covered with simple
hairs; capitula on shorter (to 3-4 cm long) peduncles; receptacle

strongly convex, almost hemispherical ................:cessseesseeeeeeeeees 42.
Leaves more or less pubescent, usually dull green; outer involucral
bracts broadly lanceolate, more or less pubeSCeMt ............s0scesesseeeeeeeees

sites omenctesl 23. P. daghestanicum (Rupr. ex Boiss.) Rupr. ex Fler.
Leaves glabrous or subglabrous; green; outer involucral bracts
lanceolate, glabrous or subglabrous ...............::cssssescesseeeeesssssesesscssensees
sisttndene ery RE once, be 24. P. aromaticum (Rupr. ex Boiss.) Tzvel.
Leaves more or less grayish or whitish from dense pubescence.
Soviet. Central Asia, southermAltaiy.i2.21.cc sees, eizeneetletesenttec sae 44.
Leaves green, less often dull green, more or less sparsely pubescent
to glabrous and subglabrous; less often (in P. lanuginosum from
Sayans) more or less sparsely floccose-tomentose ..............0008 47.

193

44.

45.

46.

47.

48.

49.

51.

185

Corona of achenes usually not more than a half (less often to two-
thirds) irregularly toothed or lobed Dzhungarian Alatau ................
ph itera eh, seeped betes) etic. 50. P. songaricum Tzvel.
Corona of achenes divided to base in oblong or linear-oblong scales
ates inate lai Lb b lellian Le sede...) 45.
Plants 3-15 cm high, covered with rather densely tomentose bracts
through which outline of laminas and involucral are scarcely visible.

Central Tien Shan ................. 58. P. leontopodium (Winkl.) Tzvel.
Plants less densely hairy; outline of laminas and involucral bracts
Ute: distinct lwo ah Bil ncn nctaie nn eee RS. 46.

Plants 5—25 cm high, with less densely hairy, usually somewhat
grayish leaves; capitula soltary; blackish-brown membranous border
on involucral bracts wide, quite distiNct .............:cseseseeeeeeeseeteeseeees
Bete sat 51. P. pyrethroides (Kar. and Kir.) B. Fedtsch. ex Krasch.
Plants 12-30 cm high, with more densely hairy, usually whitish-
pubescent leaves; capitula solitary or 2-3 on single stem; blackish-
brown membranous border on involucral bracts narrower, less
distinct because of dense pubescence .......... 52. P. neglectum Tzvel.
Corona of achenes to base, less often almost to base, divided in
oblong scales. Plants 5—25(35) cm high, with twice-pinnately cut
igames: Soviet Ceéritral Asia: .22)...caa...ageeie. aie. en ee, 48.
Corona of achenes usually to a half (less often to two thirds)
irregularly toothed or lobed. Plants (5)15—S0(70) cm high ........ 50.
Leaves usually dull green, less often pubescent to subglabrous;
bases of capitula and their peduncles always more or less pubescent
es Sele NOE 53. P. transiliense (Herd.) Rgl. and Schmalh.
Plaatentisely glabrous .......nrc2.medimna ie ee AS 49.
Plants 8-25 cm high; twice-pinnately cut, with linear-lanceolate
ahdlinear-subulate lobes i; 22...A..n mimes SRE.)
fa Oy 54. P. arassanicum (Winkl.) O. and B. Fedtsch.
Plants 5—12 cm high; leaves pinnately cut, with pinnately parted or
even pinnately lobed lobes, secondary lobes shorter ....................
notated. guxanelae:. Mom 55. P. djilgense (Franch.) Tzvel.
Plants 25-70 cm high; lamina of basal leaves to 4-5 cm wide,
twice or thrice pinnately cut; cauline leaves twice pinnately cut ...
OWI eal, ace tel, MeL TEAR ees ae Bok PEAS, Hl:
Plants 5—30 cm high; lamina of basal leaves to 1.5—2.0 cm wide,
pinnately cut, with more or less pinnately parted segments
(sometimes partly entire); cauline leaves pinnately cut, with entire
orpartly:2-5-parted sepments ik. heh Ana ands 53.
Capitula solitary, very rarely 1-2 per stem. Altai ................ccccceeeee
BOA AAR RO 44. P. krylovianum Krasch.
Capitula 2-8 per stem, very rarely solitary ..............c:ccscesesees a2:

194

186

52. Involucre 8-12 mm in dia; ligulate florets 6-18, with 7-12 mm-
long corolla limb ............... 45. P. abrotanifolium Bge. ex Ldb.

+ Involucre 10-18 mm in dia; ligulate florets 18-35, with 12-15 mm-
long corolla limb ...... 43. P. alatavicum (Herd.) O. and B. Fedtsch.

53. | Peduncles of capitula dull whitish-tomentose above; involucre 10-
15 mm in dia, whitish-tomentose only at base; ligulate florets 10—

20» Siberian. Loxvedlaincsbaeeenisme oon 46. P. pulchellum Turcz.

+ Peduncles of capitula densely tomentose; involucre 13-28 mm in
dia; involucral bracts densely tomentose at base and dorsally; ligulate

florets, 1S=395:. 08.20. eR ER ea ie gay ee 54.

54. Leaves more or less floccose-tomentose, glabrescent, usually also
near base of branches in leaf axils. Sayan Mountains .....................

ERS et CON Te 47. P. lanuginosum (Sch. Bip. and Herd.) Tzvel.

+ Plants with weakly pubescent leaves, not tomentose at base of
branches. Altai and Soviet Central Asia ............csccceesccessseeeeneeeees 55.

55. Corolla limb of ligulate florets broadly linear, corolla of tubular
florets 2-3 mm long. Soviet Central Asia .0..........ccccceseessesseseceeeteeees

sent oda 88 CE. ss cess ADAIR ORAE te vse 49, P. Karelinii Krasch.

+ Corolla limb of ligulate florets linear or narrow-linear; corolla of
tubular florets 3.0-3.5 mm long. Altai ........ 48. P. pulchrum Ldb.

Section 1. Gymnoclines (Cass.) DC. Prodr. VI (1837) 57 p. p.—
Pyrethrum sect. Leucogymnocline (Sch. Bip.) Boiss. Fl. or. HI (1875)
346.—P. sect. Gymnocline (Cass.) Sosn. subsect. Leucogymnocline
(Sch. Bip.) Sosn. in Tr. Tifl. Bot. Sada, XVII (1915) 44.—Gymnocline
Cass. in Bull. Soc. Philom. (1816) 199 and in Dict. Sc. Nat. XX
(1821) 119 p. p.—G. sect. Leucactis C. Koch in Linnaea, 339.—
Tanacetum sect. Leucogymnocline Sch. Bip. Tanacet. (1844) XXIV
(1851)/53.—Chrysanthemum sect. Gymnoclinia Benth. and Hook. f.
Gen. pl. II (1876) 427 p. p.—C. sect. Gymnocline (Cass.) Hoffm. in
Pflanzenfam. IV, 5 (1889) 278 p. p.—Herbaceous plants, 50-150 cm
high, more or less covered with short and long simple hairs, with
densely leafy stem, without short nonflowering branches; leaves
pinnatisect. Capitula numerous, from (5)10 to 100(150), aggregated at
stem apex in dense or somewhat lax corymbs; involucre narrow-
patelliform, 4-8 mm in dia; outer involucral bracts a half to two-thirds
as long as inner; receptacle weakly convex; corolla limb of ligulate
florets 2~11 mm long; corolla of tubular florets 1.8—2.4 mm long, with
slightly gradually broadened tube and lanceolate-triangular teeth in
upper half. Achenes 1.6—2.5 mm long, with 5-8 longitudinal ribs, more
or less shifted ventrally in ligulate florets; corona 0.2—0.3 mm long,
irregularly toothed to not more than middle, sometimes distinctly lobed.

195

187

Type of section: Pyrethrum macrophyllum (W. and K.) Willd.

Note. This small section is, apparently, one of the most primitive
in the genus, which is indicated by the unique habit of the species of
this section as well as by their adaptation to shady forests and extremely
sporadic distribution. Beginning with H. Cassini (l.c.), it was often,
but without sufficient justification, combined in a single section or
genus with species of the section Xanthoglossa (DC.) Sch. Bip. of the
genus Tanacetum L.

1. P. macrophyllum (W. and K.) Willd. Sp. pl. III (1803) 2154;
DC. Prodr. VI, 58; Ldb. Fl. Ross. II, 553; Boiss. Fl. or. III, 346; Sosn.
in Tr. Tifl. Bot. Sada, XVII, 44; Grossh. Fl. Kavk. IV, 139 and Opred.
Rast. Kavk. 460; Stank. in Stank. and Tal. Opred. Rast. Evrop. Ch.
SSSR, 631.—Chrysanthemum macrophyllum Waldst. and Kit. Pl. rar.
Hung. I (1802) 97, t. 94, non Biehl. 1807.—Gymnocline leucocephala
Cass. in Dict. Sc. Nat. XX (1821) 120, nom. nov.—C. macrophylla
(W. and K.) Bluff and Fingerh. Fl. Germ. II (1825) 389.—Achillea
sambucifolia Desf. Cat. paris. (1829) 180.—Tanacetum macrophyllum
(W. and K.) Sch. Bip. Tanacet. (1844) 53.—Ic.: Waldst. and Kit. l.c.;
Rchb. Ic. Fl. Germ. XVI, t. 995; Hegi, Ill. Fl. VI, 2, f. 329; Jav. and
Csap. Ic. Fl. Hung. 522.—Exs.: Fl. exs. Austro-Hung. No. 659; FI.
Bohem. et Morav. exs. No. 395.

Perennial. Plants 50-150 cm high, with thick inclined rhizome,
usually covered with long and short (to papilliform) simple hairs, lacking
short nonflowering branches; stems solitary or few, erect, densely leafy,
branched only at apex (in inflorescence). Leaves green and glabrous
above, less often pubescent, dull green beneath, more densely pubescent,
weakly punctate-glandular; basal leaves withering early; cauline leaves
sessile or sub-sessile; lamina more or less lanceolate-ovate, pinnately-
parted up to winged rachis, lobes 4-8 on each side, more or less
lanceolate, 8-25 mm wide, acute, more or less duplicicodentate along
margin, sometimes pinnatilobate. Capitula (20)40—100(150), aggregated
in dense, compound corymbs, on 4-15 mm long peduncles. Involucre
4-6 mm in dia and 3.5-4.5 mm long, more or less crisped-hairy;
involucral bracts herbaceous, outer narrow-lanceolate, inner 1.5 times,
rarely 2 times as long, oblong-linear or linear; all bracts with rather
narrow, dark brown, membranous border, apex of inner bracts
appendiculate. Ligulate florets 5-10, white, with tube about 1 mm
long and roundish or reniform limb; 2-4 mm long and 2.5-3.5 mm
wide; corolla of tubular florets 1.8-2.4 mm long. Achenes 1.6—2.0 mm
long, with 5 longitudinal ribs, in ligulate florets more or less shifted
ventrally; corona 0.2-0.3 mm long, finely toothed. Flowering July to
August.

196

188

In forests, among shrubs, subalpine meadows at altitudes to 2,500
m.—European part: Reported by Stankov (op. cit.) for western re-
gions of Ukraine; Caucasus: Eastern Transcaucasia (western part),
Western Transcaucasia, Southern Transcaucasia (northern and western
parts), Dagestan (southern part). General distribution: Scandinavia
(rarely), Central Europe (sporadically), Balkans-Asia Minor. Described
from Hungary. Type in Vienna or Budapest.

Note. P. macrophyllum var. angustifolium Ldb. (FI. Ross. Il, 554),
described from Nordman’s collections from Georgia; according to
Mandenova (in Zam. po Sist. Geogr. Rast. XXII, 1961, p. 62) it is a
synonym of Achilles latiloba Ldb.

2. P. sorbifolium Boiss. Fl. or. III (1875) 343; Manden. in FI.
Gruzii. VII (1952) 360.

Perennial. Plants 50-150 cm high, with short inclined rhizome,
more or less sparsely covered with short (often papilliform), simple
hairs, lacking short non-flowering branches; stems solitary or few, erect,
densely leafy, branches only at apex (in inflorescence). Leaves green,
more or less pubescent to entirely glabrous, lacking distinct punctate
glandular hairs; basal leaves withering fast; cauline leaves up to 15-
20 cm long and 10-12 cm wide, lower rather long-petiolate, others
short-petiolate or subsessile, their lamina ovate or oblong, pinnately
cut or pinnately parted up to narrow-winged rachis, segments 3—6 on
each side, linear-lanceolate, 4-15 mm wide, acute, acutely
duplicicodentate. Capitula (5)10—30(40), aggregated in lax compound
corymbs, on 2-6 cm long peduncles. Involucre 5-8 mm in dia and
3.5-5.0 mm long, glabrous or subglabrous; involucral bracts coriaceous-
herbaceous, outer lanceolate more or less lacking membranous border,
inner 1.5—2 times as long, linear-oblong, with appendiculate, brownish
membranous border in upper part. Ligulate florets numerous, white,
with corolla tube 1.0—1.5 mm long and oblong-linear limb 8-11 mm
long and 2—3 mm wide; corolla of tubular florets 1.8-2.4 mm long,
achenes 2.2—2.5 mm long, with 5—8 longitudinal ribs, in ligulate florets
shifted ventrally; corona about 0.2 mm long, obtusely fine-toothed
along margin. Flowering June to July.

Forests, forest glades, subalpine meadows in upper mountain
zone.—Caucasus: Western Transcaucasia (Adzharian ASSR). General
distribution: Turkey. Described from Turkey. Type in Geneva.

Note. It occupies a fairly isolated position and was included by
Boissier under a different. section of the genus. In the structure of the
capitulum it comes closer to the species of the following section.

197

189

Section 2. Balsamita (Mill.) DC. Prodr. VI (1837) 62.—P. sect.
Eupyrethrum Sosn. subsect. Balsamita (Mill.) Sosn. in Tr. Tifl. Bot.
Sada, XVII (1915) 38.—Balsamita Mill. Gard. Dict. Abridg. ed. 4
(1754); Desf. in Act. Soc. Hist. Nat. Paris, I (1792) 1.—Tanacetum
sect. Balsamita (Mill.) Sch. Bip. Tanacet. (1844) 50.—Plants
herbaceous, 20-120 cm high, more or less covered with short, appressed
bifid hairs, usually mixed with simple hairs, with densely leafy stem
and weakly developed short nonflowering branches. Leaves entire, less
often lyrate. Capitula usually numerous, aggregated at stem apices in
dense or somewhat lax corymbs, very rarely solitary. Involucre narrow-
patelliform, 5-10 mm in dia; outer bracts 2/3, less often 1/2, as long
as inner. Receptacle weakly convex, almost flat. Corolla limb in ligulate
florets 4-8 mm long; corolla of tubular florets 2.0-2.5 mm, with slightly
and gradually broadened tube in upper half and triangular, weakly
recurved teeth. Achenes 2.0-2.5 mm long, with 5—8 longitudinal ribs,
in ligulate florets more or less shifted ventrally; corona 0.1—-0.3 mm
long, irregularly toothed or lobed to middle or almost to base.

Type of section: Pyrethrum majus (Desf.) Tzvel.

Note. A widely distributed cultivated species of this section, P.
majus, without ligulate pistillate florets, was, for long, sometimes
separated a separate genus, Balsamita Mill., and sometimes combined
with the genus Tanacetum L. However, as demonstrated even by De
Candolle, this species is so close in all characters to other species of
the section included in the genus Pyrethrum that there cannot be any
doubt about the affinity of this section to this genus. All species in this
section (about 5) are close ecogeographic races of the same genus and
distributed almost exclusively (except P. majus) in the countries of
Northwest Asia.

3. P. balsamita (L.) Willd. Sp. pl. II (1803) 2153; DC. Prodr.
VI, 63; Ldb. Fl. Ross. II, 558; Boiss. Fl. or. III, 345, excl. var.; Grossh.
Fl. Kavk. IV, 134 and Opred. Rast. Kavk. 460.—Chrysanthemum
balsamita L. Sp. pl. (1763) 1252.—C. balsamita var. radiatum Schmalh.
Fl. II (1897) 69.—Matricaria balsamita (L.) Desr. in Lam. Encycl. III
(1792) 733.—Tanacetum balsamitoides Sch. Bip. Tanacet. (1844) 51,
nom. nov.—Leucanthemum balsamita (L.) Over. Fl. S. Dakota (1932)
108.

Perennial. Plants 30-120 cm high, with more or less branched,
short creeping rhizome, usually dull green from soft and appressed
bifid (mixed with simple) hairs; stems few or solitary, erect, densely
leafy, more or less branched above. Leaves dull green from rather
dense pubescence, with numerous punctate-glandular hairs, entire, more
or less obtusely toothed along margin; basal leaves rather long petiolate

198

190

(petiole often as long as lamina) with oblong, more or less abruptly
narrowed lamina at base; cauline leaves short-petiolate or subsessile,
oval, uppermost strongly reduced, often entire. Capitula 10-60, on
rather long peduncles, in somewhat lax, compound, corymbs. Involucre
6-10 mm in dia, 3-4 mm long, more or less pubescent; involucral
bracts coriaceous-herbaceous, green or greenish-brown, outer lanceolate,
almost lacking border, inner 1.5, less often 2 times as long, linear-
lanceolate, with light colored or brownish membranous border in upper
part, apically broadened appendiculately. Ligulate florets 6—20, white,
with corolla tube about 1 mm long and limb 4—6 mm; corolla of tubular
florets about 2 mm long. Achenes 2.0—2.4 mm long, about 0.6-0.7 mm
wide, with (5)6—7(8) longitudinal ribs; corona about 0.2 mm long,
more or less irregularly toothed or lobed along margin. Flowering July
to August.

Stony slopes, scrubs, meadows up to upper mountain zone, often
cultivated and escapes, found along city streets, by roadsides, in parks
and kitchen gardens.—Caucasus: Transcaucasia, Talysh. General
distribution: Asia Minor, Armenia-Kurdistan, Iran. Described from
garden specimens of stock from north-eastern Turkey. Type in London.

Note. The more alpine specimens, usually more weakly pubescent
and with a darker membranous border on the involucral bracts and
which are transitional to the next species, are described as a separate
variety—Chrysanthemum balsamita var. viride Brodz. [in Tr. Bot. Sada
Yur’evsk. Univ. XIII (1912) 24] and the form C. balsamita f. glabrescens
Nabelek [in Publ. Fac. Sc. Univ. Masaryk, Brno, LII (1925) 22}.

In the countries of Northwestern Asia; a species with lyrate-
pinnatisect leaves, which is very close to this species and, apparently,
has evolved in cultivation, also is cultivated—Pyrethrum pinnatifidum
Willd. [Willd. Sp. pl. III (1803) 2154; P. multifidum (Desf.) DC. Prodr.
VI (1837) 63; Matricaria tanacetifolia Desr. in Lam. Encycl. III (1792)
733, non Pyrethrum tanacetifolium Boiss. 1875; Chrysanthemum
multifidum Desf. Cat. horti Paris. (1829) 170; C. glaucum Pers. Syn.
pl. II (1807) 461]. This species has not yet been reported from the
Caucasus, although its occurrence as a cultivated or introduced plant
is quite probable.

Economic Importance. Widely cultivated as an aromatic vegetable
in the countries of Northwest Asia and the Caucasus.

4. P. balsamitoides (Nabel.) Tzvel. comb. nova.—Chrysanthe-
mum balsamitoides Nabel. in Publ. Fac. Sc. Univ. Masaryk. Brno, LII
(1925) 22, t. V, fig. 2.—Ic.: Nabelek, ibid.

Perennial. Plants 20-50 cm high, with strongly branched creeping
rhizome, more or less covered with appressed bifid hairs to almost

191

glabrous; stems usually few, erect or ascending at base, densely leafy,
more or less branched above, or simple. Leaves green or dull green
from pubescence, often almost glabrous, with few but not always distinct
punctate-glandular hairs, entire, more or less toothed; basal leaves on
long (often as long as lamina or longer) petioles, with oblong or obovate
lamina gradually narrowed toward base; cauline leaves oval or ovate,
lower short-petiolate, upper sessile or subsessile. Capitula solitary or
2-—10(15), in somewhat lax racemose inflorescence. Involucre 8-10
mm in dia and 4—5 mm long, more or less pubescent to subglabrous;
involucral bracts coriaceous-herbaceous, usually dark brownish-violet,
outer lanceolate, with very narrow membranous border, inner not more
than 1.5 times as long, linear-oblong, with dark brown membranous
border appendiculately broadened at apex. Ligulate florets white, with
corolla tube about | mm long and limb 6-8 mm long; corolla of tubular
florets 1.8—2.0 mm long. Achenes 1.8—2.4 mm long and about 0.5 mm
wide with 5-7 longitudinal ribs; corona 0.1—0.3 mm long, more or less
lobed or irregularly toothed. Flowering July to August.

Stony slopes, meadows at altitudes higher than 2500 m.—Caucasus:
Southern Transcaucasia. General distribution: Armenia-Kurdistan.
Described from northeastern Turkey. Type in Brno (Czechoslovakia).

Note. A high-mountain, morphologically poorly isolated race, it is
related to the previous species through transitional forms. The most
typical form is found in the alpine meadows of Aragats Mountain.

5. P. majus (Desf.) Tzvel. comb. nova.—P. tanacetum DC. Prodr.
VI (1837) 63, nom. nov.—P. balsamita var. tanacetoides Boiss. Fl. or
III (1875) 346; Sosn. in Tr. Tifl. Bot. Sada, XVII, 39; Grossh. FI.
Kavk. IV, 134.—Tanacetum balsamita L. Sp. pl. (1753) 845; Klokov
in Vizn. Rosl. URSR (1950) 538.—Balsamita major Desf. in Act. Soc.
Hist. Nat. Paris. I (1792) 3, nom. nov.—B. vulgaris Willd. Sp. pl. II
(1803) 1802, nom. nov.—B. suaveolens Pers. Syn. pl. II (1807) 408,
nom. nov.—B. balsamita (L.) Rydb. in N. Am. Fl. XXXIV (1916)
238.—Chrysanthemum majus (Desf.) Aschers. Fl. Brandenb. (1864)
329.—C. balsamita auct. non L.: Schmalh. Fl. II, 69, excl. var.—
Leucanthemum balsamita (L.) Stank. in Stank. and Tal. Opred. Rast.
Evrop. Ch. SSSR (1949) 630, non Over. 1932.—Ic.: Rchb. Ic. FI.
Germ. XVI (1854) t. 995; Syreistsch. Ill. Fl. Mosk. Gub. III, 267;
Bonnier, Fl. Compl. France, Suisse et Belg. V, t. 292; Hegi, Ill. Fl. VI,
2, 599; Jav. and Csap. Ic. Fl. Hung. 524.

Plants 30-120 cm high, with more or less branched short-creeping
rhizome, usually covered with short, appressed bifid (mixed with simple)
hairs. Stems few or solitary, erect, densely leafy, usually branched
above. Leaves usually dull green due to pubescence, sometimes

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192

subglabrous, with numerous punctate glandular hairs, entire, more or
less toothed; basal leaves long-petiolate, with oblong, more or less
abruptly narrowed lamina near base; cauline leaves short-petiolate or
sessile, oval, uppermost strongly reduced. Capitula 10—60, on shorter
peduncles, in rather dense, compound, corymbs. Involucre 6-8 mm in
dia, 3-4 mm long, more or less hairy; involucral bracts coriaceous-
herbaceous, greenish; outer bracts lanceolate, usually lacking border,
inner 1.5, less often 2 times as long, linear-oblong, with light-colored,
apically apendiculately broadened, membranous border. Ligulate florets
entirely lacking; corolla of tubular florets about 2 mm long. Achenes
1.8—2.4 mm long, about 0.6 mm wide, with 5-8 longitudinal ribs and
0.1-0.3 mm long corona, more or less irregularly toothed along margin.
Flowering July to September.

Cultivated in parks and kitchen gardens, and often escapes.—
European part: Mostly more southern and western districts; Caucasus:
Occasionally cultivated and escapes; Soviet Central Asia: Occasion-
ally cultivated. General distribution: Europe (except Far North), North-
west Asia. Described from Southern Europe. Type in London.

Note. Apparently, a species that has appeared in cultivation, dif-
fering from the original species, P. balsamita, not only in the absence
of ligulate florets, but also in having a denser inflorescence.

Economic Importance. It is an aromatic vegetable, also used in
folk medicine and has insecticidal properties.

Section 3. Parthenium (Brig.) Tzvel. comb. nova.—Tanacetum
sect. Leucoglossa § 1. Microsperma Sch. Bip. Tanacet. (1844) 55.—
T. sect. Pyrethrum subsect. Parthenium Briquet in Burnat. Fl. Alp.
Marit. VI (1916) 119.—Herbaceous plant, 15-80 cm high, more or
less pubescent (to subglabrous), with densely leafy stem and more
weakly developed, short nonflowering shoots; leaves pinnatisect, all
rather long-petiolate. Capitula usually rather numerous, in lax or more
or less dense, sometimes irregular corymbs, less often solitary. Involucre
narrow-patelliform, 6-16 mm in dia; outer involucral bracts a half to
two-thirds as long as inner. Receptacle usually strongly convex,
conically hemispherical, rarely (in P. peucedanifolium) weakly convex.
Corolla limb in ligulate florets 2.5-15.0 mm long, sometimes ligulate
florets entirely lacking; corolla of tubular florets 1.2—2.0 mm long, .
with gradually slightly broadened tube in upper half and deltoid, weakly
recurved teeth. Achenes 1.0—1.8 mm long, with 5-10 longitudinal ribs,
in ligulate florets more or less ventrally shifted; corona 0.1-0.6 mm
long, more or less irregularly toothed along margin or divided almost
to base into 5-10 lobes of different shape and size.

Type of section: Pyrethrum parthenium (L.) Sm.

193

199 Plate VIII.
1 — Pyrethrum leptophyllum Stev., habit of plant, involucre, corolla of ligulate
floret, corolla of tubular floret, achene; 2 — P. Sangaricum Tzvel., habit of
plant, corolla of ligulate floret, corolla of tubular floret: achene.

203

194

Note. About 15 species of the section are distributed mainly in the
countries of Western Asia and the Caucasus. One species is widely
distributed in the mountainous regions of Southern Europe and a large
part of Asia, while another species (P. parthenium) is a common
ornamental plant.

Series 1. Peucedanifolia Tzvel.—Receptacle weakly convex, al-
most flat. Apical lobes of leaves linear or lanceolate.

6. P. peucedanifolium (Sosn.) Manden. in Zam. po Sist. i Geogr.
Rast. Bot. Inst. Akad. Nauk Gruz. SSR, XXII (1961) 59.—P.
?parthenifolium var. peucedanifolia Sosn. in Tr. Tifl. Bot. Sada, XVII
(1915) 35.—P. svanicum M. Pop. ex Manden. I. c. in syn.

Perennial. Plants 30-80 cm high, with shorter tap root, covered
with very short, appressed, simple hairs. Stems erect, densely leafy,
usually branched only at base and apex (in inflorescence), but with
short branches in axils of middle and upper cauline leaves. Leaves dull
green from rather dense pubescence, with numerous punctate glandular
hairs, all rather long petiolate; basal and lower cauline leaves withering
very early; lamina deltoid-ovate, pinnately cut upto more or less
wingless rachis, with 2-5 lobes on each side, partly entire, but usually
lobes more or less pinnately parted; apical lobes linear or lanceolate,
usually 1.0—2.5 mm wide, narrowed toward subobtuse or subacute apex;
upper cauline leaves smaller and weakly divided. Capitula 5-20 on
single stem, on 2—8(10) cm long peduncles, in lax, sometimes irregular
corymbs. Involucre 7-10 mm in dia, 3-4 mm long, rarely puberulent;
involucral bracts coriaceous-herbaceous, outer narrow-lanceolate,
lacking border, inner lanceolate-linear and linear, with appendiculately
broadened, light colored, membranous border at apex. Ligulate florets
white, with corolla tube about 1 mm long and limb 6-8 mm long;
corolla of tubular florets 1.5—2.0 mm long. Achenes 1.2-1.6 mm long,
with 5-8 oblong ribs and 0.3-0.6 mm long corona, divided to or almost
to base into 5-10 lanceolate or lanceolate-subulate lobes of different
shapes and sizes. Flowering July to August.

River gravel beds, scrubs, forest glades up to lower mountain
zone.—Caucasus: Western Transcaucasia (northern part). Endemic.
Described from northern Georgia. Type in Tbilisi.

Note. A unique species, close to the species of the previous sec-
tion in general habit and structure of the receptacle, differing from
them, however, by smaller achenes and the nature of the pubescence.

Series 2. Parthenifolia Sosn. in Tr. Tifl. Bot. Sada, XVII (1915)
32.—Receptacle strongly convex, conically hemispherical. Apical lobes
and segments of leaves oblong to almost semicircular.

204

195

7. P. parthenifolium Willd. Sp. pl. III (1803) 2156; MB. FI.
taur.-cauc. II, 325 and III, 579; DC. VI, 58; Ldb. Fl. Ross. II, 552;
Boiss. Fl. or. III, 344; O. and B. Fedtsch. Perech. Rast. Turk. IV, 187;
Grossh. Fl. Kavk. IV, 138 and Opred. Rast. Kavk. 462.—P. parthenium
var. parthenifolium (Willd.) Lipsky, Fl. Kavk. (1899) 349.—P.
parthenifolium var. latilobum Sosn. in Grossh. Fl. Kavk. IV (1934)
138, diagnosis in Russian.—P. latilobum (Sosn.) Sosn. in Zam. po
Sist. i Geogr. Rast. Inst. Bot. Akad. Nauk Gruz. SSR, XV (1949) 6;
Manden. in FI. Gruzii, VIII, 363, Plate 398.—Chrysanthemum
parthenifolium (Willd.) Pers. Syn. pl. II (1807) 462; B. Fedtsch. Rast.
Turk. 737.—C. parthenium var. parthenifolium (Willd.) Schmalh. FI.
II (1897) 69.—Tanacetum parthenifolium (Willd.) Sch. Bip. Tanacet.
(1844) 56.—Ic.: Rcehb. Ic. Fl. Germ. XVI (1854) t. 992; Manden. op.
cit—Exs.: A. Callier, Iter. taur.-tertium, No. 636; P. Sintenis, Iter
transcasp.-pers. No. 1727.

Perennial. Plants 15-70 cm high, with shorter tap root, more or
less puberulent to subglabrous. Stem erect, densely leafy, more or less
branched near base and above. Leaves green or dull green from more
dense pubescence with numerous punctate glandular hairs, all rather
long-petiolate; basal and lower cauline leaves 10-15 cm long, often
withering early; lamina of lower cauline leaves more or less ovate,
pinnately parted or pinnately cut, with 3-7 oblong or ovate, more or
less pinnatilobate lobes; lobes entire or more or less blunt-toothed,
subacute; upper cauline leaves strongly reduced and weakly divided.
Capitula (1)5—20(30), on rather long (to 10 cm) peduncles, in lax or
somewhat lax corymbs, or corymbose-panicles. Involucre 6-9 mm in
dia, 3-4 mm long, more or less pubescent to subglabrous. Involucral
bracts coriaceous-herbaceous, outer lanceolate, almost lacking border,
inner 1.5—2 times longer, oblong-linear, with narrow, light colored,
membranous border, appendiculately broadened at apex. Ligulate florets
8-20, white, with corolla tube 1-1.5 mm long, and limb 7-10 mm
long; corolla of tubular florets 1.2-1.8 mm long. Achenes 1.2—1.8 mm
long, about 0.4 mm wide, with 5-10 longitudinal ribs and about 0.2
mm long corona, more or less irregularly obtusely toothed along margin.
Flowering June to September.

More or less shaded stony slopes, rocks, especially in forests and
scrubs up to 2,000(2,500) m.—European part: Crimea (mountains);
Caucasus: All regions, but rarely in Southern Transcaucasia; Soviet
Central Asia: Dzhungaria-Tarbagatai, Tien Shan, Amu-Darya (moun-
tains), Pamiro-Alai Region (excluding Pamir), mountainous Turkmenia.
General distribution: Mediterranean Region, Balkans-Asia Minor,
Armenia-Kurdistan, Iran, Dzhungaria-Kashgaria. Described from gar-
den specimens of unknown origin (possibly from Caucasus?). Type in
Berlin.

20

nm

196

Note. The species P. latilobum (Sosn.) Sosn., described from Geor-
gia with slightly broader leaf lobes and, on the average, larger ca-
pitula, is morphologically so weakly isolated that, in my view, it is
more an ecological form than a separate ecogeographical race. P.
parthenifolium var. anthriscifolium Sosn. (Grossh. Fl. Kavk. 1V, 138),
with thin, somewhat more strongly divided lobes of the leaves, and P.
parthenifolium var. macrophyllum Sosn. (Grossh. Fl. Kavk. IV, 138),
with larger leaves, also have hardly any taxonomic significance.

Economic Importance. Sometimes cultivated as an ornamental plant
and rarely escapes, occurring also in the more northern regions.

8. P. parthenium (L.) Sm. FI. brit. II (1800-1804) 900; MB. FI.
taur.-cauc. III, 579; DC. Prodr. VI, 58; Ldb. Fl. Ross. II, 553; Boiss.
FI. or. III, 314; Grossh. Fl. Kavk. IV, 137 and Opred. Rast. Kavk. 462;
Manden. in Fl. Gruzii, VII, 360.—Matricaria parthenium L. Sp. pl.
(1753) 890.—M. latifolia Gilib. Fl. lithuan. I (1781) 220.—
Chrysanthemum parthenium (L.) Pers. Syn. pl. II (1807) 462; Schmalh.
Fl. II, 69, excl. var.—Tanacetum parthenium (L.) Sch. Bip. Tanacet.
(1844) 55.—Leucanthemum parthenium (L.) Gren. and Godr. FI. Fr. II
(1850) 145.—Parthenium matricaria Gesn. ex Rupr. Fl. ingr. I (1860)
583.—Ic.: Rchb. Ic. Fl. Germ. XVI (1854) t. 992; B. Fedtsch. and
Fler. Fl. Evrop. Ross. 975, Fig. 976; Hegi, Ill. Fl. VI, 2, fig. 330 and
33.13

Perennial. Plants 15-50 cm high, with shorter tap root, more or
less puberulent to subglabrous. Stem erect, densely leafy, more or less
branched at base and above. Leaves as in previous species, but usually
yellowish-green and, on the average, less hairy. Capitula (1)5—20(30),
on rather long (to 8 cm) peduncles, in rather dense corymbs. Involucre
6-8 mm in dia, 3-4 mm long, more or less pubescent, subglabrous;
involucral bracts coriaceous-herbaceous, outer lanceolate, more or less
lacking border, inner 1.5—2 times as long, oblong linear, with narrow,
light colored, membranous border, more or less divided at apex. Ligulate
florets 5-15, white, with corolla tube 1 mm long and limb 2.5—6.0 mm
long, sometimes entirely lacking; corolla of tubular florets 1.2—-1.8 mm
long. Achenes 1.2-1.5 mm long and about 0.4 mm wide with 5-8
longitudinal ribs; corona about 0.2 mm long, more or less irregularly

' toothed along margin, strongly reduced in ligulate florets. Flowering

July to August.

Often cultivated and frequently as escape in gardens, parks, forests,
and in weedy places.—European part: Mainly in western and more
southern regions; Caucasus: Rarely, mostly near large cities; Eastern
and Western Siberia: More southern regions; Far East: Rarely in
southern regions; Soviet Central Asia: Rarely, near large cities. Gen-

206

197

eral distribution: Western Europe, North America, South America
(Chile), Northwest Asia; everywhere as an introduced or escaped plant.
Described from Western Europe. Type in London.

Note. Apparently, it originated as a result of the prolonged
cultivation of P. parthenifolium as an edging plant (where the
inflorescences have no ornamental value) and is associated with it
through “transitional” forms. However, there is no doubt about its
close relationship with the wild species of the P. grossheimii type,
in which ligulate florets often tend to disappear even in the wild
State.

Economic Importance. Often cultivated as an ornamental border,
mainly the variety with yellowish-green leaves—var. aureum hort. There
are a few varieties with very finely divided leaves, for example, var.
laciniatum hort. and var. selaginoides hort. However, the cultivated
form with “double” capitula is also propagated for its flowers. The
following forms or varieties differ in the degree of reduction of the
ligulate florets: the typical one—var. longiradiatum Sch. Bip. (op. cit.
55) with strongly reduced ligulate florets—var. breviradiatum Sch.;
Bip. (op. cit. 55); and entirely without ligulate florets—var. flosculosum
DC. (op. cit. 58 = var. discoideum Sch. Bip. op. cit. 55). In more
northern regions, it freezes in winter and behaves as an annual. It is
used in folk medicine.

9. P. glanduliferum Somm. and Lev. Decas. Compos. nov. cauc.
(1895) 87; Sosn. in Zhurn. Russk. Bot. Obshch. XIV (1929) 83; Grossh.
Fl. Kavk. IV, 134 and Opred. Rast. Kavk. 460; Manden. in Fl. Gruzii,
VIII, 367.—P. buschianum Sosn. in Zhurn. Russk. Bot. Obshch. XIV
(1929) 90; Grossh. Fl. Kavk. IV, 137 and Opred. Rast. Kavk. 462;
Manden. op. cit. 364.—Ic.: Dechy, Kaukasus, III (1907) t. XXV (type
specimen of species); Kolakovskii, Fl. Abkh. IV (1949) Plate XXIV.

Perennial. Plant 15—50 cm high, with shorter tap root, more or less
puberulent, often subglabrous. Stem erect, densely leafy, more or less
branched near base and above, less often simple. Leaves green or dull
green from more dense pubescence, with numerous punctate-glandular
hairs, all rather long-petiolate; lamina of lower and middle cauline
leaves oblong to broadly ovate, pinnately cut almost to wingless rachis,
with 5-9 segments, apical segments more or less connate at base,
oblong, crenate or irregularly subacutely toothed or more or less
pinnately lobed, sometimes partly pinnately parted, with more or less
toothed lobes; upper cauline leaves strongly reduced and weakly
divided. Capitula solitary or 2—15(20) on single stem, on rather long
(up to 10-15 cm) peduncles in lax, sometimes irregular, corymbs.
Involucre 9-16 mm in dia, 3-4 mm long, more or less pubescent to

198

glabrous; involucral bracts coriaceous-herbaceous, outer lanceolate or
narrow-lanceolate, almost lacking border, inner 1.5, less often 2 times
as long, linear and oblong-linear; with narrow light colored membranous
border, more or less broadened at apex. Ligulate florets numerous,
white, with corolla tube about 1.0-1.5 mm long, and limb 9-15 mm
long; corolla of tubular florets 1.5—2.0 mm long. Achenes 1.4—1.8 mm
long, about 0.5 mm wide, with 5-10 longitudinal ribs; corona 0.1—0.2
mm long, more or less irregularly obtusely toothed along margin.
Flowering July to September.

In meadows, on stony slopes, rocks at 1,500—3,000 m.—Caucasus:
Ciscaucasia, Western Transcaucasia. Endemic. Described from Glavny
[Main] Caucasus Range. Type in Florence.

Note. It is a very closely related and a more high-mountain race
in relation to P. parthenifolium with which it is linked through quite
numerous transitional forms.

Apparently, the species P. glanduliferum and P. buschianum should
be considered full synonyms. The characters, by which they have usually
been distinguished (degree of pubescence and number of capitula), do
not exhibit any clear ecological or geographical adaptation in the
extensive material.

10. P. grossheimii Sosn. in Beih. Bot. Centralbl. XLIV, 2 (1928)
243; Grossh. Fl. Kavk. IV, 137 and Opred. Rast. Kavk. 461.—?P.
parthenifolium var. discoideum Boiss. Fl. or. III (1875) 345.

Perennial. Plant 15—40 cm high, with shorter tap root, more or less
pubescent to subglabrous. Stem erect or ascending at base, densely
leafy, branched usually at base and above middle. Leaves green, rather
thin, less often pubescent, with numerous punctate-glandular hairs, all
rather long-petiolate; lamina of middle and lower cauline leaves more
or less ovate, pinnately cut to almost wingless rachis, with 2-5 segments
on each side, ovate to oblong, pinnately parted into more or less oblong,
sometimes shallow-pinnately lobed lobes, on the average, broader than
in P. sevanense. Capitula 5—20(30), on rather long (to 8 cm) slender
peduncles, in lax corymbose-panicles. Involucre 5.0—7.5 mm in dia,
2.5-3.0 mm long, less often pubescent; involucral bracts coriaceous-
herbaceous, outer narrow-lanceolate, almost lacking border, inner 1.5,
less often 2 times as long, linear or oblong linear, with narrow but
apically somewhat broadened membranous border. Ligulate florets 1-
15, sometimes entirely lacking, white, with corolla tube about 1 mm
long and limb 4.0-5.5 mm long; corolla of tubular florets 1.2-1.5 mm
long. Achenes 1-1.3 mm long, about 0.4 mm wide, with 5-10
longitudinal ribs; corona about 0.1 mm long, subobtusely toothed along
margin. Flowering May to June.

20

oo

199

Rocks, stony slopes to 2,000 m.—Caucasus: Southern Trans-
caucasia (Nakhichevan ASSR); Soviet Central Asia: mountainous
Turkmenia (rarely). General distribution: Armenia-Kurdistan, Iran
(northern part). Described from the Nakhichevan ASSR. Type in Tbilisi.

Note. In the shape of the leaves, it occupies a somewhat interme-
diate position between P. sevanense and P. parthenifolium, but in the
structure of the capitula it is close to P. parthenium.

11. P. sevanense Sosn. in Grossh. Fl. Kavk. IV (1934) 317 and
Opred. Rast. Kavk. 462.—P. palmatum (Vent.?) C. Koch in Linnaea,
XXIV (1851) 331, sphalm. (=Chrysanthemum praealtum Vent.).—P.
parthenifolium var. canescens Boiss. Fl. or. III (1875) 344 p. p.—P.
parthenifolium f. divaricata Sosn. in Tr. Tifl. Bot. Sada, XVII (1915)
36.—P. parthenifolium var. sevangensis Sosn. in Grossh. in Zhurn.
Russk. Bot. Obshch. XIV, 3 (1930) 314.—P. niveum var. divaricatum
(Sosn.) Sosn. in Grossh. Fl. Kavk. IV (1934) 137.—P. divaricatum
(Sosn.) Sosn. in Zam. po Sist. i Geogr. Rast. Bot. Inst. Akad. Nauk
Gruz. SSR, XV (1949) 5; Manden. in Fl. Gruzii, VIII (1952) 364.—
?Chrysanthemum praealtum Vent. Hort. Cels. (1800) 43, t. 43.

Perennial. Plant 15—60 cm high, with shorter tap root, more or less
puberulent. Stem erect or ascending at base, densely leafy, more or
less branched at base and above middle. Leaves dull green from rather
dense pubescence; less often subglabrous above, with numerous
punctate-glandular hairs, all rather long-petiolate; basal and lower
cauline leaves up to 15(20) cm long, 4 cm wide; lamina of lower and
middle cauline leaves more or less ovate, pinnately cut to almost
wingless rachis; leaf segments 2-5 on each side, ovate to oblong-
ovate, pinnately parted up to narrow-winged axis in more or less oblong
lobes, often in turn pinnatilobate or pinnatipartite; apical lobes ovate
to oblong, obtuse or subobtuse; upper cauline leaves strongly reduced
and weakly divided. Capitula (1)5—20(30), on rather long (up to 8-10
cm) peduncles, aggregated in lax corymbose-racemes or corymbose-
panicles. Involucre 7-10 mm in dia, 3.0-4.5 mm long, more or less
pubescent; involucral bracts coriaceous-herbaceous, outer few, narrow-
lanceolate, inner 1.5, less often almost 2 times as long, linear or oblong-
linear, with narrow but apically more or less broadened, light-colored,
membranous border. Ligulate florets numerous, white, with corolla
tube 1.0-1.2 mm long, and limb; 7-10 mm long; corolla of tubular
florets 1.4—1.8 mm long. Achenes 1.0—1.5 mm long, about 0.3-0.4 mm
wide with 5-10 longitudinal ribs and more or less subobtusely
irregularly toothed 0.1—0.3 mm long corona, usually reduced in ligulate
florets. Flowering June to August.

20

No)

200

Stony slopes, rocks at altitudes up to 2,500 m.—Caucasus: East- |
ern Transcaucasia, Southern Transcaucasia, Dagestan (southern part);
Soviet Central Asia: mountainous Turkmenia. General distribution:
Armenia-Kurdistan, Iran (northwestern part). Described from vicinity
of Lake Sevan in the Armenian SSR. Type in Tbilisi.

Note. It is very probable that the priority name of this species is
Pyrethrum praealtum (Vent.) Tzvel. comb. nova (=Chrysanthemum
praealtum Vent. |. c.). This latter species was described from garden
specimens from northern Iran and, like P. sevanense, differs from P.
parthenifolium by a very strong division of the leaves. It is to this
species that Koch (1. c.) referred his specimens from Armenia, which
undoubtedly belong to P. sevanense (the species was named by him
not “praealtum” but “palmatum” only as the result of a printing error).
However, the specimen of C. praealtum Vent. illustrated by the author
of this species as though it were the type, has broader leaf lobes in
comparison with the Caucasian specimens (possibly, as a result of
cultivation?). Hence, so far it has not been possible to establish its
complete identity with P. sevanense. There is hardly any justification
for separating the form described from Aragats Mountains, which differs
only on the average from P. sevanense by larger capitula on longer
peduncles, as an independent ecogeographical race. The Dagestan
specimens of P. divaricatum are somewhat more distinct from P.
sevanense, which are quite close to P. glanduliferum, but differ from
the former species by smaller capitula and the shape of the leaves.
Most recently, they have been separated by Mandenova as an
independent species: P. demetrii Manden. [Zam. po Sist. i Geogr. Rast.
Bot. Inst. Akad. Nauk Gruz. SSR, XXII (1961) 60]. The type of this
species: Dagestan, source of Avarian Koisu, Barbar Mountain, shale
screes, A. Dolukhanova. 7.VIII.1937. P. divaricatum var. monocephala
(Sosn.) Sosn. |. c. (1949) 6 = P. parthenifolium f. monocephala Sosn.,
established by D.I. Sosnowsky [in Tr. Tifl. Bot. Sada, XVII (1915) 35]
with solitary capitula, in my view has no taxonomic significance.

12. P. fruticulosum Biehl. Pl. nov. herb. Spreng. (1807) 37.—P.
niveum Lag. Elench. hort. Madrit. (1805?) nom. nud. and Gen. et sp.
nov. (1816) 30; DC. Prodr. VI, 57; Ldb. Fl. Ross. II, 552; Sosn. in
Zhurn. Russk. Bot. Obshch. XIV (1929) 93; Grossh. Fl. Kavk. IV, 137
and Opred. Rast. Kavk. 462; Manden. in Fl. Gruzii, VIII, 367.—P.
argenteum Adam ex Hoffm. Hort. Mosq. (1808) 34, nom. nud. non
Boiss. 1875.—P. pulverulentum Willd. Enum. hort Berol. II (1809)
906, non Lag. 1805.—P. parthenifolium var. pulverulenta (Willd.)
Karel. in Bull. Soc. Nat. Mosc. XII (1839) 158.—P. parthenifolium
var. canescens Boiss. Fl. or. III (1875) 344 p. p.—Matricaria

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201

pulverulenta (Willd.) Poir. in Lam. Encycl. meth. Suppl. HI (1814)
605.—Tanacetum niveum (Lag.) Sch. Bip. Tanacet. (1844) 56.

Perennial. Plant 15-60 cm high, with rather short tap root, more
or less canescent. Stem erect or ascending near base, densely leafy,
more or less branched mainly near base and above middle. Leaves as
in previous species, but more or less canescent. Capitula (1)5—20(30),
on rather long (to 10 cm) peduncles, in lax corymbs or corymbose-
panicles. Involucre 7-10 mm in dia and 3.0-4.5 mm long, more or less
pubescent; involucral bracts coriaceous-herbaceous, outer narrow-
lanceolate, inner 1.5, less often almost 2 times as long, linear or oblong-
linear, with narrow but apically more or less broadened, light colored,
membranous border. Ligulate florets numerous, white, with corolla
tube 1.0—-1.2 mm long and limb 7-10 mm long; corolla of tubular
florets 1.4—1.8 mm long. Achenes 1.0-1.5 mm long and about 0.3-0.4
mm wide, with 5-10 longitudinal ribs and more or less irregularly
toothed 0.1-0.3 mm long corona, more or less reduced in ligulate
florets. Flowering June to August.

On stony slopes, rocks at 1,000—2,500 m.—Caucasus: Ciscaucasia
(eastern part), Dagestan, Eastern Transcaucasia, Western Transcaucasia
(southern part?), Southern Transcaucasia (Aragats Mountain), Talysh.
General distribution: Armenia-Kurdistan. Described from Caucasus.
Type in Halle (Germany).

Note. This species is very close to the previous one and related to
it through transitional forms, but, apparently, all the same, forming a
distinct ecogeographical race, adapted mainly to northeastern part of
the Lesser Caucasus and eastern part of the Greater Caucasus. A few
specimens from the Adzharo-Imeretia Range, possibly, belong to a
separate broader-leaved race closer to P. kubense.

13. P. kubense Grossh. in Tr. Azerb. Otd. Zakavk. Fil. Akad.
Nauk SSSR, Sekt. Bot. 1 (1933) 58; Grossh. Fl. Kavk. IV, 137 and
Opred. Rast. Kavk. 462.

Perennial. Plant 20-70 cm high, with rather short tap root, canescent
(almost tomentose). Stem erect or ascending at base, densely leafy,
usually branched near base and above middle. Leaves canescent, with
indistinct punctate-glandular hairs, all rather long-petiolate; lamina of
lower and middle cauline leaves ovate or oblong-ovate, pinnately cut
almost to wingless rachis; leaf segments 2-5 on each side, oblong to
ovate, pinnatilobate or pinnatipartite; lobes and segments oblong-ovate
to almost semicircular, obtuse or subobtuse, sometimes crenate or
shallow-lobed; upper cauline leaves reduced, weakly divided. Capitula
(1)5—15(25), on long (to 15 cm) peduncles, in lax corymbose-racemes
or corymbose-panicles. Involucre 9-11 mm in dia, 3.5—4.5 mm long,

21

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202

more or less sparsely tomentose; involucral bracts coriaceous-herba-
ceous, outer narrow-lanceolate, almost lacking border, inner linear or
oblong-linear, with narrow but apically more or less broadened light-
colored, membranous border. Ligulate florets numerous, white, with
corolla tube about 1 mm long, and limb 7-10 mm long; corolla of
tubular florets 1.5—2.0 mm long. Achenes 1.2—1.8 mm long, about 0.5
mm wide, with 5-10 longitudinal ribs and more or less irregularly
toothed 0.1—0.2 mm long corona. Flowering June to August.

Rocks, stony slopes in upper mountain zone.—Caucasus: Dagestan
(southern part), Eastern Transcaucasia (northeastern part), Talysh
(reported by Grossheim). Endemic. Described from Kuba District of
Azerbaidzhan SSR. Type in Baku.

Note. Related to the previous species through transitional forms
but, all the same, apparently forming a unique ecogeographical race
differing from P. fruticulosum mainly by broader leaf lobes and a
denser pubescence.

Section 4. Leptanthemum Tzvel. sect. nova in Addenda, XXV,
871.—Plants herbaceous, 15-50 cm high, usually covered with
appressed bifid and simple hairs, usually mixed with shorter glandular
hairs, with less densely leafy stem and well developed, short,
nonflowering shoots. Leaves twice pinnately cut. Capitula usually rather
numerous, (1)4—25(40), in very lax corymbose or corymbose-paniculate
inflorescence. Involucre patelliform, 6-11 mm in dia; outer involucral
bracts 1/2—2/3, as long as inner. Receptacle weakly convex, almost
flat. Corolla limb in ligulate florets broadly oval, 5-7 mm long; corolla
of tubular florets 2.2-2.6 mm long, with weakly and gradually
broadened tube in upper half and deltoid, weakly recurved teeth.
Achenes 1.5—2.0 mm long, with (4)5(7) longitudinal ribs, in ligulate
florets shifted ventrally; corona about 0.2 mm long, usually irregularly
crenate or lobate up to middle.

A monotypic section.

Note. The only species of this section occupies a distinct position
in the genus and shows some resemblance to the species of the two
previous sections in capitulum structure. Its strong and, apparently,
not incidental resemblance to the species of section Tanacetopsis Tzvel.
of the genus Cancrinia Kar. and Kir. should also be noted.

14. P. leptophyllum Stev. in MB. FI. taur.-cauc. HI (1819) 580;
DC. Prodr. VI, 60; Ldb., Fl. Ross. II, 553; Boiss. FI. or. III, 341; Sosn.
in Tr. Tifl. Bot. Sada, XVII, 20; Grossh. Fl. Kavk. IV, 136 and Opred.
Rast. Kavk. 461.—Tanacetum leptophyllum (Stev.) Sch. Bip. Tanacet.
(1844) 48.

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203

Perennial. Plant 15-50 cm high, with creeping, more or less
branched rhizome, usually covered with adherent bifid and appressed
hairs, together with short glandular hairs. Stem erect, less densely
leafy, strongly branched from middle or slightly lower. Leaves usually
dull green from sparse (sometimes weak) pubescence, with numerous,
but not always conspicuous, punctate-glandular hairs; basal leaves up
to 15-20 cm long and 3 mm wide, on long, basally thickened petiole,
with twice or thrice pinnately cut, oblong or broad linear lamina; apical
lobes and segments broadly lanceolate to almost semicircular, to 1 mm
wide, with short cartilaginous cusp; cauline leaves reduced, lower short-
petiolate, upper sessile. Capitula (1)4—25(40) on 1.5-15.0 cm long
peduncles, in very lax, broadly spreading, corymbs or corymbose-
panicles. Involucre 6-11 mm in dia, 3-4 mm long, more or less
pubescent or glabrous; involucral bracts coriaceous-herbaceous, outer
narrow-lanceolate, usually lacking border, inner 1.5, less often 2 times
as long, linear-oblong, in upper part with narrow but apically
appendiculately broadened light colored or brownish membranous
border. Ligulate florets white, with strongly compressed corolla tube
1.0-1.5 mm long, and limb 5-7 mm long; corolla of tubular florets
2.2-2.6 mm long. Achenes 1.5-2.0 mm long and 0.5—-0.6 mm wide,
usually with 5 longitudinal ribs and irregularly crenate or lobate, about
0.2 mm long corona. Flowering July to August. (Plate VIII, Fig. 1.)

Stony slopes, screes, rocks at altitudes up to 2,500 m.—Caucasus:
Dagestan, Eastern Transcaucasia (northern part and Shakhdag Range).
Endemic. Described from southern Dagestan. Type in Helsinki; isotype
in Leningrad.

Section 5. Balsamitopsis Tzvel. sect. nova in Addenda, XXV,
871.—Plants herbaceous, 30-70 cm high, densely covered with strongly
flattened, appressed, bifid hairs, with weakly leafy stem and well
developed, short, nonflowering shoots. Basal leaves entire, cauline lyrate
to pinnately parted; all leaves more or less long-petiolate. Capitula
solitary or 2-6 on single stem, and then aggregated in lax corymbose-
racemes. Involucre narrow-patelliform, 8-15 mm in dia. Receptacle
weakly convex, almost flat. Corolla limb in ligulate florets oblong-
linear, 9-17 mm long; corolla of tubular florets about 2.5 mm long.
Achenes 2.5—2.8 mm long with 5-8 longitudinal ribs; corona 0.3-0.4
mm long, irregularly crenate.

A monotypic section.

Note. The only species of the section, known so far only from a
single locality (“Mramornaya Mountain”), does not show a clear affin-
ity with any of the known species from Soviet Central Asia and China.
Very likely it is closest to the species of section Balsamita (Mill.) DC.

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204

However, considering the large disjunction in their ranges and a series
of significant morphological differences, I prefer to include it under a
separate monotypic section.

15. P. kelleri (Kryl. and Plotn.) Krasch. in Kryl. Fl. Zap. Sib. XI
(1949) 2749.—Chrysanthemum kelleri Kryl. and Plotn. in Sb.
“Dvadtsat’ Pyat’ Let Nauchno-Pedag. i Obshch. Deyat. B.A. Kellera”*
(1931) 3.

Perennial. Plants 40-70 cm high, with creeping, more or less
branched rhizome, densely covered with strongly flattened, bifid hairs.
Stems solitary or few, erect or ascending at base, sparsely leafy, simple
or more or less branched above. Young leaves silver-grayish from
dense pubescence, older dull green, slightly sericeous, with
inconspicuous, punctate glandular hairs; basal leaves 20-25 cm long,
6-7 cm wide, on long (usually several times as long as lamina) petioles,
with entire obovate or rotund-elliptical lamina, broadly cuneate at base,
irregularly crenate along margin and near obtuse apex; cauline leaves
few (3-7), short-petiolate, with lyrate-pinnatisect lamina to 4 cm wide;
apical lobe large, broadly oval, crenate, lateral lobes small, 1-4 on
each side. Capitula solitary or 2-6 on single stem, on to 20 cm-long
peduncles, in lax corymbose-racemes. Involucre 8-15 mm in dia,
glabrous; outer involucral bracts lanceolate, inner oblong-obovate,
mainly in upper part with narrow, apically more or less broadened,
light-colored, membranous border. Ligulate florets white, limb 9-17
mm long and 3.0-3.5 mm wide. Achenes 2.5-2.8 mm long, 1 mm
wide, with 5—8 longitudinal ribs and crenate, 0.3—0.4 mm long corona.
Flowering June to July.

Dry stony slopes in lower mountain zone (at altitudes of about
1,000 m).—Western Siberia: Altai (southern part: Mramornaya
Mountain). Endemic. Described from Altai. Type in Tomsk.

Note. Unfortunately, I did not see the flowering specimens of this
unique species preserved in Tomsk.

Section 6. Cinerariifolia (Heyw.) Tzvel. comb. nova.—Tanacetum
sect. Pyrethrum subsect. Cinerariifolia Heywood in Anal. Inst. Bot.
Cavanill. Madrid, XII, 2 (1953) 325.—Plants herbaceous, 15-45 cm
high, densely covered with strongly flattened appressed, bifid hairs,
with more sparsely leafy stems and well developed, short, nonflowering
shoots. Leaves more or less pinnately cut, all usually long-petiolate.
Capitula solitary or 2-6 on single stem, and then aggregated in very

*“Twenty-five years of Scientific Pedagogic and Social Activity of B.A. Keller’ —
Translator.

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205

lax and irregular corymbs. Involucre 12-18 mm long, narrow-patelli-
form; outer involucral bracts a half to two-thirds as long as inner.
Receptacle weakly pubescent; corolla limb of ligulate florets oblong
or oblong-linear, 8-16 mm long; corolla of tubular florets 2.5-3.0
mm-long, with tube slightly broadened in upper half and deltoid, weakly
recurved teeth. Achenes 2.5—3.2 mm long, with 5—7 longitudinal ribs,
in ligulate florets more or less shifted ventrally; corona 0.6—1.0 mm
long, finely and irregularly toothed along margin.

Type of section: Pyrethrum cinerariifolium Trev.

Note. A section of relatively few species that are adapted to the
Mediterranean mountain regions.

16. P. cinerariifolium Trev. Ind. Sem. Hort. Vratisl. App. II (1820)
2 and in Nov. Act. Nat. Cur. XIII (1826) 204; DC. Prodr. VI, 55;
Grossh. Opred. Rast. Kavk. 461; Stankov in Stank. and Tal. Opred.
Rast. Evrop. Ch. SSSR, 631; Klok. in Vizn. Rosl. URSR, 539.—
Chrysanthemum turreanum Vis. Strip. Dalm. (1826) 19.—C.
cinerariifolium (Trev.) Vis. Fl. Dalm. II (1847) 88, t. 8.—Tanacetum
cinerariifolium (Trev.) Sch. Bip. Tanacet. (1844) 58.—Ic.: Vis. op.
cit.; Rchb. Ic. Fl. Germ. XVI (1854) t. 993; Hegi, Ill. Fl. VI, 2, f. 312;
Jav. and Csap. Ic. Fl. Hung. 524; Zemlinskii, Lekarstv. Rast. SSSR,
ed. 2, 230 and 231.—Exs.: Fl. Ital. exs. No. 179; Fl. exs. Austro-
Hung. No. 239.

Perennial. Plants 15-45 cm high, with short inclined rhizome,
silvery grayish from dense pubescence of strongly flattened, appressed,
bifid hairs; stems solitary or few, erect, more sparsely leafy, simple or
more or less branched (almost from base). Leaves silver-grayish from
dense pubescence, with numerous punctate-glandular hairs; basal leaves
to 10(20) cm long, long-petiolate, petioles sometimes exceeding lamina;
lamina ovate or oblong, pinnately cut up to usually wingless rachis
into cuneate segments near base, in turn pinnately or palmately parted;
apical segments and lobes linear to oblong-ovate, subobtuse or subacute;
cauline leaves smaller and weakly divided, short-petiolate. Capitula
solitary or 2—6 on single stem, and then aggregated in lax and irregular
corymbs. Involucre 12-18 mm in dia, 6-7 mm long, more or less
appressed-hairy; involucral bracts coriaceous-herbaceous, outer broad
lanceolate, subacute, lacking border, inner 1.5—2 times as long, oblong-
linear, with rather wide, light-colored, membranous border,
appendiculately broadened at apex. Ligulate florets white, with corolla
tube about 1.5 mm long and limb 8-16 mm long; corolla of tubular
florets 2.5-3.0 mm long. Achenes 2.5—3.2 mm long and 0.7-1.0 mm
wide, with 5—7 longitudinal ribs and 0.6—1.0 mm long finely toothed
corona. Flowering June to August.

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Cultivated as a medicinal (insecticidal) plant, occasionally found
as introduced or escaped plant.—European part: Southern regions;
Caucasus; Soviet Central Asia. General distribution: Balkan Peninsula
(Yugoslavia, Albania, Greece). Described from Yugoslavia. Type in
Bonn or Bratislava.

Economic Importance. It contains a high percentage of the com-
plex ester, pyrethrin, which forms the base for the “pyrethrum” insec-
ticide preparations used as solutions, powders, emulsions, etc. The
species has long been cultivated in many countries. Details about its
cultivation and uses can be found in the above cited book of Zemlinsky
(pp. 230-232), as well as in many brochures and articles by other
authors.

Section 7. Pyrethrellum Tzvel. sect. nova in Addenda, XXV,
872.—Plants herbaceous, 5-80 cm high, sometimes covered with short
bifid and simple or only simple hairs, usually more or less glabrous,
with more sparsely leafy stem and not always developed, short,
nonflowering shoots. Leaves more or less pinnately cut, lower petiolate,
upper sessile or short-petiolate. Capitula solitary, less often 2—10(40)
on single stem and then in very lax, irregular, corymbs. Involucre
narrow-patelliform, 8-20 mm in dia; outer involucral bracts not less
than 2/3 as long as inner. Receptacle usually strongly convex,
hemispherical, less often (in P. silaifolium) weakly convex. Limb of
ligulate florets oblong to linear, 7-25 mm long; corolla of tubular
florets 2.2-3.2 mm long, with slightly weakly and gradually broadened
tube in upper half and deltoid or lanceolate-deltoid, weakly recurved
teeth. Achenes 1.8-3.2 mm long, with 5-8 longitudinal ribs, more or
less shifted ventrally in ligulate florets; corona 0.1—0.5 mm long, less
often (P. komarovii) 0.5-1.3 mm long, usually irregularly toothed or
lobed to middle, less often almost to base.

Type of section: Pyrethrum coccineum (Willd.) Worosch.

Note. This Caucasian and northwest Asian section, closely related
to the previous section, shows some affinity, on the one hand, with the
species of section Parthenium through P. punctatum, and, on the other,
with some species of section Richteria (especially P. pulchellum) and
the genus Tripleurospermum Sch. Bip. through P. daghestanicum and
P. tricholobum. The generic affinity of P. tricholobum and P. komarovii,
morphologically very similar to the perennial Caucasian species of the
genus Tripleurospermum, cannot at present be established definitely
due to the absence of mature achenes. I have included both these
species (following Sosnowsky) in the genus Pyrethrum (section
Pyrethrellum) only on the basis of secondary, often insignificant
characters, and to a great extent only tentatively.

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Series 1. Stolonifera Sosn. in Tr. Tifl. Bot. Sada, XVII (1915)
27.—Plants with rooting stoloniferous branches, glabrous or with
isolated, simple papilliform hairs; stems densely leafy; leaves pinnately
parted, with broad apical lobes; receptacle strongly convex; ligulate
florets white; corona of achenes about 0.3 mm long.

17. P. punctatum (Desr.) Bordz. ex Sosn. in Zhurn. Russk. Bot.
Obshch. XIV (1929) 83; Grossh. Fl. Kavk. IV, 134 and Opred. Rast.
Kavk. 460; Manden. in Fl. Gruzii, VIII, 356.—P. palustre Willd. Sp.
pl. III (1803) 2154; DC. Prodr. VI, 54; Boiss. Fl. or. III, 341; Sosn.
in Tr. Tifl. Bot. Sada, XVII, 27.—P. jacobeiforme C. Koch, in Linnaea,
XXIV (1851) 331.—P. lilae Bordz. in Prot. Sobr. Kievsk. Obshch.
Estestv. 3, XI (1907) 27 and Fl. cauc. exs. No. 196.—Matricaria
punctata Desr. in Lam. Encycl. III (1792) 732.—Chrysanthemum
palustre (Willd.) Pers. Pers. Syn. pl. II (1807) 461.—C. punctatum
(Desr.). Bordz. in Zap. Kievsk. Obshch. Estestv. XXV (1915) 122.—
Tanacetum palustre (Willd.) Sch. Bip. Tanacet. (1844) 49.—Exs.: PI.
or. exs. No. 172; Fl. cauc. exs. No. 196; Herb. Fl. Cauc. fasc. XI, No.
54.

Perennial. Plants 20-80 cm high, with short inclined rhizome and
rooting stoloniferous shoots, glabrous or with isolated, simple,
papilliform hairs. Stem erect, densely leafy, ribbed, simple or more or
less branched above (sometimes almost from base). Leaves glabrous
or subglabrous; with numerous punctate-glandular hairs; basal and lower
cauline leaves withering early, rather long-petiolate; middle cauline
leaves to 10(14) cm long, 3(5) cm wide, sessile, oblong to oblong-
linear, pinnately parted up to narrow-winged rachis, with 4-11 lobes
on each side, usually narrowed toward base, oblong or oblong-linear,
3-10 mm wide, with large teeth or shallow-lobate along margin, with
short cartilaginous cusp. Capitula solitary or 2—20(40), on long (to 6
cm long), often leafy peduncles, in lax corymbs or corymbose-panicles.
Involucre 11-16 mm in dia, 4-6 mm long, glabrous or subglabrous;
involucral bracts herbaceous, outer lanceolate, inner 1.5, rarely less
often 2 times as long, oblong-linear; all bracts with rather wide, dark
brown or brownish membranous border, modified in inner bracts in
light colored, lacerate-fimbriate appendage at apex. Ligulate florets
white, with corolla tube 1.5—2.0 mm long, and limb 8-15 mm long;
corolla of tubular florets 2.0—2.5 mm long. Achenes 2.0—2.5 mm long,
0.7-0.8 mm wide, with 5-8 longitudinal ribs and about 0.3 mm long
subobtuse or lobed corona. Flowering June to August.

Marshes, marshy meadows at altitudes from 1,000 to 3,500 m.—
Caucasus: Eastern Transcaucasia (southwestern part), Western

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208

Transcaucasia (southern part), Southern Transcaucasia. General distri-
bution: Armenia-Kurdistan. Described from Turkey. Type in Paris.

Note. A highly polymorphic species, possibly comprising several
ecogeographical races.

Series 2. Marionia Tzvel.—Plants with weakly developed short
nonflowering shoots, subglabrous (with very short, papilliform, simple
hairs); stems more densely leafy; leaves pinnately parted, with broad
apical lobes; receptacle strongly convex; ligulate florets white; corona
of achenes 0.2—0.3 mm long.

18. P. marionii Albov in Bull. Herb. Boiss. III (1895) 92; Sosn.
in Tr. Tifl. Bot. Sada, XVII (1915) 26; Grossh. Fl. Kavk. IV, 134 and
Opred. Rast. Kavk. 460; Manden. in Fl. Gruzii, VIII, 355, Plate 394.—
Ic.: Kolakovskii, Fl. Abkh. IV (1949) Plate 23; Manden. op. cit.

Perennial. Plants 20-70 cm high, with more or less branched
inclined rhizome, more or less covered with very short, papilliform,
simple hairs, usually subglabrous. Stem erect or ascending at base,
sparsely leafy, simple or with few lateral branches from middle. Leaves
subglabrous, usually puberulent beneath, with numerous punctate-
glandular hairs; basal and lower cauline leaves to 20(25) cm long, 6(8)
cm wide, on rather long, basally thickened petiole; lamina oblong or
linear-oblong, relatively flat, pinnately parted up to narrow-winged
rachis; segments 4—7 on each side, usually narrowed toward base,
oblong to oblong-linear, pinnately lobed and irregularly sharply toothed;
apical lobes semirotund-deltoid to broadly lanceolate, 2-5 mm wide;
cauline leaves smaller, lower short-petiolate, upper sessile or subsessile,
weakly divided. Capitula solitary, less often 2—5 on single stem, forming
lax irregular corymbs. Involucre 14-19 mm in dia, 6.0—-7.5 mm long,
slightly pubescent; involucral bracts coriaceous-herbaceous, outer
broadly lanceolate, subobtuse, inner 1.5—2 times as long, linear-oblong;
all bracts with rather wide, dark brown, membranous border, apically
more or less appendiculately broadened in inner bracts. Ligulate florets
white, with corolla tube 1.6—2.0 mm long and limb 8-15 mm long;
corolla of tubular florets 2.5—3.0 mm. long. Achenes 2.0—2.6 mm long,
about 0.6 mm wide, with S—8 longitudinal ribs; corona 0.2—0.3 mm
long, toothed or irregularly lobed to middle or almost to base, strongly
reduced dorsally in ligulate florets. Flowering July to August.

Limestone rocks, stony slopes, grassy areas at altitudes of 1500-
2500 m.—Caucasus: Western Transcaucasia (Abkhazian ASSR).
Endemic. Described from Abkhazian ASSR. Type in Leningrad.

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Series 3. Rosea Sosn. in Tr. Tifl. Bot. Sada, XVII (1915) 21.—
Plants not always with developed short, nonflowering shoots, more or
less covered with short bifid and simple hairs, often subglabrous; stems
usually weakly leafy; leaves once or twice pinnately cut; receptacle
strongly convex; ligulate florets pink, of various shades; corona of
achenes 0.1-0.2 mm long.

19. P. roseum (Adam) MB. FI. taur.-cauc. II (1808) 324 and III
(1819) 578, quoad nomen.—P. carneum MB. ibid. II (1808) 325 and
III (1819) 578; DC. Prodr. VI, 55; Ldb. Fl. Ross. II, 550; Boiss. FI.
or. III, 340; Sosn. in Tr. Tifl. Bot. Sada, XX VII, 25; Grossh. Fl. Kavk.
IV, 133 and Opred. Rast. Kavk. 460; Manden. FI. Gruzii, VIII, 355.—
P. roseum var. carneum (MB.) Trautv. in Tr. Peterb. Bot. Sada, V, 2
(1878) 443.—Chrysanthemum coronopifolium Willd. Sp. pl. II (1803)
2145, non Vill. 1785.—C. roseum Adam in Web. and Mohr. Beitr. I
(1805) 70.—C. carneum (MB.) Steud. Nomencl. (1821) 192; Parsa,
Fl. Iran. III (1949) 267.—C. roseum var. carneum (MB.) Schmalh. FI.
II (1897) 70.—Tanacetum roseum (Adam) Sch. Bip. Tanacet. (1844)
58, quoad nomen.—T. carneum (MB.) Sch. Bip. ibid. 58.—Exs.: GRF
No. 774.

Perennial. Plants 20-60 cm high, with more or less branched,
inclined rhizome, usually covered with semi-erect bifid and simple
hairs, often subglabrous. Stem erect or ascending at base, sparsely
leafy, simple, less often with 1-2 lateral branches. Leaves usually sub-
glabrous, lacking distinct punctate-glandular hairs; basal and lower
cauline leaves up to 15 cm long and 4 cm wide, rather long-petiolate,
with oblong or linear-oblong, pinnately cut lamina, segments 3-7 on
each side, oblong or linear-oblong, irregularly bidentate or pinnately
lobed, up to 5-7 mm wide, terminal lobes more or less toothed; middle
and upper cauline leaves small, sessile or subsessile, usually pinnatisect,
with more or less toothed segments. Capitula solitary, less often 2-3,
on long peduncles often leafy in lower part. Involucre 12-20 mm in
dia, 4-6 mm long, more or less appressed-hairy to subglabrous;
involucral bracts herbaceous, outer linear-lanceolate, subobtuse, inner
not more than 1.5 times as long, oblong-linear; all bracts with quite
broad dark brown or brownish membranous border, somewhat broader
at apices of inner bracts. Ligulate florets in various shades of pink,
with corolla tube 1.2—2 mm long and limb 12-25 mm long; corolla of
tubular florets 2.6—3.2 mm long. Achenes 2.2—3.2 mm long, with 5-8
longitudinal ribs; corona 0.1—0.2 mm long, irregularly blunt-toothed.
Flowering June to August.

Meadows, stony slopes at altitudes of 1,500-3,000 m.—Caucasus:
Ciscaucasia, Dagestan, Eastern Transcaucasia, Western Transcaucasia
(northern part, rarely). General distribution: Armenia-Kurdistan(?).

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210

Described from Caucasus. Type in Leningrad.

Note. This species is widely known by the name P. carneum MB.
However, it, also includes the type specimens of Chrysanthemum
roseum Adam, a fact noted even by Ascherson [in Hoffman, Nat.
Pflanzenfam. IV, 5 (1889) 278], who gave “Pyrethrum roseum MB”.
a new name—“Chrysanthemum marschallii Aschers”’. In the very short
description given by Adam (1. c.), the segments of the leaves are
reported as pinnatipartite. The type specimens of Chrysanthemum
roseum Adam actually have somewhat more deeply divided leaf
segments than in typical specimens of Pyrethrum carneum MB., but,
nevertheless, their affinity to Pyrethrum carneum MB. and not to “P.
roseum MB.” is quite apparent. Adam’s specimens are missing from
the Herbarium of Marschall-Bieberstein, who, apparently, did not see
them.

Economic Importance. Sometimes cultivated as an ornamental
plant. There are several cultivars, which differ in the color of the
ligulate florets and the capitulum size, including “double-flowered”
cultivars with tubular florets, entirely or partly modified into ligulate
florets. It contains a significant percentage of the insecticidal sub-
stance pyrethrin and is cultivated occasionally to obtain different
insecticidal preparations (“Pyrethrum’’).

20. P. coccineum (Willd.) Worosch. in Spisok Sem. Gl. Bot. Sada
Akad. Nauk SSSR, IX (1954) 21.—P. raseum auct.: MB. FI. taur.-
cauc. II (1808) 324 and III (1819) 578, quoad pl.; DC. Prodr. VI, 56;
Ldb. Fl. Ross. II, 549; Sosn. in Tr. Tifl. Bot. Sada, XVII, 21, excl.
var.; Grossh. Fl. Kavk. IV, 133 and Opred. Rast. Kavk. 460; Manden.
in Fl. Gruzii, VIII, 352, non Chrysanthemum roseum Adam.—P. roseum
var. adami Trautv. in Tr. Peterb. Bot. Sada, V, 2 (1878) 443.—
Chrysanthemum coccineum Willd. Sp. pl. II (1803) 2144.—C.
marschallii Aschers. in Hoffm. in Pflanzenfam. IV, 5 (1889) 278,
nom. nov.—C. roseum auct. non Adam; Schmalh. FI. II, 70, excl.
var.—Tanacetum roseum (Adam) Sch. Bip. Tanacet. (1844) 58, quoad
pl.—Ic.: MB. Cent. Pl. rar. ruth. I (1810) t. 34; Zemlinskii, Lekarstv.
Rast. SSSR, ed. 2, 234.—Exs.: Pl. or. exs. No. 200.

Perennial. Plants 20-60 cm high, with more or less branched
inclined rhizome, somewhat covered with semi-erect bifid and simple
hairs, usually subglabrous. Stem erect or ascending at base, sparsely
leafy (more weakly than in previous species), simple, very rarely with
single lateral branch. Leaves like previous species, but more incised
segments in lower cauline and basal leaves pinnately cut or parted,
usually with pinnately parted or more or less lobate secondary lobes;
segments of middle and upper cauline leaves usually also pinnately

220

211

parted. Capitula solitary, rarely in 2 on single stem, on long (up to 10-
15 cm), more or less pubescent peduncles. Involucre 12—20 mm in dia,
4—6 mm long, more or less appressed-hairy to subglabrous; involucral
bracts like previous species, but usually with broader, dark brown,
membranous border. Ligulate florets in different shades of pink or red,
with corolla tube 1.2—2.0 mm long and limb 12-25 mm long; corolla
of tubular florets 2.6—3.2 mm long. Achenes 2.2—3.2 mm long with 5-
8 longitudinal ribs and irregularly blunt-toothed, 0.1-0.2 mm long
corona. Flowering June to August.

Meadows, stony slopes at altitudes of 1,500—3,000 m.—Caucasus:
Ciscaucasia (rarely), Dagestan (rarely), Eastern, Western and Southern
Transcaucasia (northern and western parts). General distribution:
Armenia-Kurdistan. Described from Caucasus. Type in Berlin.

Note. A highly polymorphic species, possibly comprising several
morphologically weakly separated local races, partly transitional to the
previous species. The variety “P. roseum var. album Grossh.” (FI.
Kavk. IV, 133) with white ligulate florets either belongs to one of the
following species or was described on the basis of albino plants.

Economic Importance. Like the previous species, it sometimes is
cultivated as an ornamental plant and has medicinal value (for preparing
insecticides).

Series 4. Daucifolia Tzvel.—Plants with well developed, short,
nonflowering shoots, more or less covered with short bifid and simple
hairs, sometimes sub-glabrous. Stem sparsely leafy; leaves 1-2
pinnatisect, with broader terminal lobes; receptacle slightly convex;
ligulate florets white; corona of achenes 0.1-0.3 mm long.

21. P. chamaemelifolium (Somm. and Lev.) Sosn. in Zam. po
Sist. i Geogr. Rast. Bot. Inst. Akad. Nauk Gruz. SSR, XV (1949) 3,
descr. emend.; Grossh. Opred. Rast. Kavk. 461; Manden. in Fl. Gruzii,
VIII, 356.—P. roseum var. chamaemelifolium Somm. and Lev. in Tr.
Peterb. Bot. Sada. XVI (1900) 235; Sosn. in Tr. Tifl. Bot. Sada, XVII,
25; Grossh. Fl. Kavk. IV, 133.—Ic.: Manden. op. cit. Plate 395.

Perennial. Plants 20-50 cm high, with more or less branched
rhizome, sparsely covered with semi-erect, bifid and simple hairs, often
subglabrous. Stem erect or ascending at base, very sparsely leafy,
simple. Leaves very rarely pubescent, mainly beneath, lacking distinct
punctate-glandular hairs; basal and lower cauline leaves up to 20 cm
long and 5 cm wide; on rather long (often exceeding lamina) petiole;
lamina ovate to oblong, pinnately cut to almost wingless axis, with 3-
7, sessile or short-petiolate, broadly ovate to oblong segments on each
side, pinnately parted, with lobes 2—5-parted or pinnate; terminal

221

212

segments and lobes deltoid to lanceolate-linear, up to 2 mm wide,
gradually narrowed at apex into rather long cartilaginous cusp; middle
and upper cauline leaves strongly reduced, weakly divided, upper leaves
usually sessile. Capitula solitary, on long (15-20 cm), more or less
glabrous peduncles. Involucre 12-17 mm in dia, 4.5-6.0 mm long,
only basally more or less appressed-hairy; involucral bracts herbaceous,
outer lanceolate, subobtuse, inner 1.5, rarely 2 times as long, broadly
linear; all bracts with brownish or dark brown membranous border,
appendiculately broadened at apex in inner bracts. Ligulate florets
white, with corolla tube 1.5—2.0 mm long and limb 10-18 mm long;
corolla of tubular florets 2.4-3.2 mm long. Achenes 2.4—3.0 mm long
with 8-10 longitudinal ribs and about 0.1—0.2 mm long finely toothed
corona. Flowering June to July.

Rocks, stony slopes (especially on limestone deposits) in upper
mountain zone.—Caucasus: Western Transcaucasia (northern part).
Endemic. Described from Caucasus (Rioni River basin and its upper
reaches). Type in Florence; isotype in Leningrad.

22. P. silaifolium Stev. in DC. Prodr. VI (1837) 57.—P. roseum
var. leucanthum C.A.M. Verzeichn. (1831) 73; Boiss. Fl. or. II, 340.—
P. daucifolium Stev. ex Karel. in Bull. Soc. Nat. Mosc. XII (1839)
158, nom. nud. non Ldb.—P. daucifolium (? Pers.) Ldb. Fl. Ross. II
(1845-1846) 549 p. p.; Grossh. Opred. Rast. Kavk. 461.—P.
ceratophylloides auct. non Ten.; C. Koch in Linnaea, XXIV (1851)
330.—P. paucifolium C.A.M. ex Trautv. in Tr. Peterb. Bot. Sada,
VIII, 2 (1883) 456.—P. meyerianum Sosn. in Tr. po Geob. Obsl. Pastb.
Azerb. Ser. A, VII (1931) 103-104 and in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XIX (1959) 642; Grossh. Fl. Kavk. IV, 135.—
Tanacetum silaifolium (Stev.) Sch. Bip. Tanacet. (1844) 61.—
Chaemaemelum silaifolium (Stev.) Trautv. ex Grossh. Fl. Kavk. IV
(1934) 128 and Opred. Rast. Kavk. 458.—Ic.: Sosn. I. c. (1931) t. IX.

Perennial. Plants 10-40 cm high, with branched creeping rhizome,
more or less covered with semi-erect bifid and simple hairs, often
mixed with very short papilliform hairs. Stem erect or ascending at
base, very sparsely leafy, simple or branched near middle, with 1-3
lateral branches. Leaves sparsely pubescent or glabrous, usually lacking
distinct punctate-glandular hairs; basal and lower cauline leaves up to
10 cm long and 2 cm wide, on long (often as long as lamina) petiole;
lamina oblong, pinnately cut up to wingless axis; segments 4-8 on
each side, short-petiolate or subsessile, broadly ovate to oblong, 1-2
pinnately parted or almost 3-7 palmately parted; terminal segments or
lobes broadly lanceolate to almost deltoid, up to 22 mm wide, narrowed
into rather long cartilaginous cusp; remaining cauline leaves strongly

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213

reduced, usually all short-petiolate. Capitula solitary or 2—4 on each
stem, on very long (up to 25 cm long) slender peduncles. Involucre
10-13 mm in dia and 4.0-5.5 mm long, more or less pubescent;
involucral bracts herbaceous, outer lanceolate or ovate-lanceolate, inner
not more than 1.5 times as long, oblong; all bracts with rather broad,
brownish, less often light colored, membranous border, appendiculately
broadened at apex of inner bracts. Ligulate florets white, with corolla
tube 1.0-1.5 mm long and limb 7-13 mm long; corolla of tubular
florets 2.2-3.0 mm long. Achenes 2.0-2.5 mm long with 5-8
longitudinal ribs and 0.2-0.3 mm long irregularly toothed corona.
Flowering June to August.

Shale screes, stony slopes, usually at altitudes of 1,000—2,500 m.—
Caucasus: Dagestan (southern part), Eastern Transcaucasia (northern
part), Talysh (?). Endemic. Described from eastern part of Main
Caucasian Range. Type in Helsinki; isotype in Leningrad.

Note. A very close ecogeographical race of this series—P.
daucifolium (Pers.) Ldb. [Ldb. op. cit. 549 = Chrysanthemum
daucifolium (Pers.) Syn. pl. I (1807) 462]—with larger (on the average)
capitula and a darker and broader membranous border of the involucral
bracts is found in the mountains of northern Iran (and, possibly, also
Talysh). However, I am not completely sure that this is actually
Chrysanthemum daucifolium Pers., described on the basis of garden
plants of unknown origin, and in this respect I must rely exclusively
on the authority of Ledebour. P. silaifolium was initially reported by
Steven under the name “P. daucifolium Stev.” (probably, independently
of Chrysanthemum daucifolium Pers.), but later was renamed with its
present name. P. paucifolium C.A.M. and P. meyerianum Sosn.,
described later from Meyer’s specimens, from the Eastern Caucasus
are undoubtedly synonyms of this species.

Series 5. Daghestanica Tzvel.—Plants with well developed, short,
nonflowering shoots, usually covered with short simple hairs; stems
sparsely leafy; leaves pinnatisect, with rather narrow terminal lobes;
receptacle strongly convex; ligulate florets white; corona of achenes
0.3-0.5 mm long.

23. P. daghestanicum (Rupr. ex Boiss.) Rupr. ex Flerov. Spisok.
Rast. Sev. Kavk. (1938) 563; Sosn. and Manden. in Bot. Zhurn-SSSR.
XXXIV, 3 (1949) 288.—Chamaemelum daghestanicum (Rupr. ex
Boiss.) Boiss. Fl. or III (1875) 334, excl. var.; Grossh. Fl. Kavk. IV,
128 and Opred. Rast. Kavk. 458.—Tripleurospermum daghestanicum
Rupr. ex Boiss. |. c. in syn.

223

214

Perennial. Plants 5-18 cm high, with more or less branched rhi-
zome, sparsely covered with erect or semi-erect, simple hairs, strongly
aromatic. Stem erect or ascending at base, simple, sparsely leafy. Leaves
more or less hairy, with inconspicuous punctate-glandular hairs; basal
and lower cauline leaves up to 5 cm long and 2 cm wide, on rather
long (sometimes exceeding lamina), and basally thickened petiole,
lamina broadly ovate to oblong, pinnately cut up to almost wingless
axis; segments 2-4 on each side, short-petiolate or sessile, broadly
ovate to oblong, pinnately parted up to narrow-winged axis, sometimes
almost pinnately lobed, terminal segments or lobes obovate to oblong,
1.5-2.0 mm wide, subobtuse or with very short cusp; upper cauline
leaves strongly reduced and weakly divided (sometimes partly entire),
sessile or short-petiolate. Capitula solitary, on rather densely appressed-
hairy peduncles, up to 3(4) cm long. Involucre 10-13 mm in dia, 4.0—
5.5 mm long, more or less pubescent; involucral bracts herbaceous,
outer rather broadly lanceolate, inner 1.5—2 times as long, linear ob-
long; all bracts with rather broad, brownish or dark brown, membra-
nous border, appendiculately broadened at apex of inner bracts. Ligu-
late florets white, corolla tube about 1 mm long and limb 7-11 mm
long; corolla of tubular florets 2.2-2.6 mm long. Achenes 1.8—2.2 mm
long and about 0.6 mm wide, with 5-7 longitudinal ribs, corona 0.3-
0.5 mm long, irregularly sharply toothed up to middle. Flowering July
to August.

Stony slopes, rocks at altitudes above 2000 m.—Caucasus:
Dagestan, Eastern Transcaucasia (northern part). Endemic. Described
from Southern Dagestan. Type in Leningrad.

24. P. aromaticum (Rupr. ex Boiss.) Tzvel. comb. n.—Cham.
daghestanicum var. glabrous Boiss. Fl. or. III (1875) 334.—
Tripleurospermum aromaticum Rupr. ex Boiss. ibid. in syn.—?T. nivale
Rupr. ex Boiss. ibid. in syn.

Perennial. Plants 5-15 cm high, with more or less branched
rhizome, usually sparsely covered with erect, simple hairs, often
subglabrous, strongly aromatic. Stem erect or ascending, sparsely leafy,
simple. Leaves like previous species, but usually glabrous or subglabrous
and with narrower terminal lobes (up to 1 mm wide) than in previous
species. Capitula solitary, on more or less pubescent, up to 3 cm long,
peduncle. Involucre 10-13 mm in dia and 4-5 mm long, glabrous or
subglabrous (sparsely hairy near base); involucral bracts herbaceous,
outer narrow-lanceolate, inner not more than 1.5 times as long as outer,
oblong-linear; all bracts with rather wide, dark brown, membranous
border, more or less appendiculately broadened at apex of inner bracts.
Ligulate florets white, corolla tube about 1 mm long and limb 7-11

224

215

mm long; corolla of tubular florets 2.2-2.6 mm long. Achenes like
previous species; mature achenes unknown. Flowering July to August.

Stony slopes, grassy patches, rocks above 2,500 m.—Caucasus:
Dagestan. Endemic. Described from Dagestan (Diklos-Mta Mountain).
Type in Leningrad.

Note. It is a more western or more alpine ecogeographical race.
So far only the type specimens of this species are known, which, ap-
parently, is explained by its distribution in inacessible and poorly stud-
ied regions of Dagestan. The specimens of Tripleurospermum nivale
Rupr. ex Boiss. are even smaller and were collected with still un-
opened capitula. It is possible that they belong to one of the species
of Tripleurospermum Sch. Bip.

Series 6. Trichophylla Tzvel.—Plants with well developed, short,
nonflowering shoots, usually covered with semi-erect simple hairs, often
subglabrous; stems sparsely leafy; leaves twice-pinnately cut, with very
narrow lobes; receptacle strongly convex; ligulate florets white; corona
of achenes 0.2-0.3 mm long.

25. P. tricholobum Sosn. ex Manden. nom. nov. in Zam. po Sistem.
i Geogr. Rast. Bot. Inst. Akad. Nauk Gruz. SSR, XXII (1961) 58.—
P. trichophyllum Sosn. in Zam. po Sist. i Geogr. Bot. Inst. Akad. Nauk
Gruz. SSR, XV (1949) 2, non Griseb. 1844.

Perennial. Plants 15—35 cm high, with slender branched rhizome,
usually covered with sparse semi-erect, simple hairs, usually
subglabrous. Stem erect or ascending at base, with few cauline leaves
only in lower part, simple. Leaves glabrous or subglabrous, lacking
distinct punctate-glandular hairs; basal and lower cauline leaves 10-12
cm long and 3.5 cm wide, on rather long petiole; lamina obovate to
oblong, twice pinnately cut, with 4-6 segments on each side; terminal
segments somewhat thick, linear-filiform, up to 0.6 mm wide, and 12
mm long; other cauline leaves strongly reduced, short-petiolate or
subsessile. Capitula solitary, on rather long (10-15 cm) peduncles,
more or less hairy in upper part. Involucre 8-15 cm in dia, 4-6 mm
long, more or less pubescent to subglabrous at base; involucral bracts
herbaceous, outer narrow-lanceolate, inner usually not more than 1.5
times as long as outer, oblong-linear; all bracts with wider dark brown,
membranous border, more or less expanded to broadly lanceolate
appendage at apex. Ligulate florets white, corolla tube 1.0-1.5 mm
long and limb 8-12 mm long; corolla of tubular florets 2.4-3.0 mm
long. Achenes (only immature achenes known), 1.8—2.4 mm long, with
5-6 longitudinal ribs and irregularly sharply toothed (sometimes divided
up to base) corona 0.2—0.3 mm long, about 0.1 mm in ligulate florets.
Flowering May to June.

227

216

Stony slopes, grassy patches, rocks above 2,000 m.—Caucasus:
Ciscaucasia (eastern part), Dagestan. Endemic. Described from north-
ern Ossetia. Type in Tbilisi.

Note. Morphologically, it is extraordinarily similar to the Cauca-
sian species of the genus Tripleurospermum Sch. Bip., differing only
by the very sparsely leafy stems and the structure of the achenes with
a shorter corona. Because of the absence of mature achenes, the affin-
ity of this species to the genus Pyrethrum still cannot be considered
finally proved.

Series 7. Komarovia Tzvel.—Plants with well developed, short,
nonflowering shoots, more or less covered with simple hairs, often
subglabrous; stems sparsely leafy; leaves pinnately cut, with very narrow
lobes; receptacle strongly convex; ligulate florets white; corona of
achenes 0.5—1.3 mm long.

26. P. komarovii Sosn. in Dokl. Akad. Nauk Arm. SSR, II, 4
(1945) 119; Grossh. Opred. Rast. Kavk. 461.

Perennial. Plants 5—20(25) cm high, with strongly branched
rhizome, more or less covered with erect simple hairs mixed with
pappilliform hairs, usually subglabrous. Stems erect or ascending at
base, usually numerous and often forming pulvinous turf together with
short nonflowering shoots, very sparsely leafy, simple. Leaves weakly
pubescent, often subglabrous, with rather numerous punctate glandular
hairs; basal and lower cauline leaves up to 3.5(4.0) cm long and 1.5(2.0)
cm wide, on shorter, strongly flattened and basally thickened petiole;
lamina broadly oblong to almost rotund, pinnately cut into very narrow
(up to 0.8 mm wide), linear-filiform, acute, 2—5-parted segments, 4—10
on each side; other cauline leaves strongly reduced, sessile or subsessile,
usually with more or less pinnately parted lamina. Capitula solitary,
on long (up to 10-12 cm), sparsely pubescent peduncle. Involucre 9—
13 mm in dia, 4.0-5.5 mm long, more or less pubescent to entirely
glabrous; involucral bracts herbaceous, outer lanceolate, subobtuse,
inner 1.5—2 times as long, broadly linear; all bracts with wide blackish-
brown membranous border, appendiculately broadened at apex of inner
bracts. Ligulate florets white, corolla tube 1.5—-2.0 mm long and limb
8-12 mm long; corolla of tubular florets 2.6—3.2 mm long. Achenes
(mature achenes unknown) 1.8—2.4 mm long and about 0.5 mm wide,
with 5(?) longitudinal ribs; corona of achenes of tubular florets 0.5—
1.3 mm long, strongly variable in shape and size, usually 2—5-lobed or
divided almost to base, irregularly sharply toothed along margin;
achenes of ligulate florets 0.1-0.3 mm long, often secund. Flowering
June to July.

217

Plate IX.
Habit of plant, corolla of ligulate florets, corolla of tubular floret, and achene:

225

1 — Pyrethrum tienschanicum Krasch.,; 2— P. leontopodium (Winkl.) Tzvel.

228

218

Rocks, stony slopes above 2,000 m.—Caucasus: Southern Trans-
caucasia (Zangezur and Murov-Dag ranges). Endemic(?). Described
from Zangezur Range. Type in Tbilisi.

Note. The taxonomic position of this species, which strongly re-
sembles some Transcaucasian and West Asian species of the genus
Tripleurospermum Sch. Bip., remains unclear to me. Besides the
previous species, with which P. komarovii shows a strong but,
apparently, only superficial resemblance, this species is also more or
less approaches species of section Xylopyrethrum, differing from them
by the nature of the pubescence and nonlignified branches near the
base, and some species of section Richteria (e.g., P. pulchellum).
Because of the absence of mature achenes, the affinity of P. komarovii
to the genus Pyrethrum cannot be considered as finally proved.

Section 8. Leucanthemopsis (Giroux) Tzvel. comb. nova.—
Tanacetum sect. Leucoglossa § 3. Alpina Sch. Bip. Tanacet. (1844)
59.—T. sect. Pyrethrum subsect. Leucanthemopsis Giroux in Bull. Soc.
Hist. Nat. Afr. Nord. XXIV (1933) 54.—Herbaceous plants, 5—20 cm
high, more or less covered with short bifid hairs mixed with simple
hairs, with sparsely leafy stems and well developed short nonflowering
shoots. Leaves pinnately lobed, lower petiolate, upper sessile or
subsessile. Capitula solitary. Involucre patelliform, 12-18 mm in dia;
outer involucral bracts 2/3, less often 1/2 as long as inner. Receptacle
strongly convex. Corolla limb of ligulate florets oblong, 7-15 mm
long; corolla of tubular florets 2-3 mm long, with tube fairly strongly
and abruptly broadened in upper half, toothed with lanceolate deltoid
deflexed teeth. Achenes 2-3 mm long, with pericarp very loosely
enclosing kernel and 5 weak ribs; corona 0.5—1 mm long, more or less
irregularly toothed along margin.

Type of section: Pyrethrum alpinum (L.) Schrank.

Note. This section with a small number of species is distributed
exclusively in the mountain regions of Western Europe.

27. P. alpinum (L.) Schrank, Prim. Flor. Salisb. (1792) 215; DC.
Prodr. VI, 54.—Chrysanthemum alpinum L. Sp. pl. (1753) 889.—
Leucanthemum alpinum (L.) Lam. FI. Franc. II (1778) 138.—Matricaria
alpina (L.) Desr. in Lam. Encycl. meth. III (1792) 730.—Tanacetum
alpinum (L.) Sch. Bip. Tanacet. (1844) 61.—Pontia alpina (L.) Bub.
Fl. Pyr. II (1900) 219.—Ic.: Rchb. Ic. Fl. Germ. XVI (1853) t. 990;
Hegi, Ill. Fl. VI, 2, f. 319 and 320; Jav. and Csap. Ic. Fl. Hung. 522,
t. 38.—Exs.: Schultz, Herb. norm. nov. ser. No. 2203; FI. Ital. exs.
No. 1157; Hayek, FI. Stir. exs. No. 795; Fl. exs. Reip. Bohem. Sloven.
No. 80; Fl. Pol. exs. No. 353; Pl. Pol. exs. No. 173.

229

pa Le

Perennial. Plants 5-15 cm high, with short creeping, more or less
branched rhizome, more or less covered with appressed or semi-erect,
bifid, often simple hairs. Stems few or solitary, erect or basally
ascending, weakly leafy, simple. Leaves green, more or less pubescent
to subglabrous; basal and lower cauline leaves rather long-petioles,
usually 1.5-3 times as long as lamina; lamina more or less ovate,
cuneately narrowed toward base, pinnately lobed, with 2—5 lobes on
each side; other cauline leaves strongly reduced, less deeply lobed to
linear with entire margin. Capitula solitary, on rather long, more or
less pubescent peduncle. Involucre 12—18 mm in dia, 3.5—5.0 mm long,
more or less pubescent to subglabrous; involucral bracts herbaceous,
outer oblong-linear, inner linear, 1.5—2 times as long; all bracts with
wide, blackish-brown, more or less discontinuously fimbriate,
membranous border. Ligulate florets white, corolla tube about 1.5 mm
long and limb 7-15 mm long; corolla of tubular florets 2-3 mm long.
Achenes 2-3 mm long, about 0.8-1.0 mm wide, with 5 somewhat
prominent longitudinal ribs and 0.5-1.0 mm long irregularly toothed
corona. Flowering July to August.

Stony slopes, grassy areas, rocks in upper mountain zone.—
European part: Upper Dniester (Carpathians). General distribution:
Central Europe, Balkans, Mediterranean Region. Described from the
Alps. Type in London.

Note. A highly polymorphic species, divided in several
ecogeographical races mainly differing in the shape of the leaves, size
of the capitulum and pubescence. Two very close ecogeographical
races are distributed in the Carpathians, which perhaps deserve to be
separated as independent species: Pyrethrum alpinum ssp. tatrae
(Vierh.) Tzvel. comb. nova. [= Chrysanthemum alpinum f. tatrae Vierh.
in Mag. Bot. Lapok XIII (1914) 25]—the more northern and higher
mountain race, characterized by rather dense pubescence and smaller
dimensions of the entire plant, and P. alpinum ssp. cuneifolium (Murtr.)
Tzvel. comb. nova (= Leucanthemum alpinum var. cuneifolium Murr.
in Deutsche Bot. Monatsschr. XIV (1896) 21)— the more southern and
lower mountain race, distinguished by very sparse pubescence (plants
usually almost glabrous). The species is reported by the western
European authors from the Carpathians within the Soviet Union,
although so far it has not been found here.

Section 9. Xylopyrethrum Tzvel. sect. nova in Addenda, XXV,
872.—Plants more or less woody at base, 5-20 cm high, more or less
covered with adpressed or appressed bifid hairs, often mixed with
simple hairs, with more sparsely leafy stem and well developed, short,
non-flowering shoots (usually with elongated, more or less lignified

220

base). Leaves more or less pinnately or palmately cut, with very nar-
row lobes; all leaves more or less long-petiolate. Capitula solitary (but
usually numerous on single plant). Involucre 5.5-12.0 mm in dia,
narrow-patelliform or cupuliform. Receptacle strongly convex, conical-
hemispherical. Corolla limb of ligulate florets oval to oblong, 7-12
mm long; corolla of tubular florets 2.5—-3.2 mm long, with tube fairly
weakly and gradually broadened in upper half, and lanceolate-deltoid
recurved teeth. Achenes 2—3 mm long, with 5—7 longitudinal ribs, more
or less shifted ventrally in ligulate florets; corona 0.2-1.0 mm long,
more or less irregularly toothed or lobed, in ligulate florets usually
divided dorsally almost up to base.

Type of section: Pyrethrum kotschyi Boiss.

Note. A whole series of species (including P. tenuisectum Boiss.,
P. bornmiilleri Hausskn. and others), distributed in the mountainous
regions of Western Asia, belong to this section, which does not show
a distinct affinity with other sections of the genus. The section is
divided in several distinctly isolated series.

Series 1. Kotschyana Tzvel.—Plants grayish from dense pube-
scence; involucre narrow-patelliform, 9-12 mm in dia; corona 0.5-1.0
mm long.

28. P. kotschyi Boiss. Diagn. ser. 1, VI (1845) 88 and FI. or. III,
339; Grossh. Fl. Kavk. IV, 135 and Opred. Rast. Kavk. 461.—
Chrysanthemum kotschyi (Boiss.) Nabel. in Publ. Fac. Sc. Univ.
Masaryk. Brno, No. 52 (1925) 19.—Ic.: Nabel. ibid. t.. 5.—Exs.:
Kotschy, Pl. Pers. austr. No. 770 (isotypes of species); Bornmiiller,
Iter pers. alt. No. 7443; Pl. or. exs. No. 322.

Perennial. Plants more or less woody at base, 5-20 cm high, with
strongly thickened woody rhizome, grayish from dense pubescence of
semierect bifid (often mixed with simple) hairs. Stems usually rather
numerous, erect or basally ascending, more or less branched only at
base. Leaves grayish from dense pubescence, with inconspicuous
punctate-glandular hairs; basal leaves numerous, up to 3 cm long, 1.5
cm wide, on rather long (as long as or slightly exceeding lamina)
petiole, strongly thickened at base; lamina reniform or reniform-
orbicular with narrow-cuneate base, 3—7-palmately cut or almost pinnate,
with segments usually themselves twice or thrice cut up; terminal
segments linear-subulate, up to 0.8 mm wide, with short cartilaginous
cusp; cauline leaves strongly reduced, short-petiolate, with 3-5 parted
lamina, uppermost leaves often entire. Capitula solitary (but many on
a single plant), on rather long (up to 10 cm) peduncle. Involucre 9-12
mm in dia, 4.0-5.5 mm long, finely tomentose near base; involucral

221

bracts coriaceous-herbaceous, outer lanceolate, inner 1.5, less often 2,
times as long, broadly linear; all bracts with wide, blackish-brown,
membranous border, appendiculately broadened at apex. Ligulate florets
white, with 1.2-1.8 mm-long corolla tube and 8-12 mm-long limb;
corolla of tubular florets 2.5—3.0 mm long. Achenes 2.0—2.5 mm long,
about 0.5 mm wide, with 5—6 longitudinal ribs and irregularly toothed
or lobed 0.5-1.0 mm long corona, often divided almost to base on
ventral side. Flowering June to July.

Stony slopes, rocks at 2,000-3,500 m.—Caucasus: Southern
Transcaucasia (eastern part of Nakhichevan ASSR). General
distribution: Armenia-Kurdistan, Iran (northwestern part). Described
from Iran. Type in Geneva; isotype in Leningrad.

29. P. ordubadense Manden. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XIX (1959) 358.—P. tenuisectum auct. non Boiss.: Sosn.
in Zhurn. Russk. Bot. Obshch. XIV (1929) 84; Grossh. Fl. Kavk. IV,
135 and Opred. Rast. Kavk. 461.

Perennial. Plants more or less woody at base, 5—20 cm high, with
thick woody rhizome, grayish from dense pubescence of bifid hairs
mixed with simple hairs. Stems usually rather numerous, erect or
ascending near base, more or less branched only near base. Leaves
like previous species; basal leaves sometimes larger (up to 4.5 cm
long). Capitula solitary (but numerous on a single plant), on rather
long peduncle. Involucre 9-12 mm in dia, 4.0-5.5 mm long, finely
tomentose near base; involucral bracts coriaceous-herbaceous, outer
broadly lanceolate a half, less ofte1i two-fifths as long as broadly linear
inner bracts; all bracts with rather » ‘de, brownish, membranous border,
appendiculately broadened at apex in inner bracts, slightly larger than
in previous species. Ligulate florets white, corolla tube 1.2-1.8 mm
long and limb 8-12 mm long; corolla of tubular florets 2.5—3.0 mm
long. Achenes like previous species. Flowering May to July.

Stony slopes, rocks at 1,500-3,000 m.—Caucasus: Southern
Transcaucasia (region of Ordubad). General distribution: Armenia-
Kurdistan(?). Described from Soyukh Mountain near Ordubad. Type
in Leningrad.

Note. This species, which is very close to the preceding species,
is a lower mountain ecogeographical race by comparison.

Series 2. Fruticulosa Sosn. in Tr. Tifl. Bot. Sada, XVII (1915)
16.—Plants weakly hairy; involucre cupuliform, 5.5—7.0 mm in dia;
corona 0.2—0.3 mm long.

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222

30. P. heldreichianum Fenzl. ex Tchih. As. Min. II (1860) 273.—
P. suffruticulosum Fenzl. ex Tchih. ibid. in syn.—P. fruticulosum Fenzl.
ex Boiss. Fl. or III (1875) 338; Sosn. in Tr. Tifl. Bot. Sada, XVII, 16;
Grossh. Fl. Kavk. IV, 135, non Biehl. 1807.—Tanacetum hel-
dreichianum Fenzl. ex Tchih. op. cit. 273, nom. altern.—Exs.: Sintenis,
Iter or No. 5639; Bourg. Pl. armen. No. 145.

Perennial. Plants more or less woody at base, 6—15 cm high, with
thick woody rhizome, more or less covered with semi-erect bifid and
simple hairs. Stems usually rather numerous, erect, more or less
branched only at base. Leaves usually dull green from sparser
pubescence, with numerous punctate-glandular hairs; basal leaves up
to 3.5 cm long, 1 cm wide, on rather long petiole, thickened at base;
lamina oblong to linear-oblong, pinnately cut; leaf segments 4—8 on
each side, approximate, linear-subulate, up to 0.6 mm wide, usually
entire, less often 2—3 parted, with short cartilaginous cusp; cauline
leaves strongly reduced, short-petiolate, uppermost entire. Capitula
solitary (but numerous on a single plant) on rather long (up to 8 cm)
peduncles. Involucre 5.5—7.0 mm in dia, 5.0-6.5 mm long, densely
pubescent at base; involucral bracts coriaceous-herbaceous, outer
lanceolate-ovate, a half, less often two-fifths as long as oblong-linear
inner bracts; all bracts with rather wide, dark brown, membranous
border, appendiculately broadened at apex in inner bracts. Ligulate
florets white, corolla tube 1.2—-1.8 mm long and limb 7—9 mm long;
corolla of tubular florets 2.5-3.0 mm long. Achenes 2.5—3.0 mm long,
0.6-0.7 mm wide, with 5-7 longitudinal ribs; corona of achenes of
tubular florets 0.2-0.3 mm long, usually irregularly toothed to middle,
divided to base on dorsal side, achenes of ligulate florets 0.1—-0.2 mm
long. Flowering June to August.

Rocks, stony slopes, screes in upper mountain zone.—Caucasus:
Likely to be found in south of Georgian SSR. General distribution:
Armenia-Kurdistan. Described from northeastern Turkey. Type and
isotypes in Leningrad.

Note. The species is common in the region of Artvin and Kars
near the USSR border, and its occurrence in the south of Transcaucasia
is very probable.

Section 10. Pyrethrum.—P. sect. Leucoglossa DC. Prodr. VI
(1837) 53 p. p.—P. sect. Eupyrethrum D. Sosn. in Tr. Tifl. Bot. Sada,
XVII (1915) 16 p. p.—Chrysanthemum sect. Pyrethrum (Zinn) Gaud.
Helv. V (1829) 346 p. p.; Hoffm. in Pflanzenfam. IV, 5 (1889) 278
p. p.—Tenacetum sect. Leucoglossa § 2. Macrosperma Sch. Bip.
Tanacet. (1844) 56.—T. sect. Pyrethrum subsect. Eupyrethrum Briq.
in Burnat, Fl. Alpes Marit. VI (1916) 119.—T. sect. Pyrethrum subsect.

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223

Pyrethrum (Zinn) Heywood in Anal. Inst. Bot. Cavanill. Madrid. XII,
2 (1953) 324.—Herbaceous plants, (15)20-100(150) cm high, usually
covered with rather long, bifid and simple, less often only simple
hairs, with more sparsely leafy stems and well developed, short,
nonflowering shoots. Leaves pinnately cut; lower leaves petiolate,
middle and upper sessile. Capitula usually 2-12, less often up to 20,
in lax or somewhat lax corymbs, sometimes solitary; involucre narrow-
patelliform or patelliform, 8-16 mm in dia; receptacle rather strongly convex,
almost hemispherical. Corolla limb of ligulate florets oblong-linear or
oblong, 9-16 mm long; corolla of tubular florets 1.8-2.4 mm long, tube
weakly and gradually broadened in upper half, with deltoid, weakly bent
teeth. Achenes 2.2—3.0 mm long, with 5—8 longitudinal ribs, more or less
ventrally shifted in ligulate florets; corona 0.3—1.5 mm long, irregularly
bluntly or subacutely lobed and finely toothed up to middle.

Type of section: type of genus.

Note. A small section in number of species to which, besides the
four species in the flora of the USSR described below, only a few
species of the genus from Southern Europe and Asia Minor belong.
All species are closely related ecogeographical races of a single series.

31. P. corymbosum (L.) Willd. Sp. pl. HII (1803) 2155; MB. FI.
taur.-cauc. II, 325 and III, 578; DC. Prodr. VI, 57; Ldb. Fl. Ross. I,
551; Boiss. Fl. or. III, 342, p. p.; Grossh. Fl. Kavk. IV, 136 and
Opred. Rast. Kavk. 461; Kryl. Fl. Zap. Sib. XI, 2750.—P. tauricum
Zelenetzky, Mat. Fl. Kryma (1906) 296.—Chrysanthemum corymbosum
L. Sp. pl. (1753) 890; Schmalh. Fl. Il, 69.—C. corymbiferum L. Sp.
pl. (1763) 1251.—Matricaria corymboasa (L.) Desr. in Lam. Encycl.
III (1792) 734.—Tanacetum corymbosum (L.) Sch. Bip. Tanacet. (1844)
57.—Leucanthemum corymbosum (L.) Gren. and Godr. Fl. Franc. I
(1850) 145.—Ic.: Rchb. Ic. Fl. Germ. XVI (1854) t. 993; Syreistsch.
Ill. Fl. Mosk. Gub. III, 268; Bonnier, Fl. Compl. France, Suisse et
Belg. V, t. 293; Hegi, Ill. Fl. VI, 2, t. 265, Fig. 2; Jav. and Csap. Ic.
Fl. Hung. 523; Krasheninn. in Fl. Yugo-Vost. Evrop. Ch. SSSR, VI,
345, Fig. 680; Mikhail. in Fl. Beloruss. V (1959) Plate 32.—Exs.: Fl.
exs. Austro-Hung. No. 3782; Pl. Herceg. exs. No. 194; Hayek. FI. Stir.
exs. No. 1076; Fl. exs. reip. Bohem.-Sloven. Nos. 283 and 947; PI.
Polon. exs. No. 273; Fl. Polon. exs. No. 741, GRF. No. 3265.

Perennial. Plants 30—120(150) cm high, with more or less thick,
inclined rhizome, more or less covered with rather long adpressed and
erect, bifid hairs, often mixed with simple hairs, sometimes subglabrous.
Stems solitary or few, erect, very sparsely leafy, usually more or less
branched only near tip, very rarely simple. Leaves usually sparsely
hairy, green, often completely glabrous above, with inconspicuous

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224

punctate-glandular hairs, or eglandular; basal leaves up to 30-40 cm
long, on rather long (but always shorter than lamina) petiole, with
oblong or linear-oblong lamina, pinnately cut to almost wingless axis;
leaf segments 8—20 on each side, oblong or lanceolate, more or less
pinnately lobed or pinnately parted, with sharp-toothed lobes; cauline
leaves like basal but reduced and with shorter lamina, lower petiolate,
upper and middle sessile. Capitula (1)3—15(20), in rather lax, corymbs,
on long (up to 10 cm) peduncles more or less hairy above. Involucre
8-14 mm in dia, 4-5 mm long, more or less adpressed hairy at base
and frequently also on dorsally; involucral bracts coriaceous-herbaceous,
outer lanceolate-ovate, inner 1.5—2 times longer, broadly linear; all
bracts with narrow, light or brownish, membranous border, more or
less appendiculately broadened at apex in inner bracts. Receptacle 2—
3 times wider than long. Ligulate florets white, corolla tube about 1.5
mm long and limb 10-16 mm long; corolla of tubular florets 1.8—2.5
mm long. Achenes 2.0—2.5 mm long and about 0.8—1.0 mm wide, with
5 (rarely up to 7) prominent, longitudinal ribs and 0.5—0.8 mm long
corona, bluntly toothed or lobed up to middle. Flowering June to
August.

Deciduous forests, scrubs, forest glades up to middle mountain
zone.—European part: Ladoga-IImen (near Lake Onega), Volga-Kama,
Upper Volga, Upper Dnieper, Upper Dniester, Volga-Don, Trans-Volga
Region, Bessarabia, Black Sea Region, Lower Don, Lower Volga
(northern part), Crimea (mountains); Western Siberia: Upper Tobol
(sporadic), Irtysh (sporadic), Altai (northwestern foothills); Caucasus:
Ciscaucasia, Dagestan, Eastern Transcaucasia (northern part), Western
Transcaucasia, Southern Transcaucasia (northwestern part). General
distribution: Almost all of Europe, except northern regions, Turkey.
Described from Central Europe. Type in London.

Note. The specimens from the more arid habitats have denser silky
pubescence on the lower leaf surface. Probably, P. tauricum Zelenetzky
from Crimea was described from such specimens, which, unfortunately,
I did not see. Here too, apparently, belong the specimens from Crimea
identified by N. Zelenetzky (op. cit. p. 296) as “P. poteriifolium Ldb.,”
although occurrence of the latter species in the eastern regions of
Crimea is quite probable.

P. tenuifolium Willd. [Enum. hort. berol. (1809) 906], described
from garden specimens allegedly originating from the Caucasus,
according to the report of L. Reichenbach (FI. Germ. exc. II, 1830, p.
231), is an Italian species without close affinity to P. corymbosum.
Thus P. corymbosum var. tenuifolium (Willd.) Ldb. cannot be used for
Caucasian specimens of P. corymbosum with more deeply dissected
leaves.

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225

32. P. clusii Fisch. ex Rchb. Fl. Germ. exc. II (1830) 231.—P.
subcorymbosum (Schur) Schur, Enum. PI. Trans. (1866) 337; M. Popov,
Ocherk Rast. i Fl. Karpat. 250; Klok. in Vizn. Ros]. URSR, 540.—
Chrysanthemum subcorymbosum Schur in Verh. und Mitth. Siebenb.
Ver. X (1859) 146.—C. clusii (Fisch. ex Rchb.) Kreutz, Anthochron.
Pl. Eur. Med. (1840) 60, 219; Hand.-Mazz. in Wettst. Sched. Fl. Exs.
Austro-Hung. X, 63.—C. corymbosum b. clusii (Fisch. ex Rchb.)
Halacsy, Fl. Nied.-Oesterr. (1896) 273.—Tanacetum subcorymbosum
(Schur) Simonk. Enum. FI. Trans. (1886) 312.—T. corymbosum ssp.
subcorymbosum (Schur) B. Pavl. in Pl. Polon. exs. No. 274.—Exs.: FI.
exs. Austro-Hung. No. 3783; Pl. Polon. exs. No. 274; Fl. Polon. exs.
No. 742.

Perennial. Plants 25-65 cm high, with more or less thick, inclined
rhizome, more or less covered with rather long bifid hairs, often mixed
with simple hairs, sometimes subglabrous. Stems solitary or few, erect,
very sparsely leafy, simple or more or less branched in upper part.
Leaves like previous species. Capitula solitary or 2-10 (less often to
15) on a single stem and then in lax corymb. Involucre 8-14 mm in
dia, 4-5 mm long, more or less pubescent at base, less often also,
dorsally; involucral bracts with broader, dark brown, membranous
border, otherwise like previous species. Receptacle much more convex
than in previous species, usually 1.5 times as wide as long. Ligulate
florets white, corolla tube about 1.5 mm long and limb 12-18 mm
long; corolla of tubular florets 1.8—2.5 mm long. Achenes 2.0-2.5 mm
long, about 0.8 mm wide, with 5 (less often up to 7) longitudinal ribs,
and corona more or less toothed or lobed, 0.5—0.8 mm long. Flowering
June to August.

Meadows, forest glades, scrubs above 1,500 m.—European part:
Upper Dniester (Carpathians). General distribution: Central Europe,
Balkans. Described from Carpathians. Type in Leipzig.

Note. A higher mountain ecogeographical race in comparison with
the previous species, related to it through transitional forms.

33. P. corymbiforme Tzvel. sp. nova in Addenda, XXV, 873.—
P. corymbosum auct. non L.: Kar. and Kir. in Bull. Soc. Nat. Mosc.
XV, 382; O. and B. Fedch. Perech. Rast Turk. IV, 187.—P.
corymbosum var. “fol. segmentis profundius divisis, pappo coroniformi
breviore” Ldb. Fl. Ross. IT (1845-1846) 551.

Perennial. Plants 30-100 cm high, with more or less thick inclined
rhizome, usually covered with rather long adpressed simple hairs, often
subglabrous. Stems solitary or few, erect, relatively sparsely leafy,
usually more or less branched only in upper part, less often simple.
Leaves green, usually very sparsely hairy, glabrous or subglabrous

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226

above, sparsely hairy beneath, lacking distinct punctate-glandular hairs;
basal and lower cauline leaves 20-25 cm long and 6-8 cm wide, on
rather long (but always shorter than lamina) petioles; lamina oblong to
broadly linear, usually gradually narrowed toward base, pinnately cut;
leaf segments 8—20 on each side, oblong or lanceolate, pinnately parted
or lobed, terminal segments and lobes usually more or less toothed,
with acuminate teeth; middle and upper cauline leaves strongly reduced,
sessile. Capitula (1)2—6(10) on a single stem, in lax, simple, corymbose-
racemes; peduncles up to 15-18 cm long, glabrous or subglabrous.
Involucre 8—12 mm in dia, 3.5—4.5 mm long, glabrous; involucral bracts
coriaceous-herbaceous, outer lanceolate, inner |.5—2 times as long,
lanceolate-linear; all bracts with very narrow, brownish, membranous
border, somewhat broadened at apex in inner bracts. Ligulate florets
white, corolla tube 1.0-1.5 mm long and limb 9-15 mm long; corolla
of tubular florets 2.0—-2.5 mm long. Achenes 2.0—2.5 mm long, with 5—
8 longitudinal ribs and 0.3-0.5 mm long corona, usually irregularly
lobed and finely toothed almost up to middle. Flowering June to August.

Meadows, scrubs, forest glades in middle mountain zone.—Soviet
Central Asia: Dzhungaria-Tarbagatai (Dzhungarian Alatau). General
distribution: Dzhungaria-Kashgaria(?). Described from Dzhungarian
Alatau. Type (“in rupestribus Alatau, ad fl. Sarchen,” Karelin and
Kirillov, Enum. 1841, No 441); isotypes in Leningrad.

Note. Despite great morphological resemblance to the previous
species, this species is quite distinct from them (with a completely
isolated range) by having pubescence of only simple hairs, capitula
usually aggregated in corymbose-racemes and a glabrous involucre, as
well as a shorter and more deeply divided achene corona.

34. P. poteriifolium Ldb. apud Nordm. in Bull. Acad. Sc. Petersb.
II (1837) 312 and FI. Ross. II, 550; Lipsky in Tr. Peterb. Bot. Sada,
XIII, 307; Sosn. in Tr. Tifl. Bot. Sada, XVII, 28; Grossh. Fl. Kavk.
IV, 136 and Opred. Rast. Kavk. (1949) 461; Manden. in Fl. Gruzii,
VIII, 359.—P. corymbosum auct. non Willd.: Boiss. Fl. or. III, 342, p.
p.—P. corymbosum var. oligocephalum Lipsky in Zap. Kievsk. Obshch.
Estestv. XII (1892) 359.—P. ponticum Albov in Bull. Herb. Boiss. II
(1894) 454.—P. starkianum Albov ibid. 454; Sosn. op. cit. 29; Grossh.
op. cit. IV, 134 and (1949) 460; Manden. op. cit. 359.—Chrysanthemum
poteriifolium (Ldb.) Bornm. in Fedde, Repert. LIII (1944) 341.—Ic::
Manden. op. cit. Plates 397 and 396.—Exs.: Fl. cauc. exs. No. 24.

Perennial. Plants 15-60 cm high, with thick inclined rhizome, dull-
or grayish-green from dense pubescence of rather long bifid and simple
hairs. Stems solitary or few, erect, sparsely leafy, simple or with few
lateral branches above. Leaves dull- or grayish-green from rather

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227

extensive, somewhat sparse pubescence, often more or less tomentose
beneath, with inconspicuous punctate-glandular hairs; basal leaves 15
cm long, 4 cm wide, on rather long (but shorter than lamina) petioles;
lamina oblong or oblong-linear, pinnately cut up to wingless or almost
wingless axis; leaf segments 5-12 on each side, broadly ovate to oblong,
sessile, pinnately parted or lobed; terminal segments and lobes broadly
lanceolate to oblong, up to 3 mm wide, usually more or less toothed
or lobed; lower cauline leaves like basal, middle and upper strongly
reduced, sessile. Capitula solitary or 2—4(6) on a single stem, and then
on very long (up to 20 cm) peduncles in lax corymb. Involucre 9-16
mm in dia and 4-6 mm long, rather densely adpressed hairy almost
entirely, less often sparsely hairy; involucral bracts coriaceous-
herbaceous, outer lanceolate, inner 1.5, less often 2 times as long,
broadly linear; all bracts with very narrow, light colored or brownish
membranous border, somewhat broadened at apex in inner bracts, more
or less lacking in outer. Ligulate florets white, corolla tube 1.0—1.5
mm long and limb 9-13 mm long; corolla of tubular florets 1.8-2.4
mm lorg. Achenes 2.4—3.0 mm long and about 0.8 mm wide, with 5—
8 longitudinal ribs; corona 0.8—1.5 mm long, more or less lobed and
irregularly sharply toothed. Flowering June to August.

Stony slopes, meadows, forest glades up to upper mountain zone.—
Caucasus: Ciscaucasia (western part), Western Transcaucasia
(Abkhazian ASSR). Endemic. Described from Abkhazian ASSR. Type
and isotypes in Leningrad.

Note. In the nature of the pubescence and the shape of the leaves,
the type [? typical] specimens of P. starckianum Albovy are entirely
similar to the type of P. poteriifolium Ldb., which also has only a
single capitulum and, according to the author of this species, was also
collected from the subalpine zone. These species should be considered
entirely synonymous, as noted already by Lipsky (FI. Kavk. 1899,
349); however, until recently they were considered separate species,
and, without any justification, only larger specimens with sparse pu-
bescence and few capitula on a single stem were included under P.
poteriifolium Ldb. Such specimens seem to occupy an intermediate
position between P. poteriifolium and P. corymbosum, but they hardly
deserve to be treated as a separate species, which in this case should
be called, not P. poteriifolium Ldb., but P. ponticum Albov.

The relatively rare, larger but densely hairy specimens of the
species—P. starckianum var. foliosa Albov [Prodr. Fl. Colch. (1895)
137], hardly have any taxonomic significance.

Section 11. Richteriopsis Tzvel. sect. nova in Addenda, XXV,
872.—Herbaceous plants, 15-35 cm high, densely covered with

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228

appressed bifid hairs, often mixed with simple, weakly leafy stems and
well developed, short, nonflowering shoots. Leaves twice pinnately
cut, lower more or less long-petiolate, middle and upper sessile. Capitula
solitary, less often 2—8 on a single stem, on long leafy peduncles, not
in regular corymbs; involucre 9-20 mm in dia, narrow-patelliform;
receptacle weakly convex; corolla limb of ligulate florets oblong or
oblong-linear, 7-20 mm long; corolla of tubular florets 2.2-4.0 mm
long, with long and narrow tube, slightly and gradually expanded in
upper half, with deltoid, slightly recurved teeth. Achenes 1.8-—3.2 mm
long, with 8—12 longitudinal ribs; corona 0.1—0.3 mm long, irregularly
toothed or lobed up to middle, less often almost up to base; joint
between corolla and achene inconspicuous in ligulate florets, usually
with very short corona, and ribs more or less ventrally shifted.

Type of section: Pyrethrum sericeum (Adam) M.B.

Note. Relatively few (about 8-10) species of this section, distrib-
uted in the countries of Western Asia (especially in Turkey) and in the
Caucasus, morphologically resemble some species of section Richteria,
differing from them, however, by a weakly convex receptacle, pubes-
cence almost exclusively of bifid hairs, and very short achene corona.
Section Richteriopsis differs from the equally close section Pyrethrum
by having achenes usually with 10 (and not 5) ribs and a very short
corona, weakly convex receptacle, dense pubescence of shorter hairs,
and twice pinnately cut leaves with very narrow lobes.

Series 1. Sericea Sosn. in Tr. Tifl. Bot. Sada, XVII (1915) 17.—
Outer and usually also middle involucral bracts lanceolate, acuminate,
almost lacking membranous border.

35. P. sericeum (Adam) MB. FI. taur.-cauc. II (1808) 323 and III
(1819) 577; DC. Prodr. VI, 55; Ldb. Fl. Ross. II, 551; Boiss. Fl. or.
III, 339 p. p.; Sosn. in Tr. Tifl. Bot. Sada, XVII, 17; Grossh. Fl. Kavk.
IV, 135 and Opred. Rast. Kavk. 461.—Chrysanthemum sericeum Adam
in Web. and Mohr. Beitr. I (1805) 69, non Hoffmagg. and Link, 1820.—
Tanacetum sericeum (Adam) Sch. Bip. Tanacet. (1844) 59.—Ic.: MB.
Cent. Pl. rar. ruth. II, 3 (1843) t. 78.—Exs.: Fl. cauc. exs. No. 325; Pl.
or. exs. No. 299.

Perennial. Plants 15-35 cm high, with thick, more or less branched
rhizome, sericeous-grayish (but turning brownish on drying of the
pubescence) from dense pubescence of long and short bifid hairs,
usually mixed with simple. Stems few or solitary, erect or ascending
at base, densely hairy, simple. Leaves grayish (turning brown when
dry) from dense pubescence, lacking distinct punctate glandular hairs;
basal leaves up to 20(25) cm long and 3(4) cm wide, on rather long

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2z9

(but shorter than lamina), densely villous petiole, thickened at base;
lamina broadly linear or linear-oblong, twice pinnately cut; terminal
segments narrow-linear to linear-lanceolate, up to 1.5 mm wide,
gradually narrowed apically into short cusp; cauline leaves few, strongly
reduced, lower short-petiolate, others sessile. Capitula solitary, on
peduncles up to 10-15 cm long. Involucre 9-16 mm in dia, 7-10 mm
long, densely villous; involucral bracts coriaceous-herbaceous, outer
narrow-lanceolate, acuminate, with narrow membranous border only in
upper part, inner more or less 1.5 times as long, linear, or oblong-
linear, with narrow, brownish, membranous border irregularly fimbriate
on margin and appendiculately broadened at apex. Corolla of ligulate
florets white, tube 2.0—-2.5 mm long and limb 12-20 mm long; corolla
of tubular florets 3-4 mm long. Achenes 2.5—3.2 mm long and 0.6-0.8
mm wide, with 9-12 longitudinal ribs; corona 0.1—-0.2 mm long,
irregularly blunt-toothed almost up to base. Flowering June to July.

Stony and rubbly slopes, rocks in lower and middle mountain
zones.—Caucasus: Eastern Transcaucasia (eastern part), Western
Transcaucasia (eastern part), Southern Transcaucasia (western and
northern parts). Endemic. Described from Transcaucasia. Type and
isotypes in Leningrad.

36. P. oxylepis (Bordz.) Tzvel. comb. nova.—P. aucherianum auct.
non DC.: Sosn. in Tr. Tifl. Bot. Sada, XVII (1915) 18; Grossh. FI.
Kavk. IV, 136.—P. aucherianum var. oxylepis Bordz. in Zap. Kievsk.
Obshch. Estestv. XXV, 1 (1915) 122.—Chrysanthemum oxylepis Bordz.
ibid. 122 in syn.—Exs.: Sintenis, Iter. orient. No. 5977.

Perennial. Plants 15-30 cm high, with thick, more or less branched
rhizome, grayish from dense pubescence of bifid (mixed with simple)
hairs. Stems usually rather numerous, less often solitary, erect or
ascending at base, more or less branched from middle or slightly below,
less often simple. Leaves grayish from dense pubescence, with numerous
inconspicuous, punctate-glandular hairs; basal leaves up to 10-12 cm
long and 2.0(2.5) cm wide, on rather long, basally strongly thickened
petiole; lamina oblong to broadly linear, twice pinnately cut; terminal
segments narrow-linear to linear-lanceolate, up to 0.8 mm wide, with
short cartilaginous cusp; middle and upper cauline leaves strongly
reduced, sessile. Capitula (1)2—5(8) on each stem, usually not in regular
corymbs; peduncles of capitula up to 15 cm long, densely hairy. In-
volucre 9-12 mm in dia, about 5 mm long, rather densely adpressed
hairy; involucral bracts coriaceous-herbaceous, outer broadly lanceolate,
acuminate, almost lacking border, inner 1.5, less often twice as long, oblong
or oblong-linear, with narrow, brownish membranous border in upper part,
terminating into narrow and acute appendage at apex. Ligulate florets

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230

white, corolla tube about 1.8 mm long and limb 7—12 mm long; corolla of
tubular florets 2.2-2.8 mm long. Achenes 2.2—2.6 mm long and about
0.6 mm wide, with 8-10 longitudinal ribs; corona 0.1-0.2 mm long,
irregularly blunt-toothed almost up to base. Flowering June to July.

Rocks and stony slopes of middle mountain zone.—Caucasus:
Likely to be found in Southern Transcaucasia. General distribution:
Armenia-Kurdistan. Described from Turkey. Type in Kiev.

Note. The species is common in Turkey near Kagizman close to
the USSR border, and its occurrence in Armenia or southern Georgia
is very probable. D.I. Sosnowsky (op. cit.) erroneously identified this
species with P. aucherianum. In fact, it is inuch more similar to the
previous species, with which it is also related through P. gracile Sosn.
of this series [in Vestn. Tifl. Bot. Sada, XI (1915) 17 and in Tr. Tifl.
Bot. Sada, XVII (1915) 17], described from the region of Oltu in
Turkey. P. gracile is similar to P. oxylepis in the nature of the
pubescence, but has larger solitary capitula, as in P. sericeum. The
reports of P. sericeum from Turkey (e.g., Boiss. Fl. or. III, 339),
apparently relate to this species.

Series 2. Aucheriana Tzvel.—Outer involucral bracts broadly
linear, obtuse, and like inner bracts, with broad membranous border.

37. P. aucherianum DC. Prodr. VII (1838) 297; Boiss. FI. or. III,
340.—P. leucanthemiflorum Boiss. and Huet in Boiss. Diagn. ser. II,
3 (1856) 28.—P. pseudoanthemis Boiss. and Huet ibid. (1856) 28.—
P. roopianum (Bordz.) Sosn. in Tr. Tifl. Bot. Sada, XVII (1915) 19;
Grossh. Fl. Kavk. IV, 136.—Matricaria argentea L. Sp. pl. (1763)
1256.—Chrysanthemum argenteum (L.) Willd. Sp. pl. HI, 3 (1803)
2146, non Pyrethrum argenteum Boiss. 1875.—C. roopianum Bordz.
in Tr. Bot. Sada Yur’evsk. Univ. XIII (1912) 23.—C. aucherianum
var. roopianum (Bordz.) Bordz. in Zap. Kievsk. Obshch. Estestv., XXV,
1 (1915) 121.—Tanacetum aucherianum (DC.) Sch. Bip. Tanacet.
(1844) 49.—T. tournefortii Sch. Bip. ibid. 59, nom. nov. (= Matricaria
argentea L.).

Perennial. Plants 15-35 cm high, with thick, more or less branched
rhizome, grayish from dense pubescence of bifid (mixed with simple)
hairs. Stems few or solitary, erect or ascending at base, simple, less
often with 1—2 lateral branches from middle. Leaves grayish from dense
pubescence, with inconspicuous punctate-glandular hairs; basal leaves
10-12 cm long, 2.5 cm wide, on rather long petiole; lamina oblong to
broadly linear, twice pinnately cut, terminal segments linear to broadly
lanceolate, subacute; middle and upper cauline leaves strongly reduced,
sessile. Capitula solitary, less often 2-3 on each stem; peduncles of

—

231

capitula 10-15 cm long, densely hairy. Involucre 9-12 mm in dia, 4—
5 mm long, usually more or less hairy only near base; involucral bracts
coriaceous-herbaceous, outer broadly ovate, obtuse, inner 1.5—2 times
as long, oblong; all bracts with broad, lighter colored or brownish,
membranous border appendiculately broadened at apex in inner bracts.
Ligulate florets white, corolla tube 1.5—2.0 mm long and limb 7-12
mm long; corolla of tubular florets 2.2-2.8 mm long. Achenes 2.2-2.8
mm long, about 0.6—-0.8 mm wide, with 8-10 longitudinal ribs; corona
0.1—0.2 mm long, more or less irregularly blunt-toothed along margin.
Flowering July to August.

Rocks and stony slopes.—Caucasus: Likely to be found in Southern
Transcaucasia. General distribution: Armenia-Kurdistan. Described
from Turkey. Type in Geneva.

Note. Judging from the garden specimens (particularly those sent
by Willdenow) available in the Herbarium of the Botanical Institute,
Academy of Sciences of the USSR, Matricaria argentea L., usually
considered a synonym of P. sericeum, also belongs to this species. The
description of Chrysanthemum roopianum Bordz. is mainly the result of
the imprecise identification of P. oxylepis as “P. aucherianum DC.” Later,
the author himself included C. roopianum as a variety of this latter species,
distinguishing it from typical P. aucherianum only by smaller capitula.
The last character should not be given much importance, as the
specimens of C. roopianum were collected in nonflowering condition.

Section 12. Brachyglossa Tzvel. sect. nova in Addenda, XXV,
872.—Usually basally more or less woody 10-15 cm high plants, usually
covered with rather long bifid and simple or only simple hairs, with
sparsely leafy stem and well developed, short, nonflowering shoots.
Leaves pinnately cut, with more or less pinnately lobed or pinnately
parted segments; lower leaves petiolate, upper sessile or subsessile.
Capitula solitary, less often 2—5, not forming regular corymbs; involucre
cupuliform, 6-13 mm in dia; receptacle weakly convex, usually almost
flat; ligulate florets fewer; ligule oval, less often oblong, 6-12 mm
long; corolla of tubular florets 2.5—3.2 mm long, with tube slightly and
gradually broadening in upper part, with deltoid, weakly recurved teeth.
Achenes 2.3-3.2 mm long, with 6—8 longitudinal ribs; corona 0.2—0.6
mm long, more or less lobed and toothed up to middle, less often
almost up to base, often strongly reduced in achenes of ligulate florets.

Type of section: Pyrethrum hissaricum Krasch.

Note. An endemic Central Asia section, represented by four nar-
rowly endemic species not showing close affinity with each other.
Particularly distinct is the species P. semenovii, probably deserving
separation as a monotypic section. Despite these differences, the species

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232

of this section have several features in common (fewer ligulate florets
with a more or less reduced ligule that, apparently, have a tendency to
disappear, slightly convex, almost flat receptacle, cupuliform involu-
cre, relatively weakly dissected leaves, etc.), which differentiate them
quite well from other sections of the genus.

38. P. hissaricum Krasch. in Fedde, Repert, XX VI (1929) 26.—
Chrysanthemum hissaricum (Krasch.) Bornm. in Fedde, Repert,
LXXXIX (1944) 339 in obs.

Perennial. Plants 20-40 cm high, with thick woody rhizome, grayish
from dense tomentum of flexuous bifid and simple hairs. Stems usually
rather numerous, less often solitary, erect or ascending, weakly leafy,
branched only near somewhat woody base. Leaves grayish from dense
tomentum, with inconspicuous glandular hairs under tomentum; basal
leaves up to 8-10 cm long, 2.2 cm wide, on rather long (but shorter
than lamina) petiole, strongly thickened at base in almost coriaceous
sheath; lamina oblong or broadly linear, pinnately parted or pinnately
cut up to narrow-winged axis; leaf segments numerous, sessile, oblong
or linear-oblong, pinnately lobed or pinnately parted in fewer, subobtuse
lobes and 0.6-3.0 mm long and 2-3 mm wide lobules; few cauline
leaves sessile or subsessile, strongly reduced, pinnately parted or
pinnately lobed. Capitula solitary, on long, more or less tomentose
peduncles. Involucre 8-13 mm in dia and 4-6 mm long, adpressed-
hairy; involucral bracts coriaceous-herbaceous, outer deltoid-lanceolate,
inner 1.5—2 times as long, linear-lanceolate; all bracts with rather wide,
brownish membranous border. Ligulate florets white, corolla tube 1.2—
1.5 mm long and limb 6-10 mm long and 2.5-4.5 mm wide; corolla
of tubular florets 2.6—3.2 mm long. Achenes 2.4-3 mm long and 0.6—
0.8 mm wide, with 6-8 prominent ribs; corona 0.3—0.6 mm long, cut
almost to base in irregularly toothed (with subulately pointed teeth)
lobe of various shades and size, strongly reduced in achenes of ligulate
florets, often completely absent. Flowering July to August.

Rocks and stony slopes above 2,000 m.—Soviet Central Asia:
Pamiro-Alai (Hissar Range). Endemic. Described from Hissar Range.
Type and isotypes in Leningrad.

39. P. galae M. Pop. in Tr. Turkest. Nauchn. Obshch. I (1923) 40;
Chrysanthemum galae M. Pop. ibid. 41, nom. altern.—Tanacetum galae
(M. Pop.) Nevski in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, IV (1937)
216.—Ic.: M. Pop. |. c. t. 6.

Perennial. Plants 20-50 cm high, with thick woody rhizome, usually
covered with adpressed and erect bifid hairs, mixed with simple. Stems
usually rather numerous, less often solitary, arising from branches of

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233

more or less woody caudex, weakly leafy, simple or with few (1-5)
branches, arising from axils of cauline leaves. Leaves dull green from
rather dense pubescence, with inconspicuous punctate-glandular hairs;
basal leaves 8-10 cm long and up to 2(2.5) cm wide, on rather long
(but shorter than lamina) petiole, strongly thickened at base; lamina
oblong or lanceolate-oblong, cuneately narrowed toward base,
irregularly pinnately parted (less often lobed); leaf segments 3-6 on
each side, oblong or lanceolate, usually with large irregular teeth or
pinnately lobed; few cauline leaves strongly reduced, sessile or
subsessile, usually pinnately lobed. Capitula solitary or 2—6 on long,
more or less pubescent peduncles, not in regular corymbs. Involucre
6-10 mm in dia and 6-8 mm long, more or less pubescent; involucral
bracts coriaceous-herbaceous, outer lanceolate-ovate, inner 1.5—2 times
as long, ovate-oblong; all bracts with narrow, brownish membranous
border, appendiculately broadened at apex in inner bracts. Ligulate
florets white, corolla tube 1.0-1.5 mm long, and limb 8-12 mm long
and 2.5-4.0 mm wide; corolla of tubular florets 2.5-3.0 mm long.
Achenes 2.5—3.2 mm long, 0.7—1 mm wide, with 6—8 longitudinal ribs;
corona 0.2—0.3 mm long, cut almost or right to base into acute lobes
of various shape and size; achenes of ligulate florets with shorter,
usually unilateral corona. Flowering June to July.

Sandstone and limestone rocks, dry gypsic hills in lower and middle
mountain zone.—Soviet Central Asia: Pamiro-Alai (western part of
Hissar Range). Endemic. Described from Hissar Range (Baisuntau
Mountain). Type in Tashkent; isotype in Leningrad.

40. P. mikeschinii Tzvel. sp. nova. in Addenda, XXV, 873.

Perennial. Plants 20—30 cm high, with thick woody rhizome, grayish
from rather dense pubescence of semi-erect bifid (mixed with simple)
hairs. Stems usually rather numerous, less often solitary, erect or
ascending, weakly leafy branched only near more or less woody base.
Leaves grayish-green from rather dense pubescence, with inconspicuous
punctate-glandular hairs; basal leaves up to 10 cm long and 1.3 cm
wide, on rather long (but shorter than lamina) petiole, strongly thickened
at base and almost coriaceous; lamina broadly linear, but pinnately cut
almost up to wingless axils; leaf segments oblong, sessile or subsessile,
more or less pinnately parted, sometimes partly entire or pinnately
lobed; terminal lobes linear-lanceolate, up to 1.2 mm wide, narrowed
into cartilaginous cusp; few (3-5) cauline leaves strongly reduced,
middle and upper sessile. Capitula solitary, on long, densely hairy
peduncle. Involucre 6-8 mm in dia and 5-7 mm long, sparsely
tomentose near base; involucral bracts coriaceous-herbaceous, outer
lanceolate-ovate, inner 1.5—2 times as long, oblong-lanceolate; all bracts

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234

with rather wide, brownish, membranous border, somewhat broadened
at apex in inner bracts. Fully developed ligulate and tubular florets
lacking (plant was collected before flowering). Mature achenes
unknown; corona usually about 0.2 mm long, lobed almost to base.
Flowering June to July.

Stony slopes, steppes above 2,000 m.—Soviet Central Asia: Pamiro-
Alai (Turkistan Range). Endemic. Described from Turkestan Range,
upper reaches of Isfara River 50 km south of the village of Vorukh,
northern slope in valley of tributary of Dzhiptyk above mouth of
Tamingen River, sheep’s fescue steppe, about 2,900 m (22. VI. 1938,
No. 58, G. Mikeshin). Type in Leningrad.

Note. Like the previous species, this species does not show close
affinity with other species. Morphologically, it slightly resembles
Pyrethrum pyrethroides of section Richteria. However, the nature of
the pubescence and the less dissected leaves, almost flat receptacle,
and more or less woody branches of the caudex compel us to consider
it much closer to the species of section Brachyglossa, especially P.
hissaricum and P. galae. The only two specimens of this species at our
disposal, from almost the same locality, were collected before flowering;
as a result, the details of floral and achene structure cannot be
established precisely.

41. P. semenovii (Herd.) Winkl. ex O. and B. Fedtsch. Perech.
Rast. Turk. IV (1911) 186.—Tanacetum semenovii Herd. in Bull. Soc.
Nat. Mosc. XL, | (1867) 130.—Chrysanthemum semenovii (Herd.) B.
Fedtsch. Rast. Turk. (1915) 737.

Perennial. Plants 10-35 cm high, with thick many headed rhizome,
grayish or even whitish from extensive loose tomentum of simple
flexuous hairs. Stems usually numerous, arising from more or less
woody branches of caudex, erect or ascending at base, weakly leafy,
simple. Leaves grayish or whitish-tomentose, with few distinct punctate-
glandular hairs; basal leaves usually up to 4 cm long and 0.8 cm wide,
less often (in shady specimens) up to 10-12 cm long and 2 cm wide,
on rather long (but shorter than lamina) petiole, strongly thickened at
base and with patches of axillary tomentum; lamina of basal leaves
oblong to oblong-linear, pinnately cut; leaf segments rather numerous,
sessile or short-petiolate, oblong, usually pinnately lobed or parted,
with few short lobes or segments up to 1.0—-1.5 mm wide, subacute;
few (1-3) cauline leaves strongly reduced, sessile, more or less pinnately
parted to entire. Capitula solitary (but numerous on a single plant), on
long tomentose peduncles. Involucre 7-11 mm in dia and 5-7 mm
long, more or less loosely tomentose; involucral bracts coriaceous-
herbaceous, outer lanceolate-ovate, inner 2—3 times as long, oblong;

235

245 Plate X.
1 — Spathipappus griffithii (Clarke) Tzvel., habit of plant, ligulate floret, corolla
of tubular floret, and achene; 2 — Pyrethrum semenovii (Herd.) Winkl. ex O.
and B. Fedtsch., habit of plant, leaf, corolla of ligulate floret, corona of tubular
floret, and achene.

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236

all bracts with rather wide erose-toothed light brown membranous
border. Ligulate florets white or pink, corolla tube about 1.2—-1.5 mm
long, limb 6-10 mm long and 2.5—4.0 mm wide; corolla of tubular
florets 2.5-3.0 mm long. Achenes 2.3—3.0 mm long, 0.6—0.8 mm across,
with 6-8 longitudinal ribs and more or less irregularly toothed, 0.2—
0.6 mm long corona. Flowering June to July. (Plate X, Fig. 2.)

Rocks and stony slopes in middle mountain zone.—Soviet Central
Asia: Tien Shan (Ketmen and Trans-Ili Alatau ranges), Lake Balkhash
Region (Syugaty Mountains). Endemic. Described from Trans-Ili Alatau.
Type and isotype in Leningrad.

Section 13. Trichanthemopsis Tzvel. sect. nova in Addenda, XXV,
872.—Plants more or less woody at base, 6-25 cm high, usually covered
with erect simple hairs or completely glabrous, with weakly leafy stems
and well developed, short (but often elongated base) nonflowering
shoots. Leaves more or less pinnately cut, with very narrow lobes;
basal leaves petiolate, cauline sessile. Capitula solitary (but several on
single plant); involucre narrow-patelliform, 10—16 mm in dia; receptacle
strongly convex, hemispherical; limb of corolla in ligulate florets
oblong-oval, 6-15 mm long; corolla of tubular florets 2.5-3.2 mm
long, with tube somewhat gradually broadened in upper half, with
deltoid, weakly recurved teeth. Achenes 2.5—3.5 mm long, with 5-10
longitudinal ribs; corona 1.2—2.5 mm long, almost or to base divided
into 6-10 more or less toothed scales or lobes of various shape and
size.

A monotypic section. ;

Note. The single species of this section is close to the species of
the next section, Richteria, in many respects, differing from them only
by the more or less woody bases of the branches. However, it is equally
close to some species of the genus Trichanthemis Rgl. and Schmalh.
(T. butkovii and T. litwinowii), and only the completely glabrous achenes
provide a basis to place it in the genus Pyrethrum and not in
Trichanthemis. The dual, not entirely clear taxonomic position of this
species is the main reason for placing it in a separate monotypic section.

42. P. tianschanicum Krasch. in Tr. Bot. Inst. Akad. Nauk SSSR,
Ser. 1.1 (1933) 176.—P. pyrethroides var. glabrum B. Fedtsch. ex
Krasch. ibid. 176, in syn.

Perennial. Plants 6—25 cm high, with thick woody rhizome, more
or less pubescent, often completely glabrous. Stems numerous from
branched, more or less woody caudex, usually caespitose with short
(but with elongate base) nonflowering shoots, erect or ascending near
base, sparsely leafy mainly in lower part, simple. Leaves green, less

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237

often pubescent to completely glabrous, lacking distinct punctate-
glandular hairs; basal leaves 5-8 cm long and up to 1.0-1.5 cm wide,
on rather long (but shorter than lamina) petioles strongly thickened at
base; lamina oblong-linear to oblong, pinnately cut; leaf segments
gradually reduced toward lamina base, in turn more or less palmately
cut into linear-filiform (0.2—0.4, less often up to 0.6 mm wide), 2—10
subacute lobes; cauline leaves strongly reduced, sessile, often pinnately
cut. Capitula solitary, on rather long, more or less pubescent peduncle,
but usually numerous on a single plant. Involucre 10-16 mm in dia, 5—
8 mm long, more or less pubescent at base, often with fine tomentum;
involucral bracts coriaceous-herbaceous, outer lanceolate, inner 1.5—2
times as long, oblong-linear; all bracts with rather wide, brownish or
dark-brown membranous border. Ligulate florets white, corolla tube
1.5-2 mm long, limb 6-15 mm long; corolla of tubular florets 2.5—3.2
mm long. Achenes 2.5—3.5 mm long and 0.6—0.8 mm wide with 6-10
longitudinal ribs; corona 1.2—2.5 mm long, almost or right up to base
divided in 6-10 oblong or linear-oblong scales, irregularly toothed at
apex. Flowering July to August. (Plate IX, Fig. 1.)

Rocks and stony slopes 2,000 m.—Soviet Central Asia: Tien Shan
(Ugam, Pskem and Talass Alatau ranges). Endemic. Described from
Talass Alatau. Type in Leningrad.

Note. The only specimen in the herbarium of the Botanical Institute,
Academy of Sciences of the USSR, from the Pskem River Basin (upper
reaches of Ispap-Sai, subalpine zone, 1940, No. 24, O. Knorring) has less
dissected leaves (pinnatisect, with more or less lobate segments) with
broader lobes and, perhaps, belongs to a unique ecogeographical race.

Section 14. Richteria (Kar. and Kir.) Tzvel. comb. nova.—
Richteria Kar. and Kir. in Bull. Soc. Nat. Mosc. XV (1842) 126.—
Chrysanthemum sect. Richteria (Kar. and Kir.) Clarke, Compos. ind.
(1876) 146 p. p.; B.*Fedtsch. Rast. Turk: (1915)'737.—C. sect.
Pyrethrum subsect. Richteria (Kar. and Kir.) Ling in Contr. Inst. Bot.
Nat. Acad. Peiping, III(1935) 480.—Herbaceous plants, (3)8—-40(70)
em high, usually covered with rather long and short simple hairs,
sometimes completely glabrous, with weakly leafy stem and well-
developed short, nonflowering shoots. Leaves more or less (usually
twice) pinnately cut, with very narrow lobes, lower petiolate, middle
and upper sessile. Capitula solitary, less often 2-8 on a single stem,
and then usually not in regular corymbs; involucre patelliform or
narrow-patelliform, 8-25 mm in dia; receptacle strongly convex,
hemispherical; corolla limb of ligulate florets oblong-ovate, broadly
linear and linear, 7-25 mm long; corolla of tubular florets 2.0-3.5 mm
long, with somewhat gradually (rarely abruptly) broadened tube in

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238

upper half and weakly recurved deltoid teeth. Achenes 1.8-3.0 mm
long, with 5-10 longitudinal ribs; corona 0.3—1.5 mm long, usually cut
to or almost to base (less often up to middle) in lobes or scales of
various shape and size, more or less corresponding to number of ribs.

Type of section: Pyrethrum pyrethroides (Kar. and Kir.) B. Fedtsch.
ex Krasch.

Note. It is one of the largest sections of the genus, comprising
about 25-30 species distributed in mountainous regions of Central and
Soviet Central Asia as well as in some regions of southern Siberia.
The species in this section divide themselves in two groups, which
perhaps deserve sectional rank: Richteria proper (series Pyrethroides,
almost always with a longer achene corona cut to or almost to base,
and the species with a more northern and northeastern range which are
larger on the average, with a shorter and usually also less dissected
achene corona, in habit strongly resembling species of the genus
Tripleurospermum Sch. Bip. However, a sharp morphological boundary
cannot be drawn between these groups, and, besides, there is reason to
consider the second group the progenitor of the species of the first
group, which, itself, apparently, appeared as a result of adaptation to
high-mountain conditions combined with a significantly dry climate.
The relationship of this section to other sections of the genus is quite
clear, and splitting it off as a separate genus would hardly be justified.

Series 1. Abrotanifolia Tzvel.—Plants 25-70 cm high, with thrice
pinnately cut (rarely partly twice pinnately cut) lamina of basal leaves.
Capitula 1-8 on a single stem. Corona of achenes 0.3—1.0 mm long,
irregularly toothed or lobate usually not more than to middle, less
often almost to base.

43. P. alatavicum (Herd.) O. and B. Fedtsch. in Perech. Rast.
Turk. IV (1911) 186; Kryl. Fl. Zap. Sib. XI, 2748.—Tanacetum
alatavicum Herd. in Bull. Soc. Nat. Mosc. XL, 2 (1867) 129.—
Chrysanthemum alatavicum (Herd.) B. Fedtsch. Rast. Turk. (1915)
737.—Exs.: GRF No. 3263.

Perennial. Plants 25-70 cm high, with more or less thick oblique
rhizome, sparsely covered with long, simple and short glandular hairs,
sometimes glabrous. Stems few or solitary, erect, more weakly leafy,
usually more or less branched above, less often simple. Leaves green,
more or less pubescent to glabrous, lacking punctate-glandular hairs;
basal and lower cauline leaves up to 18-20 cm long and 4 cm wide,
on rather long (but shorter than lamina), petioles strongly thickened at
base; lamina linear-oblong, 3- or partly twice pinnately cut, with 7-12
segments on each side; terminal segment linear-filiform to linear-

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lanceolate, up to 1 mm wide, gradually narrowed into rather long cusp;
other cauline leaves smaller and with shorter, less dissected lamina,
sessile. Capitula usually 2-6, on long, more or less pubescent peduncles,
in axils of upper cauline leaves, usually not forming regular corymb.
Involucre 10-18 mm in dia, 4.5-6 mm long, more or less pubescent to
subglabrous; involucral bracts herbaceous, outer linear or linear-
lanceolate, inner 1.5—2 times as long, oblong-linear; all bracts with
rather wide, blackish-brown membranous border. Ligulate flortes white,
corolla tube 1-2 mm long, limb 14-22 mm long, perhaps, less often
bearing fruits; corolla of tubular florets 2-3 mm long. Achenes 2.2-
3.2 mm long and about 0.8 mm wide, with 5-7 longitudinal ribs; corona
0.5-1.0 mm long, cut almost to base in irregularly toothed lobes or
scales. Flowering July to August.

Stony slopes, meadows above 2,500 m.—Western Siberia: Altai
(southwestern part); Soviet Central Asia: Dzhungaria-Tarbagatai, Tien
Shan (central part and Chatkal Range), Pamiro-Alai (Alai Range).
General distribution: Dzhungaria-Kashgaria. Described from Trans-Ili
Alatau. Type in Leningrad.

44. P. krylovianum Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, IX (1946) 155; Kryl. Fl. Zap. Sib. XI, 2746.—
Chrysanthemum abrotanifolium (Bge.) Kryl. Fl. Alt. III (1904) 621
p. p.—lIc.: Krasch. op. cit. 156, Fig. 1.

Perennial. Plants 25-70 cm high, with more or less thick oblique
rhizome, sparsely covered with long, simple hairs, mixed with shorter
glandular hairs, sometimes completely glabrous. Stems few or solitary,
erect, more weakly leafy, usually simple, less often with 1-2 lateral
branches. Leaves like previous species but, on the average, narrower.
Capitula solitary, less often 2-3 on a single stem, on long peduncle in
axils of upper leaves. Involucre 10-18 mm in dia, 4.5—6.0 mm long,
more or less pubescent or subglabrous; involucral bracts like previous
species. Ligulate florets white, corolla tube 1-2 mm long, limb 14—25
mm long; corolla of tubular florets about 2.5 mm long. Achenes 2-3
mm long and about 0.8 mm wide, with 5—7 longitudinal ribs and more
or less toothed 0.4-0.7 mm long corona. Flowering July to August.

Stony slopes and rocks of upper mountain zone.—Western Siberia:
Altai (mainly northern and western parts). Endemic. Described from
Altai. Type and isotype in Leningrad.

Note. It differs from the previous species in only one, not always
consistent character; the.capitula are almost always solitary, which,
with the isolated range, can still serve as a basis for segregating P.
krylovianum as a separate, though morphologically very weakly isolated,
race.

240

45. P. abrotanifolium Bge. ex Ldb. FI. Ross. II (1844-1846) 549,
non Pourr. ex Willk. and Lge. 1870; Kryl. Fl. Zap. Sib. XI, 2747.—
Chrysanthemum abrotanifolium (Bge.) Kryl. Fl. Alt. III (1904) 621
p. p.

Perennial. Plants 25-45 cm high, with thick oblique rhizome,
usually covered with very short, glandular and long, flexuous, simple
hairs, sometimes subglabrous. Stems few or solitary, erect, more
sparsely leafy, with 2—8 lateral branches mainly in upper half. Leaves
like previous species, but more densely hairy, usually with
predominance of short glandular hairs. Capitula usually 2-8, solitary
at apices of main stem and its lateral branches, not forming regular
corymb, less often solitary. Involucre 8-12 mm in dia, 4-6 mm long,
more or less pubescent at base; involucral bracts herbaceous, outer
lanceolate and linear-lanceolate, inner 1.5—2 times as long, oblong-
linear; all bracts with rather wide, brownish or dark brown, membranous
border. Ligulate florets white, corolla tube 0.8—1 mm long, limb 7-12
mm long; corolla of tubular florets about 2 mm long. Achenes 1.5—2.3
mm long, with 5-7 longitudinal ribs; corona 0.3-0.5 mm long,
irregularly toothed or lobed to middle or slightly deeper. Flowering
July to September.

Stony slopes, meadows at 2,000—3,500 m.—Western Siberia: Altai
(rarely); Soviet Central Asia: Dzhungaria-Tarbagatai. General
distribution: Dzhungaria-Kashgaria (?). Described from Altai. Type
and isotype in Leningrad.

Note. It differs from the previous species by having smaller and
more numerous capitula, as well as denser pubescence.

Series 2. Pulchella Tzvel.—Plants 10—30 cm high; lamina of basal
leaves pinnately cut, with entire or 2—4-parted segments; capitula
solitary, less often 2-3 on a single stem; corona of achenes 0.3-—0.6
mm long, irregularly toothed or lobed up to middle or almost to base.

46. P. pulchellum Turcz. in DC. Prodr. VI (1837) 55; Ldb. FI.
Ross. II, 548; Turcz. Fl. baic.-dahur. I], 46.—P. pectinatum Fisch. ex
Turcz. in Bull. Soc. Nat. Mosc. I (1838) 94, nom. nud.—Tanacetum
pulchellum (Turcz.) Sch. Bip. Tanacet. (1844) 61.

Perennial. Plants 10-30 cm high, with more or less branched,
creeping rhizome, usually covered with long flexuous hairs, often
subglabrous. Stems solitary or few, erect or ascending at base, more or
less hairy in upper part, weakly leafy, simple, less often with 1-2
lateral branches. Leaves green, glabrous, less often pubescent, lacking
punctate-glandular hairs; basal and lower cauline leaves 8—10 cm long,
and up to 1.5—2 cm wide, on rather long (but shorter than lamina),

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241

more or less winged petiole, strongly thickened at base; lamina oblong
or linear-oblong, pinnately cut; leaf segments 6-15 on each side, entire,
narrow-linear (up to | mm wide), acuminate, or 2—4 parted in similar
lobes; middle and upper cauline leaves strongly reduced, sessile, usually
with entire segments. Capitula solitary, less often 2-3 on a single
stem; peduncles long, more or less sparsely tomentose. Involucre 10-
15 mm in dia, 4.0-5.5 mm long, usually covered with whitish tomentum
at base; involucral bracts herbaceous, outer lanceolate-ovate, inner 1.5—
2 times as long, broadly linear; all bracts with very wide, blackish-
brown membranous border. Ligulate florets white, corolla tube about
1 mm long, limb 7-14 mm long, usually sterile because of partly
abortive pistil; corolla of tubular florets 1.8-2.4 mm long. Achenes
2.2-2.6 mm long and 0.6-0.7 mm wide, with 5 (rarely up to 7)
longitudinal ribs; corona 0.3-0.6 mm long, irregularly toothed and
lobed up to middle or almost up to base. Flowering July to August.

Stony slopes, bald peaks, grassy patches, in upper mountain zone.—
Western Siberia: Altai (northeastern part); Eastern Siberia: Angara-
Sayans, Dauria. General distribution: Mongolia (northern part).
Described from Lake Baikal Region. Type and isotypes in Leningrad.

Note. The very strong morphological resemblance of this species
to those of Tripleurospermum Sch. Bip. extends partly also to the
structure of the achenes in the ligulate florets (when developed), which
have prominent, almost winged ribs.

Series 3. Pulchra Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, IX (1946) 158.—Plants 5-35 cm high. Lamina of basal
leaves twice pinnately cut; cauline leaves pinnately cut, with entire or
2-5 parted segments. Capitula solitary, large, less often 2-3 on a single
stem. Corona of achenes 0.3-1.2 mm long, irregularly toothed or lobed
up to middle or almost to base.

47. P. lanuginosum (Sch. Bip. and Herd.) Tzvel. comb. nova.—
Tanacetum lanuginosum Sch. Bip. and Herd. in Pollichia, XX—XXI
(1863) 442.

Perennial. Plants 10-30 cm high, with more or less branched
creeping rhizome, more or less lanate-tomentose with flexuous simple
hairs. Stems solitary or few, but often caespitose together with short
nonflowering shoots, erect, simple, less often with 1-2 lateral branches.
Leaves usually dull or even grayish-green from rather extensive,
somewhat loose tomentum, but often sparsely hairy or subglabrous,
lacking punctate glandular hairs; basal leaves 10-15 cm long and 1.5—
2.0 cm wide, on rather long, more or less winged petiole, strongly
thickened at base and usually (but not always) lanate-tomentose in leaf

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242

axils; lamina linear or oblong-linear, narrowed at both ends, pinnately
cut; leaf segments 7-20 on each side, approximate, sessile or sub-
sessile, in turn pinnately cut or parted; terminal segments narrow-linear
and lanceolate-linear, up to 0.6 mm wide, gradually narrowed toward
apex into rather long cartilaginous cusp; cauline leaves smaller, usually
less dissected and more densely hairy, sessile or subsessile. Capitula
solitary, very rarely 2-3 on a single stem; peduncles rather long,
thickened toward apex and covered with extensive lax tomentum.
Involucre 15-28 mm in dia and 5—7 mm long, usually covered, almost
allover with lax whitish tomentum; involucral bracts herbaceous, outer
lanceolate-linear, inner 1.5—2 times as long, oblong-linear; all bracts
with very wide, blackish-brown, membranous border. Ligulate florets
white, corolla tube 2—2.5 mm long, limb 13-25 mm long narrow-linear,
often sterile; corolla of tubular florets 3—-3.5 mm long. Achenes 2-3
mm long and about 0.8 mm wide, with 5—8 longitudinal ribs; corona
0.4-0.8 mm long, usually irregularly toothed on margin not beyond
middle. Flowering July to August.

Stony slopes, bald peaks in upper mountain zone.—Eastern Siberia:
Angara-Sayans (southeastern part). General distribution: Mongolia
(northern part). Described from the eastern Sayan Range (Munku-Sardyk
Mountain). Type (or isotype?) in Leningrad.

Note. The widely distributed, weakly pubescent individuals of this
species may constitute a special ecogeographical race, which is more
or less intermediate between this and the next species.

48. P. pulchrum Ldb. Ic. pl. Fl. Ross. impr. Alt. I (1829) 20, t.
84 and FI. Alt. IV (1833) 118; DC. Prodr. VI, 56; Ldb. Fl. Ross. II,
548; Kryl. Fl. Zap. Sib. XI, 2745.—Tanacetum pulcherm (Ldb.) Rupr.
in Osten-Saken and Rupr. Sert. tiansch. (1869) 52, quoad nom.—
Chrysanthemum pulchrum (Ldb.) Winkl. in Tr. Peterb. Bot. Sada, X
(1884) 87.—Ic.: Ldb. op. cit. (1829) tab. 84.—Exs.: GRF No. 3264.

Perennial. Plants 10-30 cm high, with more or less branched
creeping rhizome, covered with flexuous simple hairs. Stems solitary
or few, but often caespitose together with short nonflowering shoots,
erect or ascending near base, simple, less often with 1-2 lateral
branches, weakly leafy. Leaves green, glabrous, or subglabrous, less
often covered with fine, somewhat sparse tomentum, lacking distinct
punctate-glandular hairs; basal leaves 10-15 cm long, and 1.5-2 mm
wide, on rather long, more or less winged petiole, always lacking
tomentum at their base; lamina of basal leaves like previous species,
but their segments, on the average, less divided (usually 2—5S-parted),
with broader (up to 1 mm wide) apical lobes; cauline leaves even more
reduced than in previous species, sessile, often with entire segments.

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243

Capitula solitary, very rarely 2-3 on single stem, on long peduncles,
more or less thickened upward and sparsely tomentose. Involucre 15—
28 mm in dia and 5—7 mm long, covered with extensive lax tomentum
near base; involucral bracts like previous species. Ligulate florets white,
corolla tube 2—2.5 mm long, limb 12—25 mm long, linear, on the
average, broader than in previous species; corolla of tubular florets 3—
3.5 mm long. Achenes 2-3 mm long and about 0.8 mm wide, with S5S—
8 longitudinal ribs; corona 0.6—1.2 mm long, usually irregularly toothed
and lobed on margin up to middle. Flowering July to August.

Stony slopes, grassy patches in upper mountain zone.—Western
Siberia: Altai. General distribution: China (northwestern part),
Mongolia (western part). Described from Altai. Type and isotypes in
Leningrad.

Note. One specimen of this species in the Herbarium of the Bo-
tanical Institute, Academy of Sciences of the USSR (the former
Oirot Autonomous Region, source of Kanas River, 2.VIII.1931, B.
Schischkin, L. Chilikina, G. Sumnevich) has a strongly tomentose
pubescence on the entire plant and, possibly, belongs to a special
ecogeographical race. It differs from the previous species in the cutting
of the leaves.

49. P. karelinii Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, IX (1946) 157.—P. pulchrum auct. non Ldb.; O. and B. Fedtsch.
Perech. Rast. Turk. VI (1911) 186.—P. richterioides var. virescens
(Winkl.) O. and B. Fedtsch. ibid. 186.—Chrysanthemum richterioides
var. virescens Winkl. in Tr. Peterb. Bot. Sada, X (1887) 86.—C.
richterioides auct. non Winkl.: B. Fedtsch. Rast. Turk. (1915) 736.

Perennial. Plants 5-35 cm high, with more or less branched,
creeping rhizome, usually covered with long, flexuous simple hairs.
Stems rather numerous, less often solitary, usually caespitose together
with short nonflowering shoots; erect or ascending at base, weakly
leafy, simple. Leaves green or dull green (to almost grayish) from
usually sparse pubescence with or without inconspicuous punctate-
glandular hairs; basal leaves 8-12 cm long and 1-2.5 cm wide, on
rather long, basally thickened petiole, always lacking tomentum in their
axils; lamina of basal leaves oblong or oblong-linear, pinnately cut;
leaf segments sessile or subsessile, pinnately parted or sometimes almost
palmately parted into 5—15 lobes; terminal segments linear or lanceolate-
linear, up to 1 mm wide, subacute; cauline leaves strongly reduced,
sessile or subsessile, usually more or less adpressed to stem, often
with entire segments. Capitula solitary, on rather long, sparsely
tomentose peduncles. Involucre 13-25 mm in dia and 5—7 mm long,
more or less tomentose near base; involucral bracts like previous

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244

species. Ligulate florets white or pinkish-white, corolla tube 1.5—2.0
mm long, limb 12-20 mm long, broadly linear; corolla of tubular florets
2-3 mm long. Achenes 2.0—2.5 mm long, with 5—8 longitudinal ribs;
corona 0.3—0.6 mm long, cut up to middle or almost up to base in
obtuse irregularly toothed lobes more or less corresponding to ribs.
Flowering July to August.

Stony slopes, grassy patches above 3,000 m.—Soviet Central Asia:
Dzhungaria-Tarbagatai (Dzhungarian Alatau), Tien Shan (except
southwestern part), Pamiro-Alai (Alai and Trans-Alai ranges, northern
Pamir). General distribution: Dzhungaria-Kashgaria. Described from
Dzhungarian Alatau. Type and isotypes in Leningrad.

Note. Through transitional forms, it is connected to a very close
eastern Tien Shan species—P. richterioides (Winkl.) Krassn. (Opyt
Ist. Razv. Fl. Vost. Tyan-Shanya* (1888) 346.—Chrysanthemum
richterioides Winkl. op. cit. 86, excl. var.), which differs mainly by
having a more dense pubescence on the entire plant (grayish from thin
tomentum) and pink-colored ligulate florets. In particular, specimens
of P. karelinii are found in Dzhungarian Alatau and the mountains of
Southeastern Kazakhstan, which do not differ from P. richterioides in
their pubescence. The differences cited by Krasheninnikov between
these species in the degree of the incision of the leaves, which, even
on the average barely exist, and the persistence of its pink color of the
flowers need verification.

Series 4. Pyrethroidea Tzvel.—Richteria Kar. and Kir. in Bull.
Soc. Nat. Mosc. XV (1842) 126, sensu stricto.—Plants 3-35 cm high;
lamina of basal leaves (usually also lower cauline leaves) twice
pinnately cut; capitula solitary, less often 2-3 on single stem; corona
of achenes 0.6—-1.5 mm long, usually cut into lobes or scales almost to
base, but sometimes not beyond middle.

50. P. songaricum Tzvel. sp. nova in Addenda, XXV, 874.—P.
pyrethroides ssp. songaricum Krasch. in herb.

Perennial. Plants 15-35 cm high, with more or less branched
rhizomes, grayish from extensive, loose tomentum of long and short
simple hairs. Stems usually few, erect or ascending at base, weakly
leafy, simple or with 1-3 lateral branches. Leaves grayish-green from
dense tomentum, with inconspicuous punctate-glandular hairs; basal
leaves up to 8-10 cm long, 3 cm wide, on rather long (usually not
more than a half as long as lamina), basally thickened petiole; their

*Experiment of Historical Analysis of the Flora of the Eastern Tien Shan—
Translator.

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245

lamina broadly linear to linear-oblong, thrice pinnately cut with linear-
lanceolate to lanceolate-ovate, up to 1 mm wide terminal segments
with short cartilaginous cusp; cauline leaves smaller, sessile, usually
twice pinnately cut. Capitula solitary or 2-4, on long, more or less
tomentose peduncles, in axils of upper cauline leaves, not in corymbs.
Involucre 10-18 mm in dia, 5—8 mm long, usually covered with fine
grayish tomentum; involucral bracts herbaceous, outer lanceolate, inner
usually not more than 1.5 times as long, linear-lanceolate; all bracts
with rather wide, dark brown, membranous border. Ligulate florets
white, corolla tube 1.5—2.0 mm long and limb; 8-14 mm long; corolla
of tubular florets 2.5—3.2 mm long. Achenes 2.5-3.0 mm long and
about 0.6 mm wide, with 6—8 longitudinal ribs; corona 1.0-1.5 mm
long, usually irregularly toothed or lobed not beyond middle. Flowering
June to August. (Plate VIII. Fig. 2.)

Stony slopes, rocks above 2,000 m.—Soviet Central Asia:
Dzhungaria-Tarbagatai (Dzhungarian Alatau), Tien Shan (Kara Tau,
Trans-Ili Alatau). General distribution: Dzhungaria-Kashgaria(?). De-
scribed from Dzhungarian Alatau. Type (Dzhungarian Alatau, Koksu
River Basin near Kuvtas Mountain, 11.VIII.1948, V. Goloskokov) in
Leningrad; isotype in Alma-Ata.

Note. This species, being, apparently, a comparative lower moun-
tain ecogeographical race, differs from P. pyrethroides on the whole
by having a more or less toothed or lobate corona, larger (on the
average) size of the whole plant, and often branched stem.

51. P. pyrethroides (Kar. and Kir.) B. Fedtsch. ex Krasch. in Tr.
Bot. Inst. Akad. Nauk SSSR, Ser. 1, I (1933) 176; Kryl. Fl. Zap. Sib.
XI, 2749.—P. transiliense var. tomentosum Rgl. and Schmalh. and
var. subsessile Rgl. and Schmalh. in Tr. Peterb. Bot. Sada, V, 2 (1878)
618.—Richteria pyrethroides Kar. and Kir. in Bull. Soc. Nat. Mosc.
XV (1842) 127; Ldb. FI. Ross. II, 519; O. and B. Fedtsch. Perech.
Rast. Turk. IV, 177 p. p.—Chrysanthemum richteria Benth. in Benth.
and Hook. f. Gen. pl. II (1873) 426.—C. artemisiaefolium Klatt. in
Sitz. Akad. Muench. (1878) 88.—C. pyrethroides (Kar. and Kir.) B.
Fedtsch. Rast. Turk. (1915) 737.

Perennial. Plants 5—25(35) cm high, with more or less branched
rhizome, grayish from extensive, loose tomentum of long and shorter
simple hairs. Stems usually rather numerous, less often solitary, erect
or ascending, weakly leafy, simple, very rarely with 1-2 lateral
branches. Leaves grayish-green from extensive tomentum, with
inconspicuous punctate-glandular hairs; basal leaves 6—8 cm long, 1.5
cm wide, on rather long (but usually less than a half as long as lamina)
petioles, strongly thickened at'base; their lamina oblong, twice pinnately

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246

cut with lanceolate-linear to lanceolate-ovate, up to 1 mm wide termi-
nal lobes with short cartilaginous cusp, in turn sometimes 2—4 parted;
few cauline leaves smaller and less incised, sessile or subsessile. Ca-
pitula solitary, very rarely 2-3 on long, more or less tomentose pe-
duncle. Involucre 7-15 mm in dia and 4—6 mm long, usually covered
with somewhat loose tomentum; involucral bracts herbaceous, outer
lanceolate, inner 1.5—2 times as long, oblong-linear; all bracts with
wide blackish-brown, membranous border. Ligulate florets white or
pink, corolla tube 1.3—2.0 mm long and limb 7-15 mm long; corolla
of tubular florets 2.5-3.0 mm long. Achenes 2.5—3.2 mm long, about
0.8 mm wide, with 5-9 longitudinal ribs; corona 0.8-1.5 mm long, cut
up to base in oblong or linear-oblong, toothed scales, usually corre-
sponding to number of ribs. Flowering July to August.

Stony slopes, rocks, screes, gravel beds in upper mountain zone.—
Western Siberia: Altai (Kurai Range), Dzhungaria-Tarbagatai, Tien Shan,
Pamiro-Alai Region. General distribution: Dzhungaria-Kashgaria, Tibet,
India-Himalayas (nothern part), Iran-Afghanistan (northeastern part).
Described from Dzhungarian Alatau. Type and isotypes in Leningrad.

Note. This widely distributed species, even after the separation of
several ecogeographical races from it, apparently, continues to be an
aggregate species, strongly varying in the form and incision of the
leaves, pubescence, length of the corona, etc.

52. P. neglectum Tzvel. sp. nova. in Addenda XXV, 874.

Perennial. Plants 12-30 cm high, with more or less branched rhi-
zome, whitish or grayish from very dense somewhat loose tomentum
of long, flexuous, simple hairs. Stems usually rather numerous, less
often solitary, erect or ascending at base, sparsely leafy, simple or
with 1-2 branches, in axils of cauline leaves. Leaves more densely
tomentose than in previous species, almost whitish from tomentum,
with inconspicuous punctate-glandular hairs; basal leaves 8-10 cm long,
2.5 cm wide, on rather long (often almost as long as lamina), basally
thickened petiole; their laminas oblong-oval, thrice pinnately cut; ter-
minal lobules lanceolate-linear to lanceolate-ovate, up to 0.6 mm wide,
with short cartilaginous cusp; few cauline leaves smaller and less di-
vided, lower short-petiolate, upper sessile. Capitula solitary or 2—3 on
long tomentose peduncle. Involucre 7-12 mm in dia, 4-5 mm long,
densely covered with somewhat loose tomentum, often covering mem-
branous border of involucral bracts; involucral bracts herbaceous, outer
lanceolate, inner 1.5—2 times as long, oblong-linear; all bracts with
rather wide (but narrower than in previous species), brownish or dark
brown, membranous border. Ligulate florets white or pinkish-white,
corolla tube about 1.5 mm long and limb 6-10 mm long; corolla

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247

of tubular florets 2.2-2.6 mm long. Achenes 1.8-2.3 mm long and
0.6—0.8 mm wide, with 6—8 longitudinal ribs; corona 0.6—-0.8 mm long,
lobed up to base in oblong scales, usually corresponding to number of
ribs. Flowering June to August.

On rocks, stony slopes at 1,500—3,000 m.—Soviet Central Asia:
Tien Shan (Chatkal and Fergana ranges), Pamiro-Alai Region (Alai
and Trans-Alai ranges, northwestern Pamir). Endemic. Described from
the southern slopes of Chatkal Range. Type (northern part of Fergana
Valley, Kuldambes mountains near Lake Sary-Chilek, 17.VII.1908, V.
Rovinsky) and isotype in Leningrad.

Note. Differs from the previous species by a more extensive (almost
whitish) pubescence of the whole plant, narrower, usually brownish
(and not blackish-brown), membranous border of involucral bracts,
often almost covered by dense pubescence, longer petioles and other
less distinct characters, apparently, representing a lower montane
ecogeographical race adapted to the mountain ranges surrounding the
Fergana Valley.

53. P. transiliense (Herd.) Rgl. and Schmalh. in Tr. Peterb. Bot.
Sada, V, 2 (1878) 618 p. p.—P. transiliense var. subvillosum Rgl. and
Schmalh. 1. c. ibid. 618.—Tanacetum transiliense Herd. in Bull. Soc.
Nat. Mosc. XL, II (1867) 129.—Chrysanthemum transiliense (Herd.)
Minkw. and Knorr. Rast. Chimkentsk. Uezda (1910) 112.—Richteria
pyrethroides var. subvillosa (Rgl. and Schmalh.) O. and B. Fedtsch.
Perech. Rast. Turk. IV (1911) 177.

Perennial. Plants 10-35 cm high, with thick, more or less branched
rhizome, more or less covered with suberect simple hairs. Stems usually
rather numerous, less often solitary, erect or ascending at base, weakly
leafy, simple, very rarely with single branch. Leaves green or sordid
green from sparse pubescence, usually subglabrous, with rather
numerous, but not always conspicuous punctate-glandular hairs; basal
leaves up to 10 cm long and 2 cm wide, on rather long (sometimes
almost as long as lamina), petioles strongly thickened at base; their
laminas oblong to oblong-linear, twice or thrice pinnately cut; terminal
segments lanceolate to linear-filiform, usually up to 0.6—0.8 mm wide,
with rather long cartilaginous cusp; few cauline leaves strongly reduced
and less divided, sessile. Capitula solitary, very rarely 2, on long
peduncle, usually hairy (to tomentose) at least in upper part. Involucre
7-15 mm in dia, 4—6 mm long, usually tomentose or hairy only near
base; involucral bracts herbaceous, outer lanceolate, inner 1.5—2 times
as long, oblong-linear; all bracts with wide, blackish-brown,
membranous border. Ligulate florets white or pinkish-white, corolla
tube 1.3-2.0 mm long and limb 7-15 mm long; corolla of tubular

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248

florets 2.5-3 mm long. Achenes 2.0-2.5 mm long and 0.6-0.8 mm
wide, with 6—8 longitudinal ribs; corona 0.6—0.8 mm long; cut almost
up to base into scales or lobes of various shape and size. Flowering
June to August.

Rocks, stony slopes, gravel beds above 2,000 m.—Soviet Central
Asia: Tien Shan (Kirgizian, Fergana, Trans-Ilian Alatau, Ketmen,
Kungei-Alatau and Terskei-Alatau ranges), Pamiro-Alai Region (rarely:
Alai, Trans-Alai and Karategin ranges). General distribution:
Dzhungaria-Kashgaria(?). Described from Trans-Ili Alatau. Type and
isotype in Leningrad.

Note. The coincidence of the range of this species with that of P.
pyrethroides, as in many other cases, is only apparent and related to
the great diversity in the conditions of growth in montane regions and
the different geology of the geographically very close montane areas.
In reality, however, both species are never found together, which does
not preclude the existence of populations with transitional features in
places of contact.

54. P. arassanicum (Winkl.) O. and B. Fedtsch. Perech. Rast.
Turk. IV (1911) 187.—P. gracillimum (Winkl. ex O. and B. Fedtsch.)
O. and B. Fedtsch. ibid. 186, nom. nud.—Chrysanthemum arassanicum
Winkl. in Tr. Peterb. Bot. Sada, XI (1890) 372.—C. gracillinum Winkl.
ex O. and B. Fedtsch. |. c. in syn.: B. Fedtsch. Rast. Turk. (1915) 737,
nom. nud.

Perennial. Plants 10—25(30) cm high, with more or less branched
rhizome, entirely glabrous. Stems usually rather numerous, less often
solitary, erect or ascending, weakly leafy, simple, less often with 1-2
lateral branches. Leaves glabrous, with inconspicuous punctate-
glandular hairs; basal leaves 6—8 cm long, 1.5 cm wide, on rather long,
petiole strongly thickened at base; their lamina oblong, twice or thrice
pinnately cut; terminal segments linear-lanceolate to lanceolate-ovate,
somewhat hard, up to 1 mm wide, with short cartilaginous cusp; few
cauline leaves smaller and less divided, usually sessile. Capitula solitary,
less often 2-3, on long glabrous peduncle. Involucre 7-15 mm in dia
and 4—6 mm long, glabrous; involucral bracts like previous species.
Ligulate florets white or pinkish-white, corolla tube 1.3—2.0 mm long
and limb 7-15 mm long; corolla of tubular florets 2.5-3 mm long.
Achenes like previous species but usually with larger, 0.6-1.0 mm
long, corona. Flowering June to August.

Stony and rubbly slopes, rocks at 2,500-3,500 m.—Soviet Central
Asia: Dzhungaria-Tarbagatai(?), Tien Shan, Pamiro-Alai. General
distribution: Dzhungaria Kashgaria(?). Described from the Kirgizian
Range. Type in Leningrad.

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249

Note. This species, first described from nonflowering specimens,
has a range more or less coinciding with that of P. pyrethroides, from
which it differs by the complete absence of pubescence. However, it
iS a rarer species, not always found in the range of P. pyrethroides;
besides, intermediates between these species are totally absent, in spite
of the fact that both species often grow together on the same stony
slope. P. transiliense could be considered as an intermediate species
on the basis of pubescence; however, it has a more limited distribution,
and its leaf blades generally are more divided than in P. arassanicum
and P. pyrethroides. Therefore, we consider it more appropriate to
accept P. arassanicum as a separate species, being a unique “twin-
species” of P. pyrethroides (as in the case of Waldheimia tomentosa
and W. stoliczkae, as well as in many other genera of the Compositae).

P. gracillimum, described by Winkler in the herbarium from
specimens collected by Komarov from the Zeravshan Range, in my
view does not differ from P. arassanicum.

55. P. djilgense (Franch.) Tzvel. comb. nova.—P. transiliense
var. glabra Franch. Pl. Turkest. (1883) 85, t. 17.—Chrysanthemum
djilgense Franch. in Bull. Mus. Hist. Nat. Par. II (1896) 345.—Richteria
pyrethroides var. djilgensis (Franch.) O. Fedtsch. in Tr. Peterb. Bot.
Sada, XXVIII, 1 (1908) 28; O. and B. Fedtsch. Perech. Rast. Turk. IV
(1911) 178.—Ic.: Franch. |. c. (1883) t. 17

Perennial. Plants 3-15 cm high, with more or less branched
rhizome, completely glabrous. Stem erect or ascending, weakly leafy,
always simple. Leaves like previous species, but smaller and less cut
up; basal leaves usually up to 4—5 cm long and | cm wide; their lamina
pinnately cut, with segments pinnately cut to pinnately lobed. Capitula
solitary. Involucre 7-12 mm in dia, 4-5 mm long, glabrous; involucral
bracts like previous species. Ligulate florets white or pinkish-white,
corolla tube 1.3—2 mm long and limb 7-12 mm long; corolla of tubular
florets 2.5—3.0 mm long. Achenes like previous species. Flowering
July to August.

Stony and rubbly slopes, gravel beds, rocks above 3,000 m.—
Soviet Central Asia: Tien Shan, Pamiro-Alai Region. General
distribution: Tibet, Iran-Afghanistan (northeastern part). Described from
Pamir. Type in Paris.

Note. In comparison with the previous species, it is a higher
montane ecogeographical race (or ecological form?), also adapted to
drier regions and related to it through transitional forms.

56. P. leontopodium (Winkl.) Tzvel. comb. nova.—Richteria
leontopodium Winkl. in Tr. Peterb. Bot. Sada, IX (1886) 421; O. and

250

B. Fedtsch. Perech. Rast. Turk. IV (1911) 178.—R. leontopodioides
B. Fedtsch. in Izv. Russk. Geogr. Obshch. IV (1898) 15.—
Chrysanthemum leontopodium (Winkl.) B. Fedtsch. Rast. Turk. (1915)
737.—Exs.: GRF No. 2776.

Perennial. Plants 3-15 cm high, with more or less branched
rhizome, whitish from extensive, somewhat loose tomentum. Stems
erect or ascending, usually rather numerous, caespitose together with
short nonflowering shoots, weakly leafy, always simple. Leaves whitish
from very dense tomentum, under which their outline is scarcely visible,
with inconspicuous punctate-glandular hairs; basal leaves up to 4 cm
long and 1.5 cm wide, on rather long (but always shorter than lamina)
petiole thickened at base; their lamina oblong or obovate, 2—3-pinnately
cut; terminal segments linear-lanceolate to oblong, up to 1 mm wide,
with or without short cusp; cauline leaves strongly reduced and less
divided, sessile. Capitula solitary, on rather long, densely whitish-
tomentose peduncles. Involucre 10-20 mm in dia and 5.5-6.0 mm
wide, covered with copious loose tomentum. Ligulate florets white,
corolla tube 1.5—2.3 mm long and limb 9-15 mm long; corolla of
tubular florets 3-4 mm long. Achenes 1.8-3.0 mm long and 0.7 mm
wide, with 7-10 elongate ribs; corona 0.8—1.3 mm long, cut up to base
into oblong scales, usually corresponding to number of ribs. Flowering
July to August. (Plate IX, Fig. 2.)

Stony and rubbly slopes, rocks, gravel beds above 3,000 m.—
Soviet Central Asia: Tien Shan (Kirgizian, Trans-Ilian Alatau, Kungei-
Alatau, Terskei-Alatau ranges). Endemic. Described from the Kirgizian
Range. Type in Leningrad.

Note. In spite of its unique “edelweiss-like” appearance, it hardly
deserves separation as an independent species, as the dense tomentose
pubescence is the only character distinguishing it from P. pyrethroides.

Economic Importance. A promising species for cultivation as an
ornamental plant for alpine gardens.

GENUS 1529. Spathipappus Tzvel. Gen. Nov.!”

Tzvel. in Addenda, XXV, 875.
Capitula solitary or 2-5, one each at apices of elongate leafy
branches, but always numerous on single plant almost at same height,

‘Treatment by N.N. Tzvelev.

"From Latin words: spatha—cover, and pappus—beard, named for the unique
shape of the pappus of the achenes, which faintly resembles the inflorescence cover
in aroids.

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251

heterogamous, with 12-40 pistillate, always sterile (as a result of par-
tially abortive pistil), ligulate ray florets, in one row, and numerous
bisexual tubular disk florets. Involucre patelliform, 10-18 mm in dia
and 4-6 mm long; involucral bracts herbaceous, imbricate, in 3-4
irregular rows; outer bracts lanceolate, subacute, with narrow, brownish
or blackish-brown membranous border, inner broadly linear, 1.5—2 times
as long, with wider membranous border. Receptacle weakly convex,
glabrous, solid, weakly punctate-tuberculate. Corolla of ligulate florets
white, tube strongly flattened but nonwinged 1.2—1.6 mm long and
limb 7-10 mm long oval-oblong or oblong-linear; corolla of tubular
disk florets yellow, 2.8-3.6 mm long, with narrow and rather long
tube, in upper half is slightly and gradually broadening in 5 straight
deltoid teeth, a seventh to a sixth as long as tube. Filaments thickened
above; anthers lacking distinct basal appendages, but with lanceolate-
ovoid subobtuse apical appendage; pollen grains roundish, spiny. Styles
bifid with linear, truncate branches. Achenes of tubular florets glabrous,
3.0-3.5 mm long, about 0.5-0.6 mm wide, with 5-6 weak elongate
ribs; pappus as entire, unilateral (dorsally up to base), apically more
or less obtuse or truncate, 1.5—2.4 mm long corona; ovary of sterile
ligulate florets connate with corolla base, at apex lacking corona or
with only reduced corona.

Perennial plants usually covered with bifid and simple hairs, with
thick many-branched woody root, usually lacking short nonflowering
shoots, with numerous, densely leafy, simple or sparingly branched,
erect stems, more or less woody at base, and alternate leaves with
twice pinnately cut blades.

Besides the type species, possibly, this genus also includes the
recently described species S. porphyrostephanus (Rech. f.) Tzvel. comb.
nova [= Chrysanthemum porphyrostephanum Rech. f. in Koie and Rech.
f. Symb. afghan. II (1955) 47] from Afghanistan with purple ligulate
florets.

Type of genus: Spathipappus griffithii (Clarke) Tzvel.

Note. This genus differs quite well from Pyrethrum Zinn. by a
unique structure of the pappus and the sterile ligulate florets whose
corolla remains fused with the ovary lacking or almost lacking pappus,
as well as in its growth characteristics. Not incidentally, both the authors
who described the species S. griffithii under two different names
(Chrysanthemum griffithii Clarke and Matricaria spathipappus Winkl.),
indicated that it probably belongs in its own genus. In my opinion, the
genus Spathipappus shows the closest affinity with Xylanthemum Tzvel.,
differing from it by the presence of ligulate florets, a patelliform
involucre, and a weakly convex receptacle.

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252

1. S. griffithii (Clarke) Tzvel. comb. nova.—Chrysanthemum
griffithii Clarke, Compos. ind. (1876) 148.—C. stoliczkae auct. non
Clarke: Hook. f. Fl. Brit. Ind. III (1881) 344 p. p.—Matricaria
spathipappus Winkl. in Tr. Peterb. Bot. Sada, X (1887) 85; O. and B.
Fedtsch. Perech. Rast. Turk. IV (1911) 185.—Pyrethrum griffithii
(Clarke) Buser in Boiss. Fl. or. Suppl. (1888) 299.

Perennial. Plants 15-45 cm high, with thick woody root, dull-
green or grayish-green from rather dense pubescence of bifid and simple
hairs. Stems numerous, arising from woody, strongly branched caudex,
densely leafy, erect or somewhat arcuately bent, simple or with few
lateral branches near base. Leaves dull-green or grayish-green from
rather dense pubescence, with inconspicuous punctate-glandular hairs;
lower cauline leaves up to 3-5 cm long and 1.5 cm wide, on short,
basally thickened petiole or sessile with twice pinnately cut lamina,
oblong-linear to oblong-ovate, terminal segments narrow-linear to
linear-lanceolate, usually up to 1 mm wide, with short cartilaginous
cusp; upper cauline leaves like lower, but usually less divided and
smaller. Capitula solitary, less often 2-5 on a single stem, but always
numerous on one plant. Involucre 10-18 mm in dia, 4—6 mm long,
usually covered with fine tomentum at base and dorsally; involucral
bracts with rather wide, blackish-brown, less often brownish,
membranous border. Ligulate florets white, limb 7-10 mm long and
limb 3—4 mm wide; corolla of tubular florets about 3 mm long. Achenes
3.0-3.5 mm long and about 0.5—0.6 mm wide; corona 1.5-—2.4 mm
long, cut to base on dorsal side, unilateral. Flowering July to August.
(Plate X, Fig.1.)

Gravel beds, screes and slope prolongations; above 3,000 m.—
Soviet Central Asia: Pamiro-Alai (excluding the Alai Range and eastern
Pamir). General distribution: Afghanistan, India (northern part).
Described from Afghanistan. Type in Calcutta or London.

Note. Due to Hooker’s error (Hooker fil. |. c.), for a long time this
species was identified with Chrysanthemum stoliczkae Clarke (I. c.
147), which is absolutely different from it. The isotype of this latter
species is preserved in the Herbarium of the Botanical Institute, Acad-
emy of Sciences of the USSR, which shows the closest affinity with
the genus Brachanthemum DC., but apparently, deserves recognition
as a separate genus together with a series of closely related Central
Asian species (of the Pyrethrum roylei DC. type).

Plants of S. griffithii from the Darvaz Range differ by having
much sparser pubescence of the whole plant, apparently, forming a
unique, morphologically very poorly isolated ecogeographical race.

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253
GENUS 1530. Waldheimia Kar. and Kir. |?

Kar. and Kir. in Bull. Soc. Nat. Mosc. XV (1842) 125.—Allardia
Decne. in Jacquem. Voyage Inde, Bot. IV (1844) 87.

Capitula solitary (but numerous on a single plant), heterogamous,
with 5-20 pistillate sterile ligulate ray florets in one row, and numerous
bisexual tubular disk florets. Involucre patelliform or narrow-
patelliform, 10-20 mm in dia and 4-6 mm long; involucral bracts
imbricate, in 3—4 irregular rows, herbaceous, with wide, blackish-brown,
membranous border, outer lanceolate or oblong, inner oblong-linear,
1.5—2 times as long. Receptacle weakly convex, solid, lacking scales,
very sparsely punctate-tuberculate. Corolla of ligulate florets white or
pink, tube weakly flattened 3-6 mm long, and limb oval or oblong (up
to linear-oblong) 7-15 mm long; corolla of tubular disk florets yellow,
3.0-5.5 mm long, tube narrow in lower half, strongly and rather abruptly
campanulately broadened in upper half, with 5 lanceolate-deltoid
recurved teeth, 1/7—1/5 as long as tube. Anthers lacking distinct basal
appendage, but with broadly lanceolate obtuse apical appendage; pollen
grains roundish, spiny. Styles bifid with linear, truncate, branches. All
achenes similar (but often undeveloped in outer ligulate florets and
then with strongly reduced pappus), glabrous or pubescent, 2.5—4.0
mm long (excluding pappus), narrow-prismatic, turbinate toward base,
weakly curved, with 5-10 prominent longitudinal ribs (of which 5
usually more prominent), testa not mucilaginous on soaking and rather
tightly enclosing achene; pappus 4—8 mm long, I-seriate, of numerous
(25-50), flattened, setaceous scales, usually connate at base in fascicles
(usually as many as ribs). Perennial plants, usually covered with long
simple hairs (to wholly glabrous), strongly branched at base, with
numerous, procumbent or ascending, flowering and nonflowering
branches, somewhat caespitose; leaves alternate, pinnate or lobate.

Of the 8 species of this genus, distributed in the mountainous
regions of Soviet Central Asia from the Altai in the north to the
Himalayan Mountains in the south, 5 are found in the USSR.

Type of genus: Waldheimia tridactylites Kar. and Kir.

Note. In spite of the unique structure of the pappus, the genus
Waldheimia is very closely related to section Richteria (Kar. and Kir.)
Tzvel. of the genus Pyrethrum Zinn and, to a lesser extent, to the
genus Trichanthemis Rgl. and Schmalh. Apparently, the pappus lobes
typical of section Richteria are divided again into numerous bristle-

'Treatment by N.N. Tzvelev.
*Named after G.I. Fischer-Waldheim (1771-1855), President of the Moscow
Society of Naturalists.

254

Plate XI.

265
1 — Waldheimia transalaica Tzvel., habit of the plant, leaf, corolla of ligulate

floret, corolla of tubular floret, and achene; 2 — Trichanthemis radiata Krasch.
and Vved., habit of the plant, corolla of ligulate floret, corolla of tubular floret,
and achene.

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255

like scales resembling the pappus hairs of many other genera of com-
posites.

The genus is divided into two very natural sections, among which
Waldheimia undoubtedly is a derivative of Allardia adapted to the
more severe conditions of very high mountains.

Economic Importance. Some species could possibly be used as
ornamental plants in alpine gardens.

1. Leaves with pinnately parted or cut lamina; ligulate florets fertile,
with normal pappus; achenes hairy; rarely glabrous. (Sect. Allardia.)

+ Leaves spatulate, apically 3—S-lobed; ligulate florets sterile, évith
strongly reduced pappus; achenes glabrous. (Sect. Waldheimia) ..

Leaves, green, glabrous ................. 2. W. stoliczkae (Clarke) ktenh
Leaves grayish from sparse but more copious tomentum ............. a)
Achenes densely hairy up to base; leaves not lobed or with 1-3
strongly reduced lateral lobes in lower half .............ccceeeeeeeee
Na Thad © vs Fal Se eer me Ac emer 3. W. transalaica Tzvel.
+ Achenes more or less hairy to subglabrous only in upper half;

leaves in lower half with reduced lateral lobes 0.0.0.0... ceceeeeeeeeseeees

RAMAN Sic IRR RR RA 1. W. tomentosa (Decne.) Rgl.
4. Plants wholly glabrous ................ 4. W. tridactylites Kar. and Kir.
+ Capitula near base (and often also peduncles) sparsely tomentose ...
ME AL CLA. OEE RA 5. W. glabra (Decne.) Rgl.

@ 2

Section 1. Allardia (Decne.) Tzvel. comb. nova.—Allardia Decne.
l. c. pro parte.—Ligulate florets fertile, with normal pappus; achenes
more or less hairy, less often glabrous. Plarts up to 10-15 cm high;
capitula on longer peduncles; leaves with pinnatipartite or pinnatisect
lamina.

Type of section: Waldheimia tomentosa (Decne.) Rgl.

Note. All species of this section are closely related ecogeographical
races.

1. W. tomentosa (Decne.) Rgl. in Tr. Peterb. Bot. Sada, VI (1879)
308; O. and B. Fedtsch. Perech. Rast. Turk. IV, 214.—Allardia
tomentosa Decne. in Jacquem. Voyage Inde, Bot. IV (1844) 87, t.
95.—Ic.: Decne. |. c.; Blatter, Beaut. Flow. of Kaschmir, I, (1927) t.
30.

Perennial. Plants up to 10-15 cm high, with strongly branched
creeping rhizome, grayish from loose but copious tomentum or long
and flexuous simple hairs. Stems numerous, more or less caespitose,

26

[o<)

256

basally covered with thin-coriaceous, brownish sheaths of strongly
reduced lower leaves, simple, terminating into single pedunculate ca-
pitulum, usually exceeding upper leaves. Leaves 4—5 cm long and 1.2—
1.6 cm wide, loosely tomentose, lacking punctate-glandular hairs, on
short-winged petiole; lamina oblong to oblong-linear, twice (rarely once)
pinnately parted up to broadly winged axis, with 5-10 lobes on each
side, gradually reduced toward lower side, often reaching up to petiole
base; terminal segments lanceolate to ovate, with oblong cusp. Involu-
cre 1.2—2.0 cm in diameter, 4-6 mm long; involucral bracts tomentose
basally and dorsally. Ligulate florets 10—20, fertile; corolla of ligulate
florets white or pinkish-white, limb 10-15 mm long and 3—5 mm wide;
corolla of disk florets 3-5 mm long. Achenes excluding pappus 2.5—
4.0 mm long and 0.6—-0.8 mm wide, more or less hairy in upper half;
pappus 4-7 mm long, often turning more or less brown (up to dark
brown). Flowering July to September.

Screes of small stones, gravel beds; rubble slopes from 3,000 to
4,500 m.—Soviet Central Asia: Tien Shan, Pamiro-Alai (Zeravshan,
Turkestan, Alai, Darvaz, Karateg, and southwestern Pamir ranges).
General distribution: India (northern part), Afghanistan, China (Ti-
bet). Described from the Himalayan Mountains. Type in Paris.

2. W. stoliczkae (Clarke) Ostenf. in Hedin, S. Tibet, VI, 3 (1922)
38.—W. korolkowii Rgl. and Schmalh. in Rgl. in Tr. Peterb. Bot. Sada,
VI (1879) 310; O. and B. Fedtsch. Perech. Rast. Turk. IV, 214.—
Allardia stoliczkae Clarke, Compos. Ind. (1876) 145.

Perennial. Plants up to 10-15 cm high, with strongly branched
glabrous or subglabrous creeping rhizome. Leaves 4—5 cm long and
1.0(1.5) cm wide, green, glabrous, lacking punctate glandular hairs.
Involucre whitish-tomentose only at base, or glabrous. Achenes sparsely
hairy mainly in upper part, sometimes completely glabrous. Other
characters like previous species. Flowering July to September.

Screes of small stones, gravel beds, rubble slopes from 3,000 to
4,500 m.—Soviet Central Asia: Tien Shan (western part, Chatkal, Talass
Alatau, and Kirgiz ranges), Pamiro-Alai Region (western part, includ-
ing western Pamir). General distribution: India (northern part), Af-
ghanistan, Tibet. Described from Himalayan Mountains. Type in
Calcutta or London.

Note. The specimen of this species from the Karakorum Moun-
tains in the herbarium of the Botanical Institute, Academy of Sciences
of the USSR, identified as “Allardia stoliczkae” by C.B. Clarke, is
identical with the type material of Waldheimia korolkowii. P. Wendelbo
(Nytt. Mag. for Bot., Vol. I, 1952, p. 65) also points out the total

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257

similarity of these two species. However, it cannot be considered as
conclusively proven.

The species is extremely close to the previous one, differing from
it only by the almost complete absence of tomentose pubescence. The
incomplete overlap of their ranges and the absence of “intermediate”
forms, nevertheless, provide a basis for retaining W. stoliczkae as a
separate species.

3. W. transalaica Tzvel. sp. nova in Addenda, XXV, 875.

Perennial. Plants up to 10 cm high, with strongly branched creeping
rhizome, grayish from loose but copious tomentum of long and flexuous
simple hairs. Stems numerous, more or less caespitose, less often few,
covered with thin coriaceous brownish sheaths of strongly reduced
lower leaves at base, simple, terminating into single capitulum, usually
exceeding upper leaves. Leaves up to 2.5-3.0 cm long and 1.0(1.5) cm
wide, sparsely tomentose, lacking punctate-glandular hairs, lamina rather
abruptly narrowed in long-winged petiole lacking or with isolated,
strongly reduced, lateral lobes; lamina more or less ovate, 2-(rarely
1-) pinnatipartite up to narrow-winged axis, with 2—5 lobes on each
side; terminal segments lanceolate to broadly ovate, up to 1 mm wide
with rather long cusp. Involucre 1.2-2.0 cm in dia, 4-6 mm long;
involucral bracts tomentose adbasally and dorsally. Ligulate florets 6-
20, fertile; their corolla white, limb 7-12 mm long and 2-4 mm wide;
corolla of disk florets 3.5-5 mm long. Achenes excluding pappus 3—
4 mm long, 0.6—0.8 mm wide, densely pilose up to base; pappus 5—7
mm long, usually light gray. Flowering August to September (Plate XI,
Fig. 1.)

Gravel beds and screes of small stones at 3,000-4,500 m.—Soviet
Central Asia: Pamiro-Alai (Trans-Alai Range). Endemic. Described
from the Trans-Alai Range. Type (Kirgizian SSR, high foothills of
Trans-Alai Range, gravel bed in upper reaches of Kapstur River,
11.VII.1933, No. 867, I. Tyshchenko) in Leningrad.

Note. Besides a separate range, it differs from W. tomentosa in the
shape of the leaves as well as by having achenes densely pilose to the
very base.

Section 2. Waldheimia.—Ligulate florets sterile, with strongly
reduced pappus; achenes glabrous; plants up to 5-6 cm high; capitula
on very short peduncles; leaves spatulate, apically 3—5-lobed.

Type of section: type of genus.

4. W. tridactylites Kar. and Kir. in Bull. Soc. Nat. Mosc. XV
(1842) 126; Ldb. Fl. Ross. II, 627; O. and B. Fedtsch. Perech. Rast.

270

258

Turk. IV, 214, p. p.—Allardia tridactylites (Kar. and Kir.) Sch. Bip.
in Pollichia, XXXXI (1863) 442; Kryl. Fl. Zap. Sib. XI, 2759.—Ic.:
Jubil. semisaec. G. Fischer de Waldheim (1847) t. 1; Pflanzenfam. IV,
5275S phe

Plants up to 6 cm high, with strongly branched woody completely
glabrous, creeping rhizome. Stems (as also nonflowering shoots)
numerous, sparsely or densely caespitose, basally covered with
coriaceous-scaly brownish sheaths of strongly reduced lower leaves,
terminating into a single capitulum, usually not exceeding upper leaves.
Leaves up to 2 cm long and 1 cm wide, spatulate, cuneately narrowed
toward base in short winged petiole, 3(5)-lobed or parted at apex;
lobes usually oblong, obtuse. Involucre 1.2—2.0 cm in dia, 4-5 mm
long, glabrous. Ligulate florets 8-15, sterile, with strongly reduced
pappus, their corolla pink or pinkish-white, limb 7-11 mm long and
2.5-4.0 mm wide; corolla of disk florets 4.0-5.5 mm long. Achenes
excluding pappus 3—4 mm long and 0.6—0.8 mm wide, glabrous; pappus
6.5—8.0 mm long, usually with reddish or brownish tinge. Flowering
July to September.

Screes of small stones, rubble slopes, rock crevices above 3,000
m.—Western Siberia: Altai; Soviet Central Asia: Dzhungaria-
Tarbagatai, Tien Shan (central and eastern parts), Pamiro-Alai (eastern
Pamir, rarely). General distribution: Mongolia (western part),
Dzhungaria-Kashgaria. Described from Dzhungarian Alatau. Type and
isotype in Leningrad.

5. W. glabra (Decne.) Rgl. in Tr. Peterb. Bot. Sada VI (1879)
310.—W. tridactylites auct. non Kar. and Kir.; O. and B. Fedtsch.
Perech. Rast. Turk. IV, (1911) 214 p. p.—Allardia glabra Decne. in
Jacquem. Voyage Inde, Bot. IV (1844) 88, t. 96.—Ic.: Decne. 1. c.;
Blatter, Beaut. Flow. of Kaschmir. I (1927) t. 30; Pampanini Fl. Caracor.
t. 6 (sub. W. tridactylites).

Perennial. Plants up to 3-4 cm high, with strongly branched, woody,
creeping rhizome, more or less tomentose, usually subglabrous. Stems
numerous, sparsely or densely caespitose, basally covered with
coriaceous-scaly brownish sheaths of strongly reduced lower leaves,
terminating into a single capitulum on very short peduncles, more or
less sparsely hairy, usually not exceeding upper leaves. Leaves up to
1.5 cm long and 0.8 mm wide, spatulate, cuneately narrowed toward
base into short-winged petiole, 3(5)-lobed or parted at apex, with obtuse
oblong lobes; upper leaves usually finely arachnoid-tomentose, others
glabrous. Involucre 1.0-1.5 cm in dia, 3.0-4.5 mm long, more or less
sparsely tomentose at base. Ligulate florets 8-15, sterile, with strongly
reduced pappus; their corollas pink of different shades to almost white,

271

259

limb 6-10 mm long and 2.5—4 mm wide; corolla of disk florets 3.5—
5 mm long. Achenes excluding pappus 2.5—3.5 mm long, 0.5—0.7 mm
wide, glabrous; pappus 6.0—-7.5 mm long, usually with pinkish-violet
or brownish tinge. Flowering July to September.

Screes of small stones, rubble slopes, rock crevices, above 3,500
m.—Soviet Central Asia: Pamiro-Alai. General distribution: India
(northern part), Afghanistan, China (southwestern regions). Described
from the Himalayas. Type in Paris.

Note. Recently, this species has often been combined with the
previous species. However, the morphological differences between
them, although not significant, along with their isolated ranges, in my
view are sufficient to retain W. glabra as a separate species—
ecogeographical race (which, in particular, was also noted by Regel,
loc. cit.). Still a third Tibetan ecogeographical race of this series can
be found: W. huegelii (Sch. Bip.) Tzvel. comb. nova (= Allardia huegelii
Sch. Bip. op. cit. 442 and Waldheimia stracheyana Rgl. op. cit. 309),
which is characterized by a more dense arachnoid-tomentose pubes-
cence over the whole plant.

GENUS 1531. Ugamia Pavl. ':”

Pavl. in Vestn. Akad. Nauk KazSSR, VIII (1950) 25.

Capitula solitary (but numerous on a single plant), on very short
(up to 6 mm long), arcuate peduncle, homogamous, with numerous,
bisexual, tubular disk florets. Involucre cupuliform, 7-12 mm in dia
and 7—10 mm long; involucral bracts coriaceous-herbaceous, imbricate,
in 4—5 irregular rows, outer deltoid lanceolate, inner 2-3 times as
long, oblong-linear; all bracts with more or less wide, brownish or
dark brown membranous border. Receptacle weakly convex, almost
flat, solid, lacking scales, sparsely punctate-tuberculate. Corolla of
tubular disk florets pale yellow, more or less brown in upper part, 4.0—
5.5 mm long, much longer than wide, with tube slightly and gradually
broadening in upper half, with 5 lanceolate-deltoid, weakly deflexed
teeth, a sixth to a fifth as long as tube. Filaments thickened in upper
part; anthers lacking distinct basal appendage but with ovoid-lanceolate
subacute apical appendage; pollen grains spherical, spiny. Styles bifid,
their branches linear, truncate. All achenes similar, 4.0—-5.5 mm long,
0.8-1.0 mm wide, with 10-15 longitudinal veins of which 5 usually

'Treatment by N.N. Tzvelev.
*Named after the Ugam Range (western Tien Shan), where the only species of the
genus is found.

Paty

260

more prominent, densely covered with semi erect hairs throughout,
prismatic-cylindrical, narrowed toward base; pappus 5—7 mm long, as
pappose corona, divided up to base in 10-20 narrow-linear scales,
finely and irregularly toothed on margin. Dwarf semi-shrubs rather
densely covered with simple hairs, strongly woody and branched at
base, with numerous, prostrate or ascending flowering and nonflowering
shoots, densely caespilose; leaves small (up to 1.2 cm long), alternate,
with pinnately cut lamina.

A monotypic genus.

Note. The only species of the genus shows a distinct affinity with
species of Trichanthemis Rgi. and Schmalh. and could even be
combined with it as a separate subgenus or section. However, the
highly distinctive appearance (particularly the strongly reduced and
arcuate peduncles of the capitula, which are not to be found in other
more or less closely related genera), cupuliform involucres, almost flat
receptacle, much longer than wide corolla of the tubular florets, and
structure of the pappus of the achene, quite similar to the tuft-like
pappus typical of Waldheimia Kar. and Kir. all provide a basis, fol-
lowing N.V. Pavlov, to consider this remarkable narrow endemic spe-
cies as a separate monotypic genus.

1. U. angrenica (Krasch.) Tzvel. comb. nova; Pav]. ex Poljak. in
Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 371, in syn.
(sphalm.).—U. trichanthemoides Pavl. in Vestn. Akad. Nauk Kazakhsk.
SSR, VIII (1950) 26.—Cancrinia angrenica Krasch. in Fedde, Repert.
XXVI (1929) 29; Polyak. op. cit. 371.—Ic.: Pavl. op. cit. 26, fig. 18.

Perennial. Dwarf semishrubs, 2—8 cm high, with thick, woody,
many-headed root, usually covered with short and long, simple hairs.
Stems (flowering shoots) numerous, with nonflowering shoots, more
or less densely cespitose, strongly woody and branched at base, prostrate
or ascending. Leaves dull-green or almost grayish-green from rather
dense pubescence, lacking punctate glandular hairs, numerous; all leaves
more or less similar, up to 1.2 cm long and 0.6 mm wide, on short
petiole strongly thickened at base; lamina oblong or oblong-linear,
pinnately cut up to almost wingless axis; leaf segments 4-10 on each
side, all or some 2-5 parted; terminal lobes linear- or lanceolate-
filiform, somewhat thick, up to 0.5-0.6 mm wide, with cartilaginous
cusp; upper cauline leaves sessile. Capitula solitary (but numerous on
a single plant), on very short (up to 6 mm long), arcuate peduncle at
apices of flowering shoots. Involucre 7-12 mm in dia, 7-10 mm long;
sparsely hairy; involucral bracts with brownish or dark brown
membranous border, broader and appendiculately broadening at apex
of inner bracts. Corolla of tubular florets 4.0-5.5 mm long. Achenes

273

274

261

4.0-5.5 mm long, with 10—15 longitudinal ribs, of which 5 usually more
prominent, densely pubescent; corona pappose, S—7 mm long, cut up to
base into 10-20 narrow-linear scales. Flowering July to August.
Rubbly and stony slopes, gravel beds above 2,000 m.—Sovier
Central Asia: Tien Shan (Chatkal, Pskem, Ugam ranges). Endemic.
Described from the Chatkal Range. Type and isotypes in Leningrad.

GENUS 1532. Trichanthemis Rgl. and Schmalh. !:?

Rgl. and Schmalh. in Tr. Peterb. Bot. Sada, V, 2 (1878) 617.—
Glossanthis Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XIX (1959) 369.

Capitula always solitary (but numerous on a single plant), on long
erect peduncle, homogamous, with numerous bisexual tubular disk
florets, or heterogamous, with many (4-22), pistillate, ligulate, outer
florets in one row, and numerous bisexual, tubular disk florets. Involucre
patelliform or broadly cupuliform, 7-22 mm in dia and 4-8 mm long;
involucral bracts coriaceous-herbaceous, imbricate, in 3-5 irregular
rows, outer lanceolate or lanceolate-ovate, inner linear-oblong or linear;
all bracts with more or less distinct dark or brownish membranous
border. Receptacle weakly or strongly convex, solid, more or less hairy
or glabrous, punctate-tuberculate. Capitula of ligulate florets yellow,
white, or pinkish-white, with flattened tube 1.8-3.0 mm long lacking
spur and with oblong-oval or linear-oblong 6-22 mm long limb; corolla
of tubular disk florets yellow, 2.4-5 mm long, with rather long tube,
usually slightly and gradually broadened in upper half, with 5 lanceolate-
deltoid or deltoid, weakly deflexed teeth, a fifth to a fourth as long as
tube. Filaments thickened in upper part; anthers lacking distinct basal
appendage, but with lanceolate-ovoid subobtuse terminal appendage;
pollen grains roundish, spiny. Style bifid; their branches linear, truncate.
All achenes similar, completely covered with more or less adpressed
directed upward hairs, 1.5—3.6 mm long and 0.6—1 mm wide, with 5-
8 prominent longitudinal ribs; pappus 0.8—6 mm long, as corona, cut
up to base into (4)5—10(12) linear oblong or ovate scales. Plants more
or less covered with long and short simple hairs, usually strongly woody
at base, less often (section Subbulbosa) herbaceous perennials with
short nonflowering shoots, sparsely leafy erect stems and alternate
leaves with pinnatisect lamina.

'Treatment by N.N. Tzvelev.
"From the Greek words: thrix (trichos)—hair; and anthemos—floret, so named
because of hairy corolla and achenes of many species of this genus.

Pai)

262

Endemic genus of the USSR with 7 species distributed in the
mountains of Soviet Central Asia.

Type of genus: Trichanthems karataviensis Rgl. and Schmalh.

Note. The species of this genus are quite distinct and differ from
each other by a whole series of rather important characters that they
can, with full justification, be divided into five sections, three of which
are monotypic. Nevertheless, on the whole, there are no doubts about
their close affinity and the naturalness of the entire genus. Convenient
but entirely unjustified with regard to a natural classification is the
splitting off of all species with ligulate florets into a separate section
(Ligulata Krasch.) or even as a separate genus (Glossanthis Poljak.).
A detailed study of the entire set of morphological characters convinc-
ingly shows, for example, that the species T. aurea is closest to T.
paradoxus, and T. radiata, despite its distinctive position, is closer to
T. karataviensis than to other species with ligulate florets.

On the whole, the genus is quite close to the genera Pyrethrum
Zinn and Cancrinia Kar. and Kir. As already mentioned (cf. note to
section Trichanthemopsis Tzvel. of Pyrethrum), even Pyrethrum
tianschanicum Krasch., which differs from Trichanthemis butkovii only
by having glabrous achenes and fewer rows of involucral bracts, perhaps
even deserves to be included in the genus Trichanthemis. However,
this species is no less closely related also to other groups of the genus
Pyrethrum. The species T. litwinowii and P. paradoxos earlier were
included in Cancrinia; however, careful comparison of them with the
other species of Trichanthemis and with the species of Cancrinia easily
convinces me of their much closer relationship to such species as
Trichanthemis aurea and T. karataviensis than to Cancrinia
chrysocephala Kar. and Kir., not to mention other species of this genus.

1. Corolla of tubular disk florets profusely hairy ............... eee 2.
+ Corolla of tubular disk florets glabrous ..............cccesesseeseeeeeeeeees 4.
2. Capitula lacking ligulate florets (section Trichanthemis) ..............+.
ae a ae ed. aed 2. T. karataviensis Rg]. and Schmalh.
+ i Capitula with ligulateiflorets;.p ete eee ca
Ligulate florets pinkish-white (section Ligulata) ..........s:csscesseeseseeees
Leta etne sd ape, seattle 1. T. aulieatensis (B. Fedtsch.) Krasch.
+ Ligulate florets yellow (section Subbulbosa) ........ccscsccseeseseeen
Sulack ees, tage eorheaae 3. T. radiata Krasch. and Vved.
4. Corona of achenes divided in (8)10(12) narrow-linear scales;
involucral bracts with narrow membranous border (section
Leianthits) s0icanih. RAR SO ee ee a:
+ Corona of achenes divided in (4)5—6(8) oblong scales; involucral
bracts (especially inner) with strongly broadened membranous border
(section Pyrethroides) ccc ih ate cea ce: teen cee 6.

a

278

263

5. Ligulate florets yellow, 4-10 ...........cccscseeeeeeeeees 4. T. aurea Krasch.
+ Ligulate florets absent ................ 5. T. paradoxos (Winkl.) Tzvel.
6. Ligulate florets white, numerous .................. 6. T. butkovii Koval.
+ Ligulate florets absent ................. 7. T. litwinowii (Krasch.) Tzvel.

Section 1. Ligulata Krasch. in Fedde, Repert. XX VI (1929) 28
p. p.; Lipschitz in Byull. Mosk. Obshch. Isp. Prir., Otd. Biol. XLV
(1936) 57 p. p.—Glossanthis Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XIX (1959) 369 p. p. non Glossanthus Klein ex
Benth., 1835.—Plants 20-40 cm high, strongly woody at base. Involucre
14-23 mm in dia; outer bracts 2/5—2/3 as long as innermost. Receptacle
strongly convex, profusely hairy; ligulate florets numerous, pinkish-
white, with linear-oblong corolla limb 12-22 mm long; corolla of tubular
disk florets profusely hairy. Achenes 2.4—3.2 mm long; corona 4-6
mm long, divided in (8)10(12) narrow-linear scales.

A monotypic section.

1. T. aulieatensis (B. Fedtsch.) Krasch. in Tr. Bot. Inst. Akad.
Nauk SSSR, Ser. 1, I (1933) 184.—Chrysanthemum aulieatense B.
Fedtsch. in Knorr. and Minkv. Tr. Pochv.-Bot. E’ksp. Peresel. Upr. VI
(1912) 187.—Glossanthis aulieatensis (B. Fedtsch.) Poljak. in Bot.
Met. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 369.—Ic.: B.
Fedtsch: she. 't. 31.

Perennial. Plants 20-40 cm high, with thick woody root, more or
less covered with scattered, long, simple hairs, usually subglabrous.
Stems rather numerous, strongly woody at base, usually branched below,
sparsely leafy, erect or ascending at base. Leaves glabrous or
subglabrous lacking punctate-glandular hairs; basal leaves up to 10 cm
long and 2 cm wide, on long (sometimes exceeding lamina) petiole,
strongly thickened at base; their lamina oblong, once or twice pinnately
cut, with 3-5 segments on each side; terminal segments linear-filiform,
somewhat thick, up to 0.8 mm wide, with rather long cusp, few cauline
leaves reduced and less divided, sessile. Capitula solitary (but numerous
on a single plant), on long (up to 20 cm), more or less hairy peduncle.
Involucre 14-23 mm in dia, 6—8 mm long, rather densely hairy near
base; involucral bracts coriaceous-herbaceous, outer broadly lanceolate,
subacute, inner oblong-linear, 1.5—-2.5 times as long; all bracts with
rather wide, dark brown or brownish membranous border,
appendiculately broadened at apex in inner bracts. Receptacle strongly
convex, densely pubescent. Ligulate florets 10-18, pinkish-white, with
corolla tube 1.8—2.0 mm long and limb 12-22 mm long; corolla of
tubular florets densely hairy, 3-5 mm long. Achenes 2.4—3.2 mm long,
with (4)5—6(7) long ribs, densely hairy; corona 4—6 mm long, divided

275

264

Plate XII.

1 — Trichanthemis karataviensis Rgl. and Schmalh., habit of plant; 2 —
Lepidolopsis turkestanica (Rgl. and Schmalh.) Poljak., habit of plant.

279

265

up to base into (8)10(12) narrow-linear, subacute scales. Flowering
May to June. (Plate XXVII, Fig. 1. )

Stony slopes, rocks in middle mountain zone.—Soviet Central Asia:
Tien Shan (Ichkele-Tau Mountains). Endemic. Described from Ichkele-
Tau Mountains (western Tien Shan). Type and isotypes in Leningrad.

Section 2. Trichanthemis.—T. sect. Eutrichanthemis Lipsch. in
Byull. Mosk. Obshch. Isp. Prir., Otd. Biol., XLV (1936) 57.—Plants
strongly woody at base, 5-20 cm high. Involucre 7-12 mm in dia;
outer bracts 1/2—2/3 as long as innermost. Receptacle rather strongly
convex, densely hairy; ligulate florets absent; corolla of tubular florets
densely hairy. Achenes 2.0-3.2 mm long; corona 0.8—2.0 mm long,
divided into (4)5—8(10) broadly ovate to oblong-linear scales, sometimes
more or less truncate.

Type of section: type of genus.

2. T. karataviensis Rgl. and Schmalh. in Tr. Peterb. Bot. Sada, V,
2 (1878) 617; O. and B. Fedtsch. Perech. Rast. Turk. IV, 180.—
Cancrinia lasiantha Winkl. in Tr. Peterb. Bot. Sada, XIII (1894) 235;
O. and B. Fedtsch. op. cit. 178.—C. karataviensis (Rgl. and Schmalh.)
Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959)
372.—Ic.: Pavlov in “Sovetsk. Kauchuk.”, III (1922) 25; Korovin,
Rast. Sredn. Azii (1934) 421.—Exs.: Herb. Asiae Med, No. 194;
Pavlov, Pl. turkest. edit. No. 90.

Perennial. Plants 5—20 cm high, with thick, woody, many-branched
root, dull-green or grayish-green from rather dense pubescence of long,
flexuous, simple hairs. Stems rather numerous, strongly woody and
more or less branched at base, simple above, very sparsely leafy, erect
or ascending at base. Leaves dull-green or even grayish-green from
rather dense pubescence, less often subglabrous with inconspicuous
punctate-glandular hairs; basal leaves numerous, very fragile when dry,
2.5(3) cm long and 0.8(1.0) cm wide, on quite long (but shorter than
lamina) petioles, strongly thickened at base; their laminas broadly linear
or linear-oblong, pinnatisect, with segments appressed or semiappressed
to main leaf axis, 2—3-parted up to or almost to base into linear-
filiform somewhat thick, up to 0.6 mm wide lobes; few cauline leaves
like basal, but sessile or subsessile. Capitula solitary (but numerous on
a single plant), on long, densely hairy peduncle. Involucre 7-12 mm
in dia, 4-5 mm long, more or less hairy; involucral bracts coriaceous-
herbaceous, outer lanceolate, subacute, usually lacking border, inner
linear-oblong, 1.5—2 times as long, with rather wide, light-colored,
membranous border. Receptacle rather strongly convex, densely
pubescent; corolla of tubular florets 2.4-3.0 mm long. Achenes 2-3

280

266

mm long and about 0.7—0.8 mm wide, densely hairy, with (4)5(8) long
ribs; corona 0.8—2.0 mm long, divided in (4)5—8(10) broadly ovate to
oblong-linear scales, sometimes more or less truncate(scales shorter
on dorsal side). Flowering May to July. (Plate XII, Fig. 1.)

Rubbly and stony, sometimes clayey slopes up to middle mountain
zone.—Soviet Central Asia: Dzhungaria-Tarbagatai (southern foothills
of Dzhungarian Alatau), Lake Balkhash Region (Chu-Ili Mountains),
Tien Shan (Karatau Mountains and adjoining regions), Pamiro-Alai
Region (foothills of Zeravshan Range). Endemic. Described from the
Karatau Mountains. Type and isotype in Leningrad.

Note. A polymorphic species, strongly varying in pubescence, size
of capitula, length of corona on achenes, etc., and, apparently, separating
into several ecogeographical races. However, we could not find stable
differences between the type specimens from the Karatau Mountains
and specimens from geographically completely isolated parts of its
range in the foothills of Zeravshan Range, from where Cancrinia
lasiantha Winkl. is described, and the foothills of Dzhungarian Alatau,
from where T. karataviensis ssp. dshungarica Rubtz. is described [in
Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, VIII, 5 (1940) 65]. On
the average, the latter subspecies in fact has larger capitula with a
wider membranous border on the involucral bracts, but similar speci-
mens are also found in the Karatau Mountains.

Economic Importance. According to Pavlov (op. cit.) its leaves
contain traces of rubber.

Section 3. Subbulbosa Tzvel. sect. nova. in Addenda XXV, 877.—
Herbaceous plants 10-35 cm high, with strongly thickened (almost
bulbous) bases of branches. Involucre 12—20 mm in dia; outer bracts
a half to two-thirds as long as inner. Receptacle strongly convex, densely
hairy; ligulate florets numerous, yellow, with oblong-oval, corolla limb
8-12 mm long; corolla of tubular florets densely hairy. Achenes 2.8—
3.6 mm long; corona 3.0—4.2 mm long, strongly truncate (often almost
to base on dorsal side), divided in (4)5—8(10) broadly ovate to oblong-
linear scales.

A monotypic section.

3. T. radiata Krasch. and Vved. in Fedde, Repert. (1929) 29.—
Glossanthis radiata (Krasch. and Vved.) Poljak. in Bot. Mat. Gerb.
Bot. Inst. Akad. Nauk SSSR, XIX (1959) 370.

Perennial. Plants 10-35 cm high, with thick, many-branched tap
root more or less bulbous at base of branches, usually covered with
long, flexuous, simple hairs and floccose tomentum of such hairs. Stems
usually rather numerous, erect or ascending at base, sparsely leafy,

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267

simple. Leaves usually dull-green or even grayish-green from isolated
hairs and floccose tomentum usually on their petioles and rachis, often
completely absent, lacking punctate-glandular hairs; basal leaves up to
8 cm long 3-4 cm wide, on rather long (but shorter than lamina)
petiole, strongly broadening at base and with dense tomentum in their
axils; their laminas oblong to oblong-linear, pinnately cut; leaf segments
3-10 on each side, divided at or near base into rather numerous, narrow-
linear (up to 0.8 mm wide), somewhat thick lobes, usually in a different
plane than leaf axis (closer to a whorl); few cauline leaves like basal,
but smaller, upper sessile. Capitula solitary, on rather long (up to 15
cm ), more or less tomentose peduncles. Involucre 12—20 mm in dia,
6-7.5 mm long, sparsely tomentose at base; involucral bracts almost
herbaceous, outer broadly lanceolate, acuminate, with very narrow
membranous border, inner 1.8—2 times as long, oblong-linear, with
wider, apically broadened, light colored or brownish, membranous
border. Receptacle strongly bulged, densely pubescent. Ligulate florets
12-25, yellow, with corolla tube 2-3 mm long and limb 8-12 mm
long; corolla of tubular florets 3.5-4.5 mm long, densely pubescent.
Achenes 2.8—3.6 mm long, densely pubescent, with 5—8 longitudinal
ribs; corona 3.0—4.2 mm long, strongly truncate, divided in (4)5—8(i0)
broadly ovate to oblong-linear scales. Flowering June to July. (Plate
XI, Fig. 2.)

Rubbly and stony slopes, rocks at 1,500 to 2,500 m.—Soviet
Central Asia: Tien Shan (Karatau Range, Talass Alatau, Kuyuk). En-
demic. Described from the Kuyuk Mountains (western Tien-Shan). Type
or isotype(?) in Leningrad.

Section 4. Leianthus (Novopokr. and Sidor.) Tzvel. comb. nova.
—T. sect. Eutrichanthemis subsect. Leianthus Novopokr. and Sidor. in
Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIII (1950) 349.—
Plants 5—25 cm high, strongly woody at base. Involucre 7-12 mm in
dia; outer bracts a third to a half as long as innermost. Receptacle
rather strongly or weakly convex, densely or sparsely hairy (to
subglabrous). Ligulate florets 4-10, yellow, with oblong-oval corolla
limb 6-12 mm long, or entirely absent. Corolla of tubular florets
glabrous. Achenes 1.5—2.4 mm long; corona 1.3—3.4 mm long, divided
into (8)10(12) narrow-linear scales.

Type of section: Trichanthemis paradoxos (Winkl.) Tzvel.

4. T. aurea Krasch. in Fedde, Repert. XXVI (1929) 28-29.—T.
radiata auct. non Krasch. and Vved.: Lipsch. in Byull. Mosk. Obshch.
Isp. Prir. Otd. Biol. XLV (1936) 59 p. p.—Glossanthis aurea (Krasch.)
Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959)
370.

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268

Perennial. Plants 5-15 cm high, with thick, woody many-branched
root, dull-green or grayish-green from rather dense pubescence of long,
flexuous, simple hairs. Stems rather numerous, strongly woody and
more or less branched at base, simple above, very sparsely leafy, erect
or ascending at base. Leaves dull-green or almost grayish-green from
rather dense pubescence, lacking punctate-glandular hairs; basal leaves
up to 2 cm long and 0.5 cm wide, on rather long (but shorter than
lamina) petiole, strongly thickened at base, tomentose in axil; lamina
of basal leaves broadly linear to linear-oblong, pinnately cut; leaf
segments quite distant, 3-5 on each side, 2—5-pinnately or palmately
parted into narrow-linear and lanceolate, subacute lobes up to 0.5 mm
wide, usually in a different plane than rachis; few cauline leaves strongly
reduced, uppermost sessile. Capitula solitary, on rather long (up to 10
cm), finely tomentose peduncle. Involucre 7-12 mm in dia and 5-7
mm long, densely pubescent near base; involucral bracts coriaceous-
herbaceous, outer ovate, obtuse, inner 2-3 times as long, linear-oblong;
all bracts with rather wide, light colored, membranous border on margin.
Receptacle strongly convex, densely pubescent. Ligulate florets 4-10,
yellow, with corolla tube about 1.8 mm long and limb 6-12 mm long;
corolla of tubular florets 2.5—3.2 mm long, glabrous. Achenes 2.0-2.4
mm long, about 0.6-0.7 mm wide, more sparsely hairy, with 5-8
longitudinal ribs; corona 2.5—3.4 mm long, divided in (8)10(12) narrow-
linear scales. Flowering May to June.

Stony (predominantly shaly) slopes of lower mountain zone.—
Central Asia: Syr-Darya (northern foothills of Alai Range). Endemic.
Described from northern foothills of Alai Range. Type in Leningrad.

5. T. paradoxos (Winkl.) Tzvel. comb. nova.—T. glabriflora
Novopokr. and Sidor. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XIII (1950) 347.—Cancrinia paradoxos Winkl\. in Tr. Peterb. Bot.
Sada, XI, 12 (1891) 371; O. and B. Fedtsch. Perech. Rast. Turk. IV,
178.—C. paradoxa Winkl. ex Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XIX (1959) 371.—Ic.: Novopokr. and Sidor. op.
cit. 348.

Perennial. Plants 6-25 cm high, with thick, woody, many-branched
root, usually covered with long simple hairs. Stems rather numerous,
strongly woody and more or less branched at base, simple above, very
sparsely leafy, erect or ascending at base. Leaves dull green from more
or less dense pubescence, lacking punctate-glandular hairs; basal leaves
numerous, 2.0-2.5 cm long, and 0.6 cm wide, on rather long (but
shorter than lamina) petiole, strongly thickened at base (but not
tomentose in axil) and persisting long after withering of leaves; their
laminas linear-oblong or oblong, pinnately cut; leaf segments pinnately

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269

or palmately 2-5 parted into narrow-linear and oblong somewhat thick
up to 0.8 mm wide lobes, usually in a different plane than rachis; few
cauline leaves strongly reduced, short-petiolate. Capitula solitary, on
rather long peduncle. Involucre 8-12 mm in dia, 5—6.5 mm long,
glabrous or subglabrous; involucral bracts coriaceous-herbaceous, outer
lanceolate-ovate, subacute, with very narrow border, inner 2-3 times
as long, oblong-linear, with rather wide light colored or brownish,
membranous border, appendiculately broadened at apex. Receptacle
weakly convex, sparsely pubescent to subglabrous. Ligulate florets
absent; corolla of tubular florets 2.4-3 mm long, glabrous. Achenes
1.5-2.2 mm long, about 0.6—0.7 mm wide, rather densely pubescent,
with 5 (rarely to 8) longitudinal ribs, corona 1.3-—2.2 mm long, divided
into (8)10(12) narrow-linear scales. Flowering July to August.

Rubbly and stony slopes, steppes up to 2,000 m—Soviet Central
Asia: Tien Shan (central part and Kuramin Range). Endemic. Described
from Central Tien Shan. Type in Leningrad.

Note. Despite the geographic isolation, the plants of T. glabriflora
Novopokr. and Sidor. from the Kuramin Range are identical with the
typical specimens of 7. paradoxos from the centrai Tien Shan.

Section 5. Pyrethroides Tzvel. op. cit. 877.—Plants strongly woody
at base, 15-40 cm high. Involucre 9-12 mm in dia; outer bracts 1/3-
1/2 as long as innermost. Receptacle weakly convex, glabrous or
subglabrous; ligulate florets numerous, white, with corolla limb oblong-
ovate, 8-12 mm long or entirely absent; corolla of tubular florets
glabrous. Achenes 2-3 mm long; corona 1.5—2.0 mm long, divided
into (4)5—6(8) oblong scales.

Type of section: Trichanthemis litwinowii (Krasch.) Tzvel.

6. T. butkovii Koval. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
Uzb. SSR, XVI (1961) 32.—Ic.: Koval. op. cit. p. 33.

Perennial. Plants 20-40 cm high, with thick, woody many-branched
root, usually covered with long, flexuous, simple hairs, often
subglabrous. Stems rather numerous, strongly woody, and more or less
branched at base, simple above, sparsely leafy, erect or ascending at
base. Leaves glabrous or sparsely hairy; basal leaves up to 6 cm long
and 2 cm wide, on short petioles strongly thickened at base; their
lamina oblong-obovate, twice or thrice pinnately cut, with more distant
segments; terminal segments narrow-linear, somewhat thick, terminating
into short cusp; cauline leaves reduced and less divided, except
lowermost-sessile. Capitula solitary, 3.0—-3.5 cm in dia, on long peduncle
more or less hairy in upper part. Involucral bracts sparsely hairy or
subglabrous; outer bracts lanceolate-ovate, with rather narrow

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270

membranous border, inner 2—3 times as long, linear-oblong, with very
wide, light colored membranous border, appendiculately broadened at
apex. Receptacle weakly convex, glabrous. Ligulate florets white,
corolla limb oblong-oval; corolla of tubular florets glabrous. Achenes
2.0-2.5 mm long, with (4)5(8) longitudinal ribs, rather densely hairy;
corona about 1.7 mm long, divided into oblong scales. Flowering June
to July.

Rocks of middle and upper mountain zones.—Soviet Central Asia:
Tien Shan (Chatkal River basin). Endemic. Described from western
Tien Shan (Ak-Bulak River in Chatkal River basin). Type in Tashkent.

Note. This species diagnosis was kindly provided by S.S.
Kovalevskaya prior to its publication. T. butkovii shows a quite clear
relationship with both Trichanthemis litwinowii and Pyrethrum
tianschanicum, as though occupying with these species an intermediate
position between the genera Pyrethrum and Trichanthemis.

7. T. litwinowii (Krasch.) Tzvel. comb. nova.—Cancrinia
litwinowii Krasch. in Bot. Mat. Gerb. Glavn. Bot. Sada, III (1922) 81;
Poljakov in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959)
372.—Pyrethrum simulans Pavl. in Vestn. Akad. Nauk Kazakhsk. SSR,
3 (1950) 36.—Ic.: Pavl. ibid. 37, fig. 12.

Perennial. Plants 15-30 cm high, with thick, woody, many-branched
root, usually covered with long simple hairs. Stems rather numerous,
strongly woody and more or less branched at base, simple above,
sparsely leafy, erect or ascending at base. Leaves sparsely hairy or
subglabrous, lacking punctate-glandular hairs; basal leaves usually up
to 2.5 cm long, 0.6 cm wide, less often up to 5 cm long and 1.5 cm
wide, on very short petioles strongly thickened at base; their lamina
broadly linear or linear-oblong, palmately cut up to almost wingless
axis; leaf segments 5-10 on each side, strongly reduced toward base
of lamina, 2—5-parted or partly entire; terminal lobes linear- or
lanceolate-filiform, somewhat thick, up to 0.6 mm wide, with short
cartilaginous cusp; cauline leaves smaller and less divided, upper sessile.
Capitula solitary, on long, sparsely pubescent. peduncle. Involucre
9-12 mm in dia, 6-8 cm long, with more or less pubescent near base;
involucral bracts coriaceous-herbaceous, outer lanceolate-ovate,
subobtuse, inner 2—3 times as long, oblong-linear and oblong; all bracts
with very wide, light colored (silvery), membranous border at apex,
appendiculately broadened at apex of inner bracts. Receptacle weakly
convex, glabrous or subglabrous (with isolated fragile hairs). Ligulate
florets absent; corolla of tubular florets 3-4 mm long, glabrous. Achenes
2-3 mm long, about 0.6-0.7 mm wide, with S—8 longitudinal ribs,
quite densely (not very densely) hairy; corona 1.5—2.0 mm long, divided

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271

in (4)5—6(8) oblong, more or less irregularly toothed scales. Flowering
July to August.

Rocks, stony slopes, mainly in middle mountain zone.—Soviet
Central Asia: Tien Shan (Chatkal and Pskem ranges). Endemic.
Described from the Chatkal Range. Type and isotype in Leningrad.

Note. The isotype of Pyrethrum simulans Pavl. in the Herbarium
of the Botanical Institute, Academy of Sciences of the USSR, described
from the Pskem Range, differs from the typical specimens of T.
litwinowii from the Chatkal Range by having larger (up to 5 cm long)
leaves and more densely hairy achenes. Nevertheless, it probably
constitutes a separate ecogeographical race.

GENUS 1533. Xylanthemum Tzvel. Gen. Nov. !?

Tzvel. in Addenda, XXV, 877.

Capitula solitary or numerous, on rather long (up to 15 cm)
peduncle, one each at apices of leafy branches, sometimes at same
height as though forming lax corymb, homogamous, with rather
numerous (15-50), bisexual, tubular disk florets. Involucre cupuliform
or narrow-cupuliform (poculiform), 4-8 mm in dia and 4.5-8.0 mm
long; involucral bracts coriaceous-herbaceous, imbricate, in 4-5
irregular rows, outer ovate or broad-lanceolate, subobtuse, with very
narrow membranous border, inner broadly linear, 2.5—4 times as long,
obtuse, with more or less wide membranous border. Receptacle flat or
almost flat, glabrous, solid, punctate-tuberculate. Corolla of tubular
disk florets yellow, 2.6-3.2 mm long, with very narrow, long, tube
slightly and gradually broadened in upper 2/3, with 5 deltoid teeth,
almost not deflected, a seventh to a fifth as long as tube. Filaments
thickened in upper part; anthers lacking distinct basal appendages, but
with lanceolate-ovoid obtuse apical appendages; pollen grains roundish,
spiny. Styles bifid; their branches linear, truncate. All achenes similar,
glabrous, 1.5-3.5 mm long and about 0.6-0.7 mm wide, prismatic,
gradually narrowed toward base, with 5, less often 6, prominent
longitudinal ribs; pappus as corona, 1.2—2.2 mm long, entire, but incised
dorsally to or almost to base, unilateral or divided up to base in 3-6
oblong scales, more or less shifted ventrally. More or less hairy
semishrubs (hairs bifid, mixed with simple) with thick woody root,

'Treatment by N.N. Tzvelev.
2From the Greek words: xylon—wood, and anthemos—flower; named for the
stem which is woody over a large part in the species of this genus.

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272

strongly branched, stems woody long way at the base, and alternate
leaves with pinnately cut lamina.

The 5 species of this genus are distributed in Afghanistan,
northeastern Iran, and the southern part of Soviet Central Asia. Three
of these are found in the USSR.

Type of genus: Xylanthemum fischerae (Aitch. and Hemsl.) Tzvel.

Note. The species of this genus, being closely related
ecogeographical races of a single series, constitute a completely separate
group, well distinguished by their habit from the other genera of the
tribe Anthemideae. Potjakov (in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XIX, 1959, 371) includes the genus in Cancrinia Kar. and Kir.
emend. Poljak., based on the homogamous capitula borne singly at the
apices of leafy branches, without regard to a whole series of other
equally important characters. In our circumscription the genus Xylanthemum
differs from Cancrinia Kar. and Kir. in particular by the structure of
the pappus, shape of the corolla, cupuliform multiseriate involucre
with very small outer bracts, flat receptacle, and stems that are strongly
lignified in the lower part. The genus Trichanthemis Rgl. and Schmalh.
is closer to it, but Xylanthemum is distinguished even from this genus
quite clearly by the glabrous achenes, structure of the pappus, shape of the
corolla, cupuliform involucre, and flat receptacle, not to mention the
differences in habit. Xylanthemum differs from the other, possibly no
less closely related genus Spathypappus Tzvel. by the form and structure
of the involucre and the absence of ligulate florets.

1. | Achenes with corona of 3-6 oblong scales, somewhat approximate
and largerson \ventral side). of iachene,..2.:.2040:..008 Vices. auch
NSS AEAIEGD, . Sa, SLEYES Pace EMO IER Tee 1. X. pamiricum (Hoffm.) Tzvel.
+ Achenes with entire corona, incised dorsally up to base, unilateral
dipdedteedap hae Ata EA RRG BALA ABET, TOL CTCL, OY Ra 2:
2. Involucre cupuliform, 5-8 mm in dia and 5—7 mm long; peduncles
ofcapitula’ 12=152cmilome i.) Anke AL AL, Be ae eee
Pe ee, tettsssssstsssssssssssssseses 2e Ke fischerae (Aitch. and Hemsl.) Tzvel.
+ Involucre narrow-cupuliform, 4—5 mm in dia and 4.5—5.5 mm long;
peduncles -of:capitula;3—4 cm Lome .s.21t 2.5. AU eet asteen deaveneecbeenae nents
Eth), LO, EN, 3. X. rupestre ( M. Pop. ex Nevski) Tzvel.

1. X. pamiricum (Hoffm.) Tzvel. comb. nova.—Chrysanthemum
pamircum Hoffm. in Pauls, Pl. coll. in As. med. and Pers. (1903)
149.—Tanacetum kuschakewiezii O. Fedtsch. Fl. Pamira (1903) 121.—
T. pamiricum (Hoffm.) Bornm. in Mitteil. Thiiring. Bot. Ver. XXII
(1907) 48; O. and B. Fedtsch. Perech. Rast. Turk. IV, 205.—Cancrinia
pamirica (Hoffm.) Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk

287

273

SSSR, XIX (1959) 371.—Ic.: B. Fedtsch. in Bull. Herb. Boiss. ser. 2,
V (1905) t. IV; Pauls. Stud. Veg. Pamir. (1920) 47.

Perennial. Semishrubs, 5-20 cm high, with thick woody root,
usually lacking short nonflowering shoots, dull-green or grayish from
rather dense appressed, bifid (transitional to simple) hairs. Stem,
strongly branched near base, woody over long distance, uniformly leafy,
erect or ascending. Leaves more or less appressed hairy to subglabrous,
with punctate-glandular hairs; cauline leaves up to 2 cm long and 0.8
cm wide, on rather long petiole, distinctly thickened at base, often
with solitary narrow-linear lobes as auricles; their laminas obovate or
broadly oval, cuneately narrowed toward base, usually with short lateral
lobes extending over petiole, pinnately cut up to narrow-winged axis,
with 2-6 approximate lateral segments, in turn almost palmately 2-5
parted; terminal lobes broadly lanceolate to linear, somewhat thick,
subacute, more or less revolute, to 1.0-1.5 mm wide; uppermost leaves
reduced, sessile, less divided (to entire). Capitula usually numerous on
a single plant, one each at apices of leafy branches, on long (up to 6
cm) erect peduncles, often at equal height as though in corymbs.
Involucre 5-8 mm in dia and 5~7 mm long, more or less appressed
hairy; involucral bracts with narrower brownish, membranous border
or almost lacking it; outer bracts a third to two-fifths as long as inner.
Corolla of tubular florets yellow, usually somewhat hairy at apex, 2.6—
3.2 mm long. Achenes 2.4—3.6 mm long (excluding corona) and about
0.7 mm wide; corona 1.2—1.8 mm long, divided to base into 3-6 oblong
scales, more or less ventrally shifted. Flowering July to September.

Stony and rubbly slopes, gravel-beds; above 3,000 m.—Soviet
Central Asia: Pamiro-Alai Region (Pamir). General distribution: Iran-
Afghanistan (Afghanistan, northern part). Described from Pamir. Type
in Copenhagen; isotype in Leningrad.

Note. The report of this species from Iran relates to Pyrethrum
kermanense Bornm. (in Oesterr. Bot. Zeitschr. XLVII, 1897, p. 291,
nom. nudum), subsequently accepted by the author as a synonym of X.
pamiricum, but apparently constituting a separate ecogeographical race.

2. X. fischerae (Aitch. and Hemsl.) Tzvel. comb. nova.—
Tanacetum fischerae Aitch. and Hemsl. in Journ. Linn. Soc. XIX (1882)
170.—Pyrethri ‘n fischerae (Aitch. and Hemsl.) Buser in Boiss. FI. or.
Suppl. (1888) 300.—Chrysanthemum fischerae (Aitch. and Hemsl.)
Rech. f. in Kdie and Rech. f. Symb. afghan. II (1955) 45.

Perennial. Semishrubs, 10-25 cm high, with thick woody root,
dull green from appressed bifid hairs. Stem like previous species. Leaves
like previous species, but usually less hairy, usually with entire lobes.
Capitula usually numerous, one each at apices of leafy branches, on

28

foe}

274

long (3-15 cm long) erect peduncles. Involucre 5—8 mm in dia and 5-
7 mm iong, more or less hairy; involucral bracts usually with wider,
dark brown, membranous border, outer a fourth to a third as long as
inner. Corolla of tubular florets yellow, 2.5-3.2 mm long. Achenes
2.4-3.2 mm long, about 0.6—0.7 mm wide; corona 1.5—2.2 mm long,
entire, unilateral, sinuate-toothed at apex. Flowering August to
September.

Stony and rubbly slopes, gravel beds; above 3,000 m.—Soviet
Central Asia: Pamiro-Alai (south western Pamir). General distribution:
Afghanistan. Described from Afghanistan. Type in London; isotype in
Leningrad.

Note. Apparently it replaces the previous species in Afghanistan.
In the USSR, so far it is known only from one locality (Khorog-Osh
highway, near the village [“Kishlak”] of Morch, right bank of the
Tobuz-Bulak River, about 3,200 m above sea, 4.IX.1955, No. 5127, S.
Ikonnikoy).

3. X. rupestre (M. Pop. ex Nevski) Tzvel. comb. nova.—
Tanacetum rupestre M. Pop. ex Nevski in Tr. Bot. Inst. Akad. Nauk
SSSR, Ser. 1, IV (1937) 282.—Chrysanthemum rupestre (M. Pop. ex
Nevski) Rech. f. in Koie and Rech. f. Symb. afghan. II (1955) 47, non
Matsum. and Koidz. 1910.—Cancrinia rupestris (M. Pop. ex Nevski)
Poljak. in Fl. Turkm. VII (1960) 104.—Ic.: Nevski, op. cit. 285, fig.
8.

Perennial. 6-15 cm high semishrub with thick woody root, usually
grayish from rather dense pubescence of appressed bifid hairs. Stem
and leaves like previous species, but leaves usually more densely hairy,
with usually entire segments; uppermost leaves entire, linear. Capitula
usually numerous, one each at apices of leafy branches, on shorter (up
to 3-4 cm long) erect peduncles. Involucre 4-5 mm in dia and 4.5-5.5
mm long, more or less hairy; involucral bracts with rather wide, dark-
brown, membranous border, outer a fourth to a third as long as inner.
Corolla of tubular florets yellow, 2.5—-3.2 mm long. Achenes 1.8-—2.5
mm long and about 0.6 mm wide; corona 1.2-1.6 mm long, entire,
unilateral, apically irregularly truncate or obtuse. Flowering July to
August.

Stony and rubbly slopes, rocks; from 2,000 to 3,000 m.—Soviet
Central Asia: Pamiro-Alai (Kugitang Mountains). General distribution:
Afghanistan(?). Described from Kugitang. Type and isotype in
Leningrad.

Note. Apparently replaced in Afghanistan by the closely related
species X. polycladum (Rech. f.) Tzvel. comb. nova (= Chrysanthemum
polycladum Rech. f. op. cit. 47).

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275
GENUS 1534. Lepidolopha Wink. ':?

Winkl. in Tr. Peterb. Bot. Sada, XIV (1893) 236.

Capitula homogamous, discoidal, florets tubular; involucre
cylindrical, imbricate, 2—3-seriate; receptacle convex, glabrous; corolla
regular, 5-toothed, with limb slightly broader than tube; anthers almost
entire at base, with 2 very short cusps. Style branches, cylindrical
above, weakly thickened, truncate, brushlike. Achenes compressed,
linear, obpyramidal; pappus membranous, with 8-10 lanceolate scales,
almost entire or 2—3-lobed. Shrubs with flowering shoots more densely
leafy below, sparsely above; lower leaves ternate, upper entire. Capitula
medium, in corymbose inflorescence.

Types of genus: L. komarowii Wink.

The genus comprises 6 species endemic in Central Asia.

1. Capitula aggregated in dense corymbs, 5-20 on short peduncles,
solitary stems bearing 3-5 capitula each; leaves deeply divided,

BEES NE) 2. RUN RecA ora cicie st RES a
+ Capitula aggregated in lax inflorescence, on long peduncles, or
solitary; leaVés Scattered or ‘entifé's):.0..).2..5.0...00. WOR. ZO... 3
2. Lobes of leaves 2.5-4.5 mm wide; achenes pyramidal, 4.0-4.5 mm
eS Se ee ee ee 1. L. komarowii Winkl.

+ Lobes of leaves 5-10 mm broad; achenes 3 mm long .................
wet tee OS) ee ea 2 2. L. filifolia Pavl.

3: Leaves not deeply ternate in broad lobes or deeply dissected and
compound ternate with narrow-linear lobes ..............eeeeeeeeeeeees 4.
+ Leaves entire; age in lax inflorescence, 8-9 mm long, 4 mm in
IRR OA: BOR AR, A RR! 6. L. fedtschenkoana Knorr.

4. Capitula usually solitary, less often in clusters of 2-3; leaves deeply
divided into linear lobes, 2.0-2.5 cm long and | mm wide, upper
CIHEBAG: AML SORA ae 5. L. Karatavica Pavl.

+ Capitula in more lax inflorescence on long peduncles ................. =

Leaves more or less deeply ternate, with linear or linear-lanceolate

lobes; lobes of lower leaves 1.0-1.5 cm long, 1.0—-2.5 mm wide;

upper leaves entire; capitula 9 mm long, 3 mm Wide .............::seseeeeeees

TORU. MING... AAR REMAR. BNE ER, .tedtensd 4. L. nuratavica Krasch.

+ Leaves not deeply ternate, with rather wide lobes, 1.0—1.5 cm long,
3—5 mm wide; upper leaves both entire and incised; capitula 7-10
mm long, 5—7 mm Wide ...........ceeeeeeeeee 3. L. mogoltavica Krasch.

4

'Treatment by O.E. Knorring.
*From the Greek words: lepis—scale, and lophos—crest; named for the nature of
the pappus.

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276

Section 1. Congestae Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, IX (1946) 162.—Capitula smaller, cylindrical, on rather
short peduncles, in groups of 5—20 in dense corymbs; stems solitary
bearing 3-5 capitula each.

1. L. komarowii Winkl. in Tr. Peterb. Bot. Sada, XIII (1894)
234.—Exs.: GRF No. 1875.

Perennial. 15-35 cm high, branched semishrub, older branches
covered with grayish bark, year-old stems erect, forked in middle and
above, weakly pubescent, densely leafy in lower half. Leaves petiolate;
lamina deeply divided, with linear or oblong lobes, 15-25 mm long,
2.5—-4.5 mm wide, with prominent midrib; upper leaves entire; all leaves
weakly pubescent. Capitula in corymbs on 5-10 mm long peduncle,
cylindrical—7 mm long, 3 mm in dia. Involucre many-rowed, finely
pubescent; outer involucral bracts broadly ovate, obtuse, 3 mm long,
2 mm wide; middle bracts 5 mm long, and 2 mm wide. Receptacle
glabrous. Florets yellow at apex. Achenes pyramidal, 4.0-4.5 mm long;
pappus coronate, 1.5—2.0 mm long, membranous, deeply divided, scales
mostly entire, less often toothed. Flowering July to August.

Stony slopes.—Soviet Central Asia: Tien Shan (western Karatau
Range), Pamiro-Alai. Described from Hissar Range. Endemic. Type in
Leningrad.

2. L. filifolia Pavl. in Vestn. Akad. Nauk KazSSR (1950) 24.

Perennial. 25—45 cm high semishrub. Stem branched, old branches
covered with grayish-brown bark, young weakly pubescent, densely
leafy in lower half, densely or sparsely in middle; leaves deeply
dissected, petiolate, with prominent midrib; leaf lobes linear, 2-3 cm
long and 5-10 mm wide; upper leaves smaller, entire less often twice
or thrice pinnate, 7-10 mm long. Capitula in dense corymbs, on short
5-9 mm long peduncles, small, cylindrical, 6-7 mm long and 2 mm in
dia. Involucre many-rowed, densely scaly, outer and middle bracts 3
mm long and 2 mm wide, inner 4 mm long and 1 mm wide. Receptacle
glabrous, flat. Florets yellow. Achenes 3 mm long, ribbed; pappus of
membranous scales, divided up to base, or in turn lobed above or
toothed, 1 mm long.

Rubbly and stony slopes.—Soviet Central Asia: Tien Shan
(western). Endemic. Described from western part of Karatau Range.
Type in Alma-Ata; cotype in Leningrad.

Section 2. Divaricatae Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, IX (1946) 182.—Capitula in depauperate, very lax inflo-
rescence on long peduncles or solitary, only sometimes on separate
stems crowded together at the top in twos or threes.

277

3. L. mogoltavica Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, IX (1946) 182.—L. komarowii Winkl. var. mogoltavica
Krasch. in Bot. Mat. Gerb. Bot. Sada, III (1922) 79.—Exs.: H.F.A.M.
No. 195, sub L. komarowii Winkl.

Perennial. 30-40 cm high semishrub, strongly branched, old
branches covered with gray bark, year-old stems erect or slightly
divergent, divaricate in upper half, puberulent, densely leafy below.
Leaves weakly ternate, with subobtuse, rather wide lobes, 1.0—-1.5 cm
long and 3-5 mm wide, or leaves partly almost entire, spatulate, with
prominent midrib, appressed hairy. Capitula cylindrical, 7-10 mm long,
5-7 mm in dia, on long peduncles, 2.5-3.5 cm long, in depauperate
corymbs of 2-3, or solitary. Involucre many-rowed, compact, grayish-
pubescent; outer involucral bracts 3—S mm long, 2.5 mm wide, middle
5-5.5 mm long and 3 mm wide, inner 6 mm long and 1.5 mm wide.
Receptacle glabrous, flat. Florets yellow, more or less glandular.
Achenes ribbed; pappus coronate, of scales incised or 2-3 toothed.
Flowering June.

Sandstone and limestone mountain slopes.—Soviet Central Asia:
Pamiro-Alai. Endemic. Described from Mogoltau Mountains. Type in
Leningrad.

4. L. nuratavica Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, IX (1946) 179; Ic.: ibid. 181, Fig. 11.—Exs.: Gerb. FI.
SSSR No. 3360.

Perennial. 80-100 cm high semishrub, strongly branched, old
branches covered with grayish bark; solitary stems erect, forked in
middle and above, appressed hairy, densely leafy below. Cauline leaves
and leaves of nonfertile shoots petiolate, deeply ternate above, with
linear or linear-lanceolate lobes, 1.0-1.5 mm long and 1.0-2.5 mm
wide; upper leaves linear, entire, S-15 mm long. Capitula in very lax
inflorescence, on 2.0—3.5 cm long peduncle, narrow-cylindrical, 9 mm
long and 3 mm wide. Involucre many-rowed, grayish-pubescent; outer
involucral bracts ovate-orbicular or subacute, 2-3 mm long and 2.0—
2.5 mm wide, middle ovate-lanceolate, 3-5 mm long and 2-3 mm
wide, subobtuse, inner lanceolate or linear-lanceolate, 5-8 mm long
and 1 mm wide. Receptacle glabrous, flat. Florets yellow at apex,
more or less glandular. Achenes obpyramidal, ribbed, glandular; pappus
coronate, 1.5 mm long, of scales, more or less deeply 4—6-parted,
incised, sometimes scales toothed or entire. Flowering July to August.

Stony mountain slopes, on exposed granite and limestone deposits.
—Soviet Central Asia: Pamiro-Alai. Endemic. Described from the
Nuratau Mountains. Type in Leningrad.

292

278

Note. This species is not always very sharply distinguishable from
L. mogoltavica Krasch., and transitional forms with shorter peduncles
as well as broader leaf lobes are found where they come in contact
with each other.

5. L. karatavica Pavl. in Bull. Soc. Nat. Mosc. (1933) 127.—
Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, IX (1946)
182.

Perennial. Spreading 20-30 cm high semishrub, old branches
covered with grayish bark, young branches weakly pubescent, densely
leafy below. Leaves deeply incised or ternate into narrow linear lobes,
20-25 mm long and 1 mm wide, with prominent midrib, densely
puberulent; upper leaves entire, linear, 7-9 mm long. Capitula usually
solitary, less often in groups of 2—3 on long peduncles, 2-4 cm long,
broadly cylindrical, up to 10 mm and more long and up to 1 cm in dia.
Involucre compact; involucral bracts many-rowed, outer 3 mm long
and 3 mm wide, middle 6 mm long and 3.5 mm wide, inner 8 mm long
and 2 mm wide. Receptacle flat, glandular-hairy. Corolla yellow.
Achenes 3—3.5 mm long, ribbed, with raised glandular hairs; pappus of
scales divided of lobed. Flowering June to July.

Stony and rubbly mountain slopes.—Soviet Central Asia: Tien Shan
(western). Described from the Karatau Mountains. Type in Moscow.

6. L. fedtschankoana Knorr. in Bot. Mat. XIX (1959) 382.

Perennial. 25-35 cm high semishrubs, divaricately branched, with
numerous stems; old branches covered with dark gray bark, year-old
branches slightly deflected, yellowish brown, puberulent, leafy from
base to middle. Leaves entire, linear, with grooved midrib, pubescent;
lower leaves 25 mm long and 2.5-3.5 mm wide; leaves below
inflorescence 3.0—-3.5 mm long and 0.5 mm wide. Capitula on more or
less long peduncle, in depauperate cylindrical inflorescence, 8-9 mm
long, 4 mm in dia. Involucre many-rowed; outer involucral bracts 4
mm long and 2 mm wide, middle 5 mm long and 2 mm wide, inner 6
mm long and 1.5-2 mm wide. Receptacle flat. Florets yellow. Achenes
obpyramidal, with sunken ribs, 3.5 mm long; pappus coronate, 1.5—2.0
mm long, of scales, lobed up to base, apically entire or short-toothed.

In mountains at 1,800 m.—Soviet Central Asia: Pamiro-Alai
(western part). Described from Kugitang Mountains. Endemic. Type in
Leningrad.

293

279
GENUS 1535. Cancriniella Tzvel. Gen. Nov. !:?

Tzvel. in Addenda, XXVI, 876.

Capitula solitary (usually numerous on single plant) on rather long,
erect peduncles, at apices of strongly reduced branches, homogamous,
with numerous bisexual tubular disk florets. Involucre broadly
cupuliform, 6-10 mm in dia and 3.5-6 mm long; involucral bracts
herbaceous, fewer, imbricate, in 1-2 rows, almost equal; outer
lanceolate, acute, more or less lacking membranous border, inner
oblong-lanceolate, subobtuse, with rather wide, light colored,
membranous border in upper part; usually few innermost bracts short,
linear, resembling floral bracts. Receptacle strongly convex, almost
hemispherical, solid, sparsely pubescent, weakly punctate-tuberculate.
Capitula of tubular disk florets yellow, 2.2-3.0 mm long, tube slightly
and gradually broadening in upper two-thirds, with 5 almost nondeflexed
deltoid teeth, two-sevenths to two-ninths as long as tube. Filaments
thickened in upper part; anthers lacking distinct basal appendage, but
with lanceolate-ovoid obtuse apical appendage; pollen grains roundish,
spiny. Styles bifid; their branches linear, truncate. Achenes similar,
glabrous, more or less terete, narrowed toward base, 2.4-2.8 mm long,
0.6—-0.7 mm wide, with 10 prominent, almost uniformly distributed
longitudinal ribs, lighter than achene surface; pappus 0.4—0.6 mm long,
as corona, divided to (or almost to ) base in 6-10 irregular subobtuse
scales along margin. Perennial herb, densely covered with long erect
simple hairs (but often woody at base) with thick many-branched root
and numerous, strongly reduced and usually closely clustered branches,
terminating in rosula of leaves with pinnatipartite or palmatipartite
lamina, and capitula on long peduncles.

A monotypic genus.

Note. The only species of this genus, an endemic of the Chu-Ili
Mountains, is well distinguished by its unique morphology from all
other genera of the Chrysanthemum L. sensu lato group. It is well
distinguished also from Cancrinia Kar. and Kir., the closest genus to
it, in the structure of the involucre, with relatively few, almost equal,
broad involucral bracts and achenes with 10 prominent ribs; from the
other closely related genus, Pyrethrum Zinn, in the structure of the
involucre and the absence of pistillate ligulate florets; and from
Trichanthemis Rg]. and Schmalh., in the structure of the involucre and
the glabrous achenes.

'Treatment by N.N. Tzvelev.
?Diminutive of the generic name Cancrinia Kar. and Kir.; named for the similarity
with species of that genus.

294

280

1. C. krascheninnikovii (Rubtz.) Tzvel. comb. nova.—
Brachanthemum krascheninnikovii Rubtz. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XI (1949) 208.—Pyrethrum krascheninnikovii
(Rubiz.) Pavl. in Vestn. Akad. Nauk Kazakhsk. SSR, 5 (1952) 93.—
Cancrinia krascheninnikovii (Rubtz.) Poljak. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XIX (1959) 371.—Ic.: Rubtz. op. cit. 209.

Perennial. Plants 3-10 cm high, with thick many-branched root,
grayish-green or dull green from dense pubescence of long, erect, simple
hairs; branches numerous, usually closely clustered, strongly reduced,
densely covered with sheaths of dead leaves and terminating into leafy
rosula. Leaves up to 1.5 cm long and 0.5 cm wide, usually grayish-
green from dense pubescence, lacking punctate-glandular hairs, on rather
long (sometimes longer than lamina), petiole strongly thickened at base,
with pinnately or palmately 3—7-parted laminas, cuneately narrowed to
petiole; terminal lobes oblong to narrow-linear, up to 1.2 mm wide,
obtuse or subacute. Capitula solitary (but numerous on a single plant),
on long (2-8 cm long), erect, leafless or almost leafless (with 1-2
narrow-linear leaves) peduncles, arising from leafy rosulas. Involucre
6-10 mm in dia, 3.5-6.0 mm long; involucral bracts almost equal.
Corolla of tubular disk florets 2.2-3.0 mm long. Achenes 2.4—2.8 mm
long, about 0.6-0.7 mm wide, with 10 prominent longitudinal ribs and
0.4-0.6 mm long corona. Flowering May to June. (Plate XIII, Fig. 2.)

Rubbly and stony slopes of lower mountain zone.—Soviet Central
Asia: Lake Balkhash Region (Chu-Ili Mountains). Endemic. Described
from Chu-Ili Mountains. Type and isotype in Leningrad.

GENUS 1536. Cancrinia Kar. and Kir. Emend., Tzvel. !:?

Kar. and Kir. in Bull. Soc. Nat. Mosc. XV(1842) 124.—
Lepidolopsis Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XIX (1959) 374 p. p.

Capitula usually quite numerous, up to 40, less often (section
Polychrysum up to 80-100, in lax or more or less dense corymbs or
corymbose-panicles, or solitary (section Cancrinia), homogamous, with
numerous, bisexual, tubular disk florets. Involucre patelliform, less
often (section Polychrysum) cupuliform, 3-17 mm in dia and 3-6 mm
long; involucral bracts herbaceous, imbricate, in 3-4 (less often almost
2) irregular rows; outer bracts lanceolate-linear or lanceolate, sometimes

‘Treatment by N.N. Tzvelev.
*Named for the Finance Minister E.F. Cancrin (1774-1845), who helped in some
scientific investigations; C.F. Ledebour dedicated Flora Rossica to him.

297

281

few, inner 1.5—2 times as long, lanceolate-oblong to broadly linear; all
bracts with more or less distinct light colored or dark membranous
border; innermost bracts often very narrow, resembling floral bracts,
sometimes merging with outer bracts. Receptacle strongly or weakly
convex, solid, glabrous or sparsely pubescent, weakly punctate-
tuberculate, sometimes distinctly alveolate. Corolla of tubular disk
florets yellow, 1.2-3.0 mm long, with short wide tube, rather strongly
and abruptly broadened in upper half, with 5 slightly decurved triangular
or lanceolate-triangular teeth, a fifth to a third as long as tube. Filaments
thickened in upper part; anthers lacking distinct basal appendage but
with lanceolate-ovoid subobtuse apical appendage; pollen grains
spherical, spiny. Styles bifid; their branches linear, truncate. All achenes
similar, glabrous, less often sparsely hairy (section Cancrinia),
prismatic-terete, narrowed toward base, 1.3-2.4 mm long, about 0.6—
0.8 mm wide, with 5, less often 6-7, prominent longitudinal ribs
(sometimes more or less converging on ventral side of outer achenes);
pappus 0.4—1.0 mm long, less often (section Cancrinia) up to 3 mm
long, as corona, often divided up to base in lobes or scales (usually as
many as ribs) or various shapes and size. Perennial herbs, more or less
covered with long simple hairs (in section Tanacetopsis sometimes
mixed with bifid hairs), usually with short vegetative shoots, more
sparsely leafy, antrorse (sometimes strongly reduced) cauline and
alternate leaves with pinnately cut, less often palmately cut lamina.

Apparently, the genus, includes about 30 species, distributed in
the mountainous regions of Central and Middle Asia from the Altai in
the north to Afghanistan in the south. Of these, 17 species are found
in the USSR.

Type of genus: Cancrinia chrysocephala Kar. and Kir.

Note. In the circumscription accepted by us, this genus occupies,
as it were, an intermediate position between the genera Pyrethrum
Zinn and Tanacetum L. Perhaps, it is closer to the former genus, dif-
fering from it mainly in the absence of ligulate florets. As stated ear-
lier (cf. note to the genus Pyrethrum Zinn), Pyrethrum majus (Desf.)
Tzvel. a species with homogamous capitula, which arose in cultiva-
tion, could be formally included in the genus Cancrinia, and generally
there is a basis to consider Cancrinia as having originated from the
ancestral type with white or pink ligulate florets by their complete
reduction. Nevertheless, keeping in mind that on the basis of other
characters (e.g., shape of corolla) Cancrinia is a fully isolated group
of species with a definite area of distribution, we prefer to consider it
as a separate genus. From the other closely related genus, Tanacetum L.,
Cancrinia differs mainly by having homogamous capitula as well as in-
volucres almost always patelliform and not cupuliform. Besides, the slight

282

295 Plate XIII.
Habit of plant, leaves, involucre, corolla of tubular floret, and achenes: 1—
Cancrinia chrysocephala Kar. and Kir.; 2 — Cancriniella krascheninnikovii.
(Rubtz.) Tzvel.

298

283

tendency towards hairiness of the achenes and receptacle in Cancrinia,
which would seem to associate it with the genus Trichanthemis Rg.
and Schmalh., must be noted, as it is never found in the genera
Pyrethrum and Tanacetum.

The genus is divided into four completely isolated sections, which,

perhaps, would be treated more properly as independent genera.
However, in my opinion, there is no doubt about their close affinity
which, in particular, is confirmed by the similarity in structure of the
florets and achenes and, according to the observations of V.A.
Simanovich, also the pollen grains.

1.

~ +

2

Capitula 3-4 mm in dia, always numerous (up to 100), in dense
corymbs. Plants up to 150 cm high. (sect. Polychrysum.) ............4+
VTA IR LR IAL RS 1. C. tadshikorum (Kudr.) Tzvel.
Capitula 6-17 mm in dia, less numerous. Plants up to 60 cm high...2.
Corona of achenes 2.3-3 mm long, divided into 8-12 linear or
lanceolate-linear scales; achenes more or less sparsely hairy. Plants
up to 10 cm high with solitary capitula. (sect. Cancrinia) ........... 3.
Corona 0.4—1.5 mm long, entire or divided; achenes glabrous

Achenes sparsely hairy all over. Central Tien Shan .........0..
Pecnabebl ie Mevslie eveQsetworsaveles sedis 18. C. tianschanica (Krasch.) Tzvel.
Achenes sparsely hairy only at apex. Dzhungarian Alatau .................
RRMA Retina i IA 17. C. chrysocephala Kar. and Kir.
Receptacle strongly convex, conically hemispherical; corolla tube
very short and wide. Biennial, less often perennial (?) plants 5—20
cm high, with slender root, which is easily pulled out, and solitary
eapinilal® (sects Matricariotdes.)) G26. XA AUER RAAB RR..n chek
POY GRRL i sdelivesiconiendseneves 16. C. discoides (Ldb.) Poljak.
Receptacle usually weakly convex; corolla tube narrower. Perennial
(8) 15-60 cm high plants with thicker roots and usually numerous,
less often solitary capitula. (sect. TanacetopSis.) .....:c.ssccesseeeseee =)

Corona of achenes about 0.1 mm long, subobtusely small toothed
along margin; capitula numerous (more than 20), at apices of
divaricate branches of wide lax panicle. Paropamisus Mountains
(Turkmenian SSSR) ............... 14. C. paropamisica (Krasch.) Tzvel.
Corona of achenes 0.3-1.5 mm long, more or less (often up to
base) divided into lobes of various shapes and sizes ................. 6.

Stem densely lanate-tomentose at base in axils of sheaths of dead
RODE DALAL, RR ip
SienPHSt tomentose atase 0. SA SR f.,
Capitula 10-40, in more dense, compound corymbs; leaves glabrous
or subglabrous. Trans-Ili and Dzhungarian Alatau ............c:seseseeeeees
a atin nv ncturoeniin i Muslaaaaer 2. C. goloskokovii (Poljak.) Tzvel.

299

10.

ae

Lo:

16.

Capitula fewer, usually up to 10, less often up to 15 on a single

stem, leaves more or less hairy. Other regions ...............:ccssseeeeses 8.
Corolla densely hairy in upper part) sis. cds teecet 01.48 ee 9.
Corolla glabrous)....:.cc.-siss-s+0+ssonssedbeas anette eeae sits can ea eae 10.

Corona of achenes 0.3-0.6 mm long, irregularly toothed up to
middle... Turkestan and. Zetavshany;.:i:sdi.0%3..ad.. dba nearer:
becahey. pomeld shies Sls te: 11. C. urgutensis (M. Pop.) Tzvel.
Corona of achenes 0.6—1 mm long, usually divided up to base in
acuminate unequal lobes. Western spurs of Hissar Range. .................
sasnecedb te bt ats Hie CaM, Se tak 12. C. botschantzevii (Koval.) Tzvel.
Achenes 1.2—1.5 mm long; leaves dull green from sparser
ae Gissar, Darvaz, Vakhsh and other ranges ................
ba dusted ERASE 0 “coh 10. C. pamiroalaicum (Koval.) Tzvel.
ettahan 1.8—2.4 mm long; leaves usually grayish-green from sparse
tomentum. Kugitang and Daisuntau mountains .................s:cccceeeesees
doe Sedewsl. eutets. Pesbetuiie. ..tepeesd (pera, Bee Geena! 13. C. nevskii Tzvel.
Capitula solitary or 2-5, on long deflexed peduncles, usually not in
regular corymb. Plants of plains and lower hills, 8-30 cm high,
with more). slendery faproOtyvvs...scice!. .epy ceded eende dd ~eaepevisets denn ceueereee
Sepsipna Raa ada 15. C. sentoana (Krasch., M. Pop. and Vved.) Poljak.
Capitula usually more numerous. Taller (20-70 cm tall), mountain

plants with thick many-branched root ...........csscsssceseeseseeeeeeeeeenes f2.
Capitula on shorter (usually to 3, less often to 6 cm long) peduncle,
inrégulariand rather denSe COrymDs . ...........:--»-csossnasnudedinayyoroeees 13.

Capitula on longer (to 10-15 cm) peduncles, in very lax corymbose-
panicles, less often (in C. mucronata var. corymbulosa) more dense
iota. hide Sandleve lipo. ai. dpi Noes atheele, Asien. .ghe alpen en ep
Leaves glabrous or subglabrous. Karatau and Nuratau mountains ...
Cicths Oy eo Bceneatlie. oR, NE onan hsaseterpsaeecestac deans 9. C. karatavica Tzvel.
Leaves: denselyshaityicii:.; sis.spseics get hehe shor sss. ae) a epewelatedee aeee tee 14.
Lobes of corona on achenes obtuse. Shugnan Range ..............:e000+
dae 0h dale anbas Le GERROLeAa neeE 3. C. subsimilis (Rech. f.) Tzvel.
Lobes of corona on achenes acuminate. Karatau Mountains .......
bis Re ATE. Sees. mush oe, a teen eae 8. C. pjataevae (Koval.) Tzvel.
Only innermost involucral bracts with very narrow, membranous
border along margin and apex, others lacking border. Western Tien

Shan cal. yey: aocsy.., eel. Crees. 7. C. submerginata (Koval.) Tzvel.
Middle and inner involucral bracts with rather wide membranous
border along’ margin) i¢24. ds -cisccobhvbntecst an tenyenad pe biaeneals: weenie >) ese 16.

Outer involucral bracts numerous, very narrow, more or less linear-
subulate, usually lacking membranous border. Fergana and Chatkal
FAN LES)... Maisie Ah Ae 6. C. ferganensis (Koval.) Tzvel.

300

285

+ Outer in involucral bracts few, wider, lanceolate or lanceolate-linear,
usually with narrow membranous border. Pamiro-Alai................ AT:

17. Leaves sparsely hairy (to subglabrous ) with terminal segments up
to 1 mm wide; involucre appressed hairy; outer involucral bracts

not less than two-thirds as long AS INNET ............:cssssssesessseseeseesseesesensens
Bbbtegeh acti diewks Seycenlouats, 5. C. setaces (Rgl. and Schmalh.) Tzvel.

+ Leaves densely hairy, with terminal segments up to 0.6 mm wide;
involucre sparsely hairy, often subglabrous; outer involucral bracts

a half to two-thirds as lomg aS iMMer ..........cccscecsssceeseeeeseeeeeeeeees
pebthdaetcanaas aenead tapas sf 4. C. mucronata (Rgl. and Schmalh.) Tzvel.

Section 1. Polychrysum Tzvel. sect. nova in Addenda, XXV,
876.—Perennial, monocarpic, 30-150 cm high plants. Capitula rather
numerous (more than 40), at stem apices in dense corymbs, up to 25
cm in dia; involucre cupuliform, 3-4 mm in dia; receptacle as flat
arola near margin, weakly convex in middle, glabrous, punctate-
tuberculate, but distinctly alveolate; corolla tube usually with short
glandular hairs in lower, narrower half. Achenes 1.3-1.8 cm long,
rather densely covered with very short, glandular hairs; corona 0.4—0.7
mm long, lobed almost to base.

A monotypic section.

Note. The most isolated section of the genus which, apparently,
deserves to be split off as a separate genus. Its only species is
morphologically very similar to the species of Hendelia Heimerl. and
Pseudohandelia Tzvel.; however, according to the structure of the
achenes, florets, and pollen grains, it is much closer to the species of
section Tanacetopsis Tzvel. of Cancrinia, with which I, too, relate it.

1. C. tadshikorum (Kudr.) Tzvel. comb. nova.—Tanacetum
tadshikorum Kudr. in Bull. Univ. As. Centr. XIX (1934) 163.—
Chrysanthemum tadshikorum Kudr. ibid. 163, in syn., nom. altern.—
Pyrethrum tadshikorum Kudr. ibid. 163, in syn., nom. altern.—
Lepidolopsis tadshikorum (Kudr.) Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XIX (1959) 375.—Ic.: Kudr. op. cit. 169.

Perennial. Monocarpic, 30-150 cm high plant, with thick taproot,
more or less sparsely floccose-tomentose with long, flexuous, simple
hairs. Stem usually solitary, erect, strongly thickened at base and
covered with numerous sheaths of dead leaves with dense tomentum in
their axils, more sparsely leafy, branched only at apex (in inflorescence).
Leaves more or less grayish from lax tomentum, with inconspicuous
punctate-glandular hairs; basal leaves 30-35 cm long, 10 cm wide,
often withering before flowering, on petioles strongly thickened at
base, but often with reduced lateral segments close to their base, with

301

286

2—4-pinnately cut lamina, oblong, or linear-oblong; apical lobes some-
what thick, lanceolate or linear-lanceolate, usually to 1 mm, less often
to 1.5 mm wide, gradually narrowed toward apex into rather long,
cartilaginous cusp; lower cauline leaves like basal, middle and upper
reduced, sessile. Capitula quite numerous (more than 40), in dense,
compound, corymbs, to 25 cm in dia. Involucre 3—4 mm in dia and 3-
4 mm long, usually pubescent to subglabrous; involucral bracts herba-
ceous, outer lanceolate-subulate, more or less lacking border, inner
linear, 1.5—2 times as long, with rather wide, light colored, membra-
nous border. Corolla of tubular florets 2.3-3 mm long, usually covered
in lower half with short-stalked glandular hairs. Achenes 1.3—1.8 mm
long, about 0.4 mm wide, with 5—7 longitudinal ribs; corona 0.4—0.7
mm long, irregularly divided almost up to base into subulate lobes of
various shapes and sizes. Flowering May to July.

Gravel beds, stony slopes, scrubs in lower and middle mountain
zones.—Soviet Central Asia: Pamiro-Alai (western Tadzhikistan
ranges). Endemic. Described from Hissar Range. Type in Tashkent.

Section 2. Tanacetopsis Tzvel. sect. nova in Addenda, XXV,
876.—Perennial, 20-70 cm high polycarpic plants. Capitula usually
numerous, (1)2—25(40), in lax, less often dense, corymbose-panicles
or corymbs; involucre patelliform, 6-14 mm in dia; receptacle weakly
convex, punctate-tuberculate, glabrous, less often sparsely hairy; corolla
tube shorter and wide, usually glabrous (hairy in two species). Achenes
1-2.4 mm long, glabrous; corona 0.1-1.5 mm long, often strongly
truncate (narrower on ventral side), more or less divided along margin
into lobes of various shapes and sizes or irregularly toothed.

Type of section: Cancrinia mucronata (Rgl. and Schmalh.) Tzvel.

Note. The largest section of the genus, like section Xanthoglossa
(DC.) Sch. Bip. of the genus Tanacetum L., comprising a large number
of close ecogeographical races, which are often difficult to distinguish.
The species of the section, also similar in their habit to those of section
Xanthoglossa, are easily distinguished from them by their homogamous
capitula, patelliform involucres (at anthesis), and pubescence consisting
almost exclusively of simple hairs.

Series 1. Goloskokoviae Tzvel.—Stem tomentose at base in axils
of sheaths of dead leaves; capitula 10-40, in regular corymbs; corona
of achenes 0.4—0.6 mm long, strongly truncate and 5—7-lobed up to
base.

2. C. goloskokovii (Poljak.) Tzvel. comb. nova.—Lepidolopsis
goloskokovii Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XIX (1959) 376.

302

287

Perennial. Plants 30-70 cm high, with thick many-branched
rhizome, usually with occasional, long, simple hairs to more or less
glabrous. Stems more or less numerous, erect, sparsely leafy, whitish-
tomentose at base in axils of sheaths of dead leaves, branched only
above. Leaves glabrous or subglabrous, lacking distinct punctate
glandular hairs; basal leaves up to 25 cm long and 3-4 cm wide, on
rather long, basally thickened petiole, with twice or thrice pinnately
cut, broad linear lamina; terminal segments linear or lanceolate, up to
1.2 mm wide, and 1-6 mm long, with short cartilaginous cusp; cauline
leaves like basal, but smaller, middle and upper sessile. Capitula 10—
40, on up to 2.5 cm long peduncle, less often to 5 cm, in compound
corymbs. Involucre 6-9 mm in dia, 3.5-4 mm long, glabrous or
subglabrous; involucral bracts herbaceous, outer broadly lanceolate,
subobtuse, almost lacking border, inner 1.5—2 times as long, oblong-
linear, with rather wide, light colored, membranous border,
appendiculately broadening at apex. Corolla of tubular florets 1.8—2.2
mm long. Achenes 1.0-1.2 mm long, about 0.4 mm wide, with 5-7
longitudinal ribs; corona strongly truncate, 0.4—0.6 mm long, inequally
5-7 lobed up to base. Flowering June to July.

Stony slopes, rocks in middle mountain zone.—Soviet Central Asia:
Dzhungaria-Tarbagatai (Belgain Mountains in Dzhungarian Alatau),
Tien Shan (eastern spurs of Trans-Ili Alatau). Endemic. Described
from Trans-Ili Alatau. Type and isotypes in Leningrad; isotypes in
Alma-Ata.

Series 2. Mucronatae Tzvel.—Stems not tomentose at the base;
capitula (1)2—15(20), in lax corymbose-panicles or corymbose-racemes,
less often in regular corymbs; corona of achenes 0.4—1.5 mm long,
more or less truncate and divided up to middle or base in lobes of
various shapes and sizes, less often irregularly toothed.

3. C. subsimilis (Rech. f. ) Tzvel. comb. nova.—Chrysanthemum
subsimile Rech. f. in Koie and Rech. f. Symb. afghan. II (1955) 49.—
Tanacetum subsimile (Rech. f.) Koval. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk Uzb. SSR, XVI (1961) 23.—Ic.: Rech. f. op. cit. 50, fig.
39.

Perennial. Plants 25-60 cm high, with thick many-branched
rhizome, dull green or grayish-green from rather dense pubescence of
long, erect, simple hairs. Stems usually rather numerous, erect or
ascending at base, sparsely leafy, more or less branched mainly above.
Leaves dull-green or grayish-green from rather dense pubescence, with
inconspicuous punctate-glandular hairs; basal leaves up to 15 cm long
and 3 cm wide, on rather long, petiole strongly thickened at base, with

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288

2-3 pinnatisect broadly linear or oblong-linear lamina, terminal seg-
ments lanceolate or linear, somewhat thick, up to 1 mm wide, with
rather long cartilaginous cusp; cauline leaves like basal but reduced,
middle and upper sessile. Capitula (1)2—10(12), on up to 1-1.5 cm
long peduncle in rather dense, compound corymbs, sometimes almost
umbellate. Involucre 7-11 mm in dia, 3.5-5.0 mm long, more or less
pubescent; involucral bracts herbaceous, outer lanceolate, with narrow
membranous border, inner 1.5-2 times as long, oblong-linear, with
wide, brownish, usually light colored border, appendiculately broadened
at apex. Corolla of tubular florets 2.0-2.4 mm long. Achenes 1.6—2.2
mm long and about 0.6 mm wide, with 5-6 longitudinal ribs; corona
distinctly truncate, 0.5—0.9 mm long, divided up to middle or slightly
deep in to wide obtuse lobes or various shapes and sizes. Flowering
July to August.

Screes, rocks, stony slopes above 3,000 m.—Soviet Central Asia:
Pamiro-Alai Region (region of Shugnan Range). General distribution:
Iran-Afghanistan (Afghanistan). Described from Afghanistan. Type in
Vienna.

Note. The species was repeatedly collected from the Shakh-Dara
River basin, but reported as a new species for the USSR only recently
by S.S. Kovalevskaya. Specimens from the USSR correspond fully to
the original description and diagram of the species.

4. C. mucronata (Rgl. and Schmalh.) Tzvel. comb. nova.—
Pyrethrum mucronatum Rgl. and Schmalh. in Tr. Peterb. Bot. Sada, V,
2 (1878) 619, p. p.; O. and B. Fedtsch. Perech. Rast. Turk. IV, 187.—
Chrysanthemum mucronatum (Rgl. and Schmalh.) Minkw. and Knorr.
Rast. Chimkentsk. Uezda (1910) 112; B. Fedtsch. Rast. Turk. 737.—
Tanacetum mucronatum (Rgl. and Schmalh.) M. Pop. in Tr. Uzbeksk.
Gos. Univ. Nov. Ser. No. 27, Biol. XIV (1940) 80.—Lepidolopsis
mucronata (Rgl. and Schmalh.) Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XIX (1959) 375.—Exs.: GRF No. 3267.

Perennial. Plants 20-50 cm high, with thick many-branched
rhizome, dull green from sparse pubescence of long, erect, simple
(mixed with bifid) hairs. Stems usually numerous, erect or ascending
at base, sparsely leafy, more or less branched above middle. Leaves
dull green from sparse pubescence, besides short papillae, with
inconspicuous punctate-glandular hairs; basal leaves up to 15 cm long,
3 cm wide, on rather long petiole basally strongly thickened, with
twice or thrice pinnately cut oblong-linear lamina; terminal segments
lanceolate or narrow-linear, up to 0.6 mm wide, somewhat thick, with
rather long cartilaginous cusp; cauline leaves strongly reduced, middle
and upper (often also lower) sessile. Capitula (1)2—10(12), on long (up

289

to 10-15 cm long) slender peduncles arising from axils of upper leaves,
in depauperate corymbose-raceme of corymbose-panicles, less often in
more dense corymbs (var. corymbulosa Winkl.). Involucre 7—-10(12)
mm in dia, 3-4 mm long, more or less sparsely hairy to subglabrous;
involucral bracts herbaceous, outer lanceolate, with narrow membranous
border, inner 1.5—2 times as long, oblong-linear, with wide, brownish,
usually light colored, membranous border, appendiculately broadened
at apex. Corolla of tubular florets 1.8-2.5 mm long. Achenes 1.6—2.4
mm long, about 0.6 mm wide, with 5, less often 6-7, longitudinal ribs;
corona slightly truncate, 0.8—1.5 mm long, divided up to or almost to
base into 4-7 unequal, more or less aristate lobes. Flowering July to
August.

Screes, rocks, stony slopes above 2,000 m.—Soviet Central Asia:
Pamiro-Alai Region (Kugitang, Hissar, Zeravshan, and Turkestan
ranges). Endemic. Described from the Zeravshan River basin. Lecto-
type (collections of O. Fedtschenko) in Leningrad; isotypes in Taskents.

Note. This polymorphic species was divided only very recently by
S.S. Kovalevskaya into several completely separate ecogeographical
races, differing mainly in the position of the capitula and the structure
of the involucre. Possibly specimens with a denser and almost regular
corymbose inflorescence are close to the previous species and form
the particular higher mountain race, called “Tanacetum mucronatum
var. corymbulosum Winkl.” in the herbarium of C. Winkler. Their
range coordinates fully with that of the typical form.

Very close to it is the Afghan species C. tripinnatifida (Oliv.)
Tzvel. comb. nova. [=7Tanacetum tripinnatifidum Oliv. in Hook Icon.
pl. XXIV (1894) t. 2306], which differs from C. mucronata by having
weaker pubescence and longer terminal segments of the leaves. The
isotype of C. tripinnatifida is preserved in the Herbarium of the
Botanical Institute, Academy of Sciences of the USSR.

5. C. setacea (Rgl. and Schmalh.) Tzvel. comb. nova.—Pyrethrum
setaceum Rgl. and Schmalh. in Izv. Obshch. Lyubit. Estestv., Antrop.
i E’tnogr. XXXIV, 2 (1882) 44; O. and B. Fedtsch. Perech. Rast.
Turk. IV, 187.—?P. setifolium Rgl. in Delect. Sem. Horti. Petrop.
(1876) 36, nom. nud.; Trautv. in Tr. Peterb. Bot. Sada. VIII, 2, 456,
nom. nud.—Tanacetum setaceum (Rgl. and Schmalh.) Oliv. in Hook.
Icon. pl. XXIV (1894) sub tab. 2306 in obs. non Tausch. 1829.—T.
regelii Koval. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk Uzb. SSR,
XVI (1961) 27, nom. nov.—Chrysanthemum setaceum (Rgl. and
Schmalh.) B. Fedtsch. Rast. Turk. (1915) 737.—Lepidolopsis setacea
(Rgl. and Schmalh.) Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XIX (1959) 376.

30

mm

290

Perennial. Plants 25-70 cm high, with thick many-branched rhi-
zome, usually covered with long, flexuous simple (mixed with bifid)
hairs, sometimes subglabrous. Stems usually rather numerous, erect or
ascending at base, sparsely leafy, more or less branched above. Leaves
green, more or less sparsely hairy to subglabrous, with inconspicuous
punctate-glandular hairs; basal leaves 25-30 cm long, 6 cm wide,
otherwise like previous species, but terminal segments usually broader
(up to 1 mm wide). Capitula (1)4—12(15); on long (up to 15 cm)
slender peduncles arising from axils of upper leaves, in depauperate
corymbose-panicles (rarely racemes). Involucre 7—10(12) mm in dia
and 3-4 mm long, with semi-erect hairs at base and dorsally; involucral
bracts herbaceous, outer narrow-lanceolate, usually lacking membranous
border, inner not more than 1.5 times as long, oblong-linear, with rather
wide, light colored or brownish, membranous border, appendiculately
broadened at apex. Corolla of tubular florets 2-3 mm long. Achenes
1.5-2.5 mm long, more or less 0.6 mm wide, with 5-7 longitudinal
ribs and weakly truncate 0.8—1.3 mm long corona, divided to or almost
to base into 4—7 aristate-toothed lobes. Flowering July to August.

Rocks, stony slopes above 2,000 m.—Soviet Central Asia:. Pamiro-
Alai Region (northern slope of Turkestan Range and western Alai
Range). Endemic. Described from Turkestan Range. Type in Leningrad.

6. C. ferganensis (Koval.) Tzvel. comb. nova.—Tanacetum
ferganense Koval. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk Uzb. SSR,
XVI (1961) 31.—Pyrethrum mucronatum auct. p. p.

Perennial. Plants 25-60 cm high, with thick many-branched
rhizome, more or less covered with short and long, erect, simple (mixed
with bifid) hairs. Stems usually rather numerous, erect or ascending at
base, sparsely leafy, more or less branched above. Leaves usually dull-
green from rather dense pubescence, with quite distinct punctate-
glandular hairs; basal leaves 18-20 cm long, 5-6 cm wide, on rather
long petiole, strongly thickened at base, with twice or thrice pinnately
cut, oblong or linear-oblong lamina; terminal segments lanceolate or
linear, up to 1 mm wide, with very short, cartilaginous cusp; cauline
leaves strongly reduced, except lowermost sessile. Capitula (1)2—6(1),
on long (up to 10-15) slender peduncles, in depauperate corymbose-
racemes. Involucre 7-14 mm in dia; 3-4 mm long, more or less hairy;
involucral bracts herbaceous, outer numerous, lanceolate-linear, lacking
border, inner not more than 1.5 times as long, oblong-linear, with
wide, light-colored, membranous border, more or less broadened at
apex. Corolla of tubular florets 1.8-2.5 mm long. Achenes 1.8—2.4 mm
long, about 0.6 mm across, with 5—7 longitudinal ribs and distinctly
truncate 0.8—1.2 mm long corona, divided up to or almost to base into

306

291

more or less aristate-toothed lobes of various shapes and sizes. Flow-
ering July to August.

Rocks, stony slopes above 2,000 m.—Soviet Central Asia: Tien
Shan (Fergana and Chatkal ranges). Endemic. Described from Chatkal
Range. Type in Tashkent.

7. C. submarginata (Koval.) Tzvel. comb. nova.—Tanacetum
submarginatum Koval. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk. Uzb.
SSR, XVI (1961) 30.—Pyrethrum mucronatum auct. p. p.

Perennial. 25-60 cm high plants, with thick many-branched
rhizome, more or less covered with long, erect, simple (mixed with
bifid) hairs. Stems usually rather numerous, erect or ascending at base,
sparsely leafy, more or less branched above. Leaves green, usually
sparsely hairy, with inconspicuous punctate-glandular hairs; basal leaves
up to 15 cm long and 4 cm wide, on rather long petiole strongly
thickened at base, with twice or thrice pinnately cut, usually oblong
lamina; terminal segments lanceolate or lanceolate-linear, up to 1.2
mm wide, with rather long, cartilaginous cusp; cauline leaves strongly
reduced, lowermost sessile. Capitula (1)3—12(15), on long (up to 10-
15), slender peduncles, in depauperate corymbose-racemes or
corymbose panicles. Involucre 6-10 mm in dia, 3-4 mm long, more or
less hairy to subglabrous; involucral bracts herbaceous, outer numerous,
lanceolate-linear or lanceolate-subulate, like middle, lacking
membranous border, inner oblong-linear, 1.5, less often 2 times as
long, with very narrow light-colored, membranous border; corolla of
tubular florets 1.8-2.5 mm long. Achenes 1.6—2.2 mm long, 0.6-0.8
mm wide, with 5, less often 6—7, longitudinal ribs and slightly truncate
0.7—1.2 mm long corona, divided to or almost to base into more or less
aristate-toothed lobes of various shapes and sizes. Flowering July to
August.

Rocks, stony slopes above 2,000 m.—Soviet Central Asia: Tien
Shan (Pskem, Ugam, Chatkal, and Tashkent Alatau ranges). Endemic.
Described from Tashkent Alatau (Greater Chimgan). Type in Tashkent.

8. C. pjataevae (Koval.) Tzvel. comb. nova.—Tanacetum
pjataevae Koval. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk Uzb. SSR,
XVI (1961) 27.—Pyrethrum mucronatum auct. p. p.

Perennial. Plants 20-50 cm high, with thick, many-branched
rhizome, dull green from sparse pubescence of long, erect, simple
(sometimes mixed with bifid) hairs. Stems usually rather numerous,
erect or ascending at base, sparsely leafy, more or less branched above
middle. Leaves dull green from sparse pubescence. Often also with
short papillae, with inconspicuous punctate-glandular hairs; basal leaves

307

292

up to 15 cm long, 3 cm wide, on rather long, basally strongly thick-
ened petiole, with twice or thrice pinnately cut, oblong-linear lamina;
terminal segments lanceolate or lanceolate-linear, up to 1 mm wide,
with rather long, cartilaginous cusp; cauline leaves strongly reduced,
middle and upper sessile. Capitula (1)5—15(20), on peduncles up to 4
cm, less often to 6 cm long, in depauperate compound corymbs. In-
volucre 7-10 mm in dia, more or less hairy; involucral bracts herba-
ceous, outer lanceolate-linear, lacking membranous border, inner ob-
long-linear, 1.5 times, less often 2 times as long, with quite wide,
light-colored, membranous border. Corolla of tubular florets 1.8—2.5
mm long. Achenes 1.3—-1.8 mm long, about 0.6 mm wide, with 5, less
often 6-7 longitudinal ribs and weakly truncate 0.4—1.0 mm long co-
rona, divided up to or almost to base into more or less aristate-toothed
lobes of various shapes and sizes. Flowering June to August.

Rocks, stony slopes above 2,000 m.—Soviet Central Asia: Tien
Shan (Karatau Mountains). Endemic. Described from Central Karatau.
Type in Tashkent.

9. C. karatavica Tzvel. nomen. novum.—Tanacetum karataviense
Koval. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk Uzb. SSR, XVI (1961)
28, non Cancrinia karataviensis Poljak., 1959.

Perennial. Plants 20-50 cm high, with thick, many-branched root,
usually covered with sparse, long, erect, simple hairs, sometimes
glabrous. Stems usually rather numerous, erect or ascending at base,
sparsely leafy, more or less branched above. Leaves like previous
species, but glabrous or subglabrous (with occasional hairs on midrib)
and with rather numerous punctate-glandular hairs. Capitula (1)5-
15(20), on up to 2 cm, less often 4-5 cm long peduncles, more or less
dense, compound corymb. Involucre 6—8(10) cm in dia and 3-4 mm
long, glabrous; involucral bracts herbaceous, outer lanceolate-linear,
lacking membranous border, inner oblong-linear, 1.5 times, less often
2 times as long, with quite wide, light-colored, membranous border.
Corolla of tubular florets 1.8-2.4 mm long. Achenes 1.5—2.0 mm long
and about 0.6 mm wide, with S, less often 6-7, longitudinal ribs and
weakly truncate 0.4-1.0 mm long corona, divided to or almost to base
into more or less aristate-toothed lobes of various shapes and sizes.
Flowering May to July.

Rocks, stony slopes in middle mountain zone.—Soviet Central
Asia: Tien Shan (Karatau Mountains), Pamiro-Alai (Nuratau and Aktau
mountains). Endemic. Described from eastern Karatau. Type in
Tashkent.

Note. The lowest montane ecogeographical race of the series. De-
spite the considerable gaps in the range, the specimens from Karatau

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293

and Nuratau are very similar and were quite rightly placed in one
species by Kovalevskaya.

Series 3. Pamiroalaicae Tzvel.—Stem densely lanate-tomentose
at base in axils of sheaths of dead leaves; capitula (1)2—10(20) on a
single stem, in depauperate corymbose-raceme or corymbose-panicle;
corona of achene 0.3—0.7 mm long, more or less divided or irregularly
toothed, sometimes strongly truncate.

10. C. pamiralaica (Koval.) Tzvel. comb. nova.—Tanacetum
pamiralaicum Koval. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk Uzb.
SSR, XVI (1961) 35.

Perennial. Plants 20-50 cm high, with thick many-branched
rhizome, dull green from rather dense pubescence of long and short
simple (with bifid) hairs. Stems usually rather numerous, erect, densely
lanate-tomentose at base in axils of leaf sheaths, sparsely leafy, more
or less branched above. Leaves dull green from rather dense pubescence,
with inconspicuous punctate-glandular hairs; basal leaves up to 15 cm
long, 3 cm wide, on quite long (sometimes almost as long as lamina),
petiole strongly thickened at base, with twice or thrice pinnately cut,
usually oblong lamina; leaf segments 4-8 on each side, somewhat
distant, with narrow rachis; terminal segments lanceolate to linear, to
1.2 mm wide, with short cartilaginous cusp; cauline leaves strongly
reduced and less dissected except lower, sessile. Capitula (1)2—10(12),
on quite long (up to 10 cm), slender peduncles, in depauperate
corymbose-racemes or corymbose-panicle. Involucre 6-10 mm in dia
and 3-4 mm long, more or less finely tomentose near base and dorsally
on bracts; involucral bracts herbaceous, outer lanceolate, lacking
membranous border, inner 1.5—2 times as long, oblong-linear, with
quite narrow, but apically more or less broadened, light colored,
membranous border. Receptacle glabrous. Corolla of tubular florets
1.8-2.3 mm long. Achenes 1.2—1.8 mm long and about 0.6 mm wide,
with 5—7 longitudinal ribs and strongly truncate (lobes on ventral side
almost a half as long as on dorsal) 0.4—0.7 mm long corona, divided
almost to base into 5-7 irregularly toothed lobes. Flowering July to
August.

Rocks, stony slopes, screes above 2,000 m.—Soviet Central Asia:
Pamiro-Alai Region (Darvaz, Vakhsh, Rushan, Yazgulem, and Hissar
ranges). Endemic. Described from Hissar Range. Type in Tashkent.

Note. According to Kovalevskaya, it differs from the very close Afghan
species C. eriobasis (Rech. f.) Tzvel. comb. nova. [ = Chrysanthemum
eriobasis Rech. f. in Koie and Rech. f. Symb. afghan. II (1955) 44] by
having the capitula on much longer peduncles which are aggregated in
very lax corymbose-racemes (and not rather dense corymbs).

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294

11. C. urgutensis (M. Pop.) Tzvel. comb. nova.—Pyrethrum
urgutense M. Pop. in Tr. Uzbeksk. Gos. Univ. Nov. Ser. No. 27, Biol.
XIV (1941) 80, diagnosis in Russian and in Addenda, XXV, 874.—
Tanacetum urgutense (M. Pop.) Koval. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk Uzbeksk. SSR, XVI (1961) 36.

Plants 18-40 cm high, with thick many-branched rhizome, dull
green from rather dense pubescence of long, flexuous, simple hairs.
Stems usually rather numerous, erect or ascending, densely lanate
tomentose at base in axils of leaf sheaths, sparsely leafy, more or less
branched above. Leaves dull green from rather dense pubescence,
lacking punctate-glandular hairs; basal leaves up to 10 cm long, 2.5
cm wide, on rather long (sometimes almost as long as lamina) petiole,
with twice or thrice pinnately cut, linear-oblong lamina; leaf segments
distant, 4-8 on each side; terminal segments lanceolate or linear, up to
1 mm wide, with short cartilaginous cusp; cauline leaves strongly
reduced and less divided except lower, sessile. Capitula (1)2—6(8), on
long (up to 10-15 cm), slender peduncles, in very lax, corymbose-
racemes. Involucre 7-12 mm in dia, 3.5-5 mm long, more or less
appressed hairy; involucral bracts herbaceous, outer lanceolate, few,
lacking membranous border, inner oblong-linear, 1.5 times, less often
2 times as long, with quite wide, light colored, membranous border.
Receptacle sparsely hairy, less often glabrous. Corolla of tubular florets
2.2-2.8 mm long, densely hairy in upper part. Achenes 2.0-2.4 mm
long, about 0.6 mm wide, with 5, less often 6, longitudinal ribs and
usually nontruncate 0.3-0.6 mm long corona, irregularly sharply toothed
usually not more than to middle. Flowering June to August.

Rocks, stony slopes at 2,000-3,000 m.—Soviet Central Asia:
Pamiro-Alai Region (Turkestan and Zeravashan ranges). Endemic.
Described from Zeravshan Range. Type in Tashkent.

Note. As also in the next species, the corolla densely hairy in the
upper part, a character of wide occurrence in the genus Trichanthemis
Rgl. and Schmalh. but not found in Tanacetum L. and Pyrethrum Zinn.

12. C. botschantzevii (Koval.) Tzvel. comb. nova.—Tanacetum
botschantzevii Koval. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk Uzb.
SSR, XVI (1961) 36.

Perennial. Plants 25-45 cm high, with thick many-branched
rhizome, dull green or grayish-green from rather dense pubescence of
flexuous simple hairs. Stems usually rather numerous, erect, lanate-
tomentose at base in axils of sheaths of dead leaves, sparsely leafy,
strongly branched from middle or below. Leaves lacking punctate-
glandular hairs; basal leaves up to 10 cm long and 2 cm wide, on
rather long petiole, with twice (sometimes more or less thrice) pinnately

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295

cut, oblong-linear or broadly linear lamina; leaf segments including
terminal segments like previous species; cauline leaves more or less
reduced and less dissected. Capitula 3-8 on a single stem, on very
long (up to 15 cm), slender peduncles in lax corymbose-racemes.
Involucre 7-10 mm in dia, 3—4 mm long, more or less appressed hairy;
involucral bracts like previous species. Receptacle sparsely hairy to
subglabrous. Corolla of tubular florets 1.8—2.4 mm long, densely hairy
above. Achenes 2.0-2.4 mm long, about 0.6 mm wide, with 5(6)
longitudinal ribs and strongly truncate, corona 0.6—1.0 mm long, usually
irregularly divided almost to base into unequal acuminate lobes.
Flowering July.

Screes and rocks at 2,000 m.—Soviet Central Asia: Pamiro-Alai
Region (western spurs of Hissar Range). Endemic. Described from
Baisuntau Mountains (upper reaches or Yakkabag-Darya River in
vicinity of the village of Tash-Kurgan, 12.VII.1936, Nos. 902 and
910, V. Botschantzev and A. Butkov). Type in Tashkent.

13. C. nevskii Tzvel. nom. nov.—Tanacetum krascheninnikovii
Nevski in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, IV (1937) 282, non
Cancrinia krascheninnikovii (Rubtz.) Poljak. 1959.—Lepidolopsis
krascheninnikovii (Nevski) Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XIX (1959) 376.—Ic.: Nevski, op. cit. 283, fig. 7.

Perennial. Plants 20-50 cm high, with thick many-branched
rhizome, grayish or dull green from rather dense pubescence of long,
flexuous, simple hairs. Stems usually rather numerous, erect, densely
lanate-tomentose at base in axils of leaf sheaths; sparsely leafy, more
or less branched above. Leaves usually grayish from dense, almost
tomentose pubescence, with inconspicuous punctate-glandular hairs;
basal leaves up to 20 cm long, 3-4 cm wide, on rather long petiole
strongly thickened at base, twice or thrice pinnately cut, linear-oblong
lamina; leaf segments quite distant, 4-10 on each side; terminal
segments lanceolate or linear, up to 1.5 mm wide, somewhat thick,
with very short cartilaginous cusp; cauline leaves strongly reduced and
less dissected, except lower, sessile. Capitula (1)5—15(20), on rather
long (up to 10 cm) peduncles in depauperate corymbose-racemes or
corymbose-panicles. Involucre 6-10 mm in dia and 3—4.5 mm long,
more or less finely tomentose to subglabrous; involucral bracts
herbaceous, outer few, lanceolate-ovate, subobtuse, almost lacking
membranous border, inner oblong-linear and oblong, 1.5 times, less
often 2 times as long, with quite narrow, light colored, membranous
border. Receptacle glabrous or sparsely hairy. Corolla of tubular florets
1.6—2.2 mm long. Achenes 1.8—2.4 mm long and 0.6-0.8 mm wide,
with 5, less often 6, longitudinal ribs and weakly truncate 0.3-0.7 mm

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long corona, divided almost up to base into longitudinal, more or less
aristate-toothed lobes. Flowering June to July.

Stony slopes, rocks (especially on gypsum rocks) at 1,500—2,500
m.—Soviet Central Asia: Pamiro-Alai Region (Kugitang and Baisuntau
mountains). Endemic. Described from Kuguitang Mountains. Type and
isotypes in Leningrad.

Series 4. Platyrachides Tzvel.—Stems densely lanate-tomentose
at base in axils of sheaths of dead leaves. Capitula numerous (more
than 20 on a single stem), at apices of divaricate branches in very lax,
panicles or corymbose-panicle; corona of achenes about 0.1 mm long,
blunt-toothed along margin.

14. C. paropamisica (Krasch.) Tzvel. comb. nova.—Pyrethrum
paropamisicum Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
IX (1946) 166.—Lepidolopsis paropamisica (Krasch.) Poljak. in Bot.
Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 376.—Tanacetum
paropamisicum (Krasch.) Koval. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk Uzb. SSR, XVI (1961) 30.—Ic.: Krasch. op. cit. 169, fig. 7.

Perennial. Plants 30-70 cm high, with strongly thickened, many-
branched tap root, dull green or grayish-green from rather dense
pubescence of long, flexuous, simple hairs, usually on papilliform’
tubercles, and shorter, appressed simple hairs. Stems less numerous or
solitary, always clustered at base and densely lanate-tomentose in axils
of leaf sheaths, erect, dichotomously branched from middle or slightly
below. Leaves dull green or grayish-green from rather dense pubescence,
mixed with inconspicuous punctate-glandular hairs; basal and lower
cauline leaves up to 15 cm long and 3-4 cm wide, on rather long
petioles strongly thickened at base, with twice or thrice pinnately cut,
oblong to oblong-linear lamina; leaf segments 3-8 on each side, quite
distant, alternate lobes less distant; terminal segments broadly lanceolate
to linear, up to 1.5 mm wide, with short cartilaginous cusp; middle and
upper cauline leaves strongly reduced or less dissected sessile. Capitula
numerous (20-50), on rather long (up to 6-8 cm) peduncle in
depauperate and broadly paniculate or corymbose-paniculate
inflorescence. Involucre 6-10 mm in dia, 3.0-4.5 mm long, finely
grayish tomentose; involucral bracts herbaceous, outer lanceolate, fewer,
lacking membranous border, inner 1.5 times, less often 2 times as
long, oblong-linear, with rather narrow, light colored, membranous
border. Receptacle sparsely hairy. Corolla of tubular florets 2.0—2.5
mm long. Achenes 2.0-2.5 mm long, about 0.6 mm wide, with 5-7
longitudinal ribs and about 0.1 mm long blunt-toothed corona. Flowering
May to June.

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Stony slopes, rocks in lower mountain zone.—Soviet Central Asia:
Mountainous Turkmenia (Paropamisus Mountain). General distribution:
Iran(?). Described from the Paropamisus Mountain. Type and isotype
in Leningrad.

Note. So far the species is known only from the type specimens,
which are without sheaths on the basal leaves and tomentum in their
axils. However, there is dense tomentum in the very closely related
(perhaps, even identical) Iranian species C. platyrachis (Boiss.) Tzvel.
comb. nova [= Pyrethrum platyrachis Boiss. Fl. or. Ill (1875) 356],
differing from C. paropamisica by only a slightly larger size of the
capitula and the whole plant.

Series 5. Santoanae Tzvel.—Stems not tomentose at base; capitula
solitary or few (up to 5 on a single stem) and then in lax corymbose-
raceme; corona of achenes 0.3—-0.4 mm long, irregularly aristate-toothed
along margin up to middle.

15. C. santoana (Krasch., M. Pop. and Vved.) Poljak. in Bot.
Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 370.—Pyrethrum
santoanum Krasch. M. Pop. and Vved. in Tr. Sredneaz. Gos. Univ.
Ser. VIIc, III (1928) 102.—Chrysanthemum santoanum Krasch. M.
Pop. and Vved. ibid. nom. altern.—Exs.: H.F.A.M. No. 476 (isotypes).

Perennial. Plants 8—30 cm high, with thickened tap root, grayish-
green from rather dense pubescence of long, flexuous, simple hairs.
Stems less numerous or solitary, always closely clustered, erect, simple
or with few lateral branches. Leaves grayish-green from rather dense
erect pubescence, with inconspicuous punctate-glandular hairs; basal
and lower cauline leaves up to 5—6 cm long, 0.8 cm wide, on rather
long petiole, strongly thickened at base, with pinnatisect lanceolate-
linear to oblong-linear lamina; leaf segments S—10 on each side, usually
3-5 parted, less often entire or pinnately cut; apical segments linear to
oblong, up to 1.2 mm wide, with short cartilaginous cusp;. middle and
upper cauline leaves few, strongly reduced, sessile or sub-sessile.
Capitula solitary or 2-5, on rather long peduncles, arising from axils
of middle and upper cauline leaves, and then in very lax corymbs.
Involucre 7-10 mm in dia, 3.5—5.5 mm long, more or less appressed
hairy; involucral bracts herbaceous, outer narrow-lanceolate, lacking
or more or less lacking membranous border, inner 1.5 times, less often
almost 2 times as long, oblong-linear, with rather wide, light-colored,
membranous border, appendiculately broadened at apex. Receptacle
glabrous, very rarely hairy. Corolla of tubular florets, 1.8—2.5 mm
long. Achenes 1.2—1.6 mm long, about 0.4 mm wide, with 5 longitudinal
ribs and irregularly aristate-toothed (usually not deeper than middle)
0.3-0.4 mm long corona. Flowering May to June.

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Stony and clayey slopes, gravel beds up to lower mountain zone.—
Soviet Central Asia: Syr-Darya, Pamiro-Alai Region (northern and
northwestern foothills). Endemic. Described from northwestern foothills
of Pamiro-Alai (near village of Santo). Type in Tashkent; isotypes in
Leningrad.

Section 3. Matricarioides Tzvel. sect. nova. in Addenda, XXV,
876.—Biennial or perennial(?) polycarpic 5—20 cm high plants. Capitula
solitary at apices of stem and leafy branches arising from its base;
involucre patelliform, 7-12 mm in dia; receptacle strongly convex,
conically hemispherical, punctate-tuberculate, glabrous; corolla tube
very short and wide, glabrous. Achenes 1.8—2.2 mm long, glabrous;
corona 0.8-1.0 mm long, weakly truncate, irregularly obtusely lobed
(lobes as many as number of ribs) almost to middle (or slightly deeper),
small toothed on margin.

A monotypic section.

Note. This section is closely related through C. santoana to the
previous section, representing, as it were, the ultimate stage of the
gradual ephemeralization of the mesophilic high-mountain ancestor,
culminating with the formation of two desert ephemerals—species of
the genus Microcephala Pobed. However, the relationship of this section
to section Cancrinia also is no less close, one of whose species—C.
lasiocarpa Winkl.—is identical with C. discoidea, differing from it
only by having hairy achenes with a strongly elongate corona.

16. C. discoidea (Ldb.) Poljak. comb. nova. in litt.—C.
brachypappus Winkl. in Tr. Peterb. Bot. Sada, XII (1892) 29.—
Pyrethrum discoideum Ldb. Ic. pl. Fl. Ross. impr. Alt. II (1830) t. 153
and Fl. Alt. IV (1834) 119; DC. Prodr. VI, 59; Ldb. Fl. Ross. II, 556;
O. and B. Fedtsch. Perech. Rast. Turk. [V, 188.—Tanacetum ledebourii
Sch. Bip. Tanacet. (1844) 47, nom. nov. non T. discoideum Rchb. f.
1853.—Matricaria songorica Bge. in Mém. Ac. Sc. Pétersb. VII (1854)
335, nom. nov. non M. discoidea DC. 1837.—M. ledebourii (Sch.
Bip.) Schischk. in Fl. Zap. Sib. XI (1949) 2733.—Chrysanthemum
discoideum (Ldb.) B. Fedtsch. Rast. Turk. (1915) 737, non All. 1785.—
C. ledebourianum Ling in Contr. Inst. Bot. Nat. Ac. Peiping, III (1935)
474, nom. nov.—Ic.: Ldb. op. cit. (1830) t. 153.

Biennial or perennial. Plants 5-20 cm high with slender tap root,
more or less sparsely floccose-tomentose with long simple hairs. Stem
usually branched at base, simple above, very sparsely leafy (with few
strongly reduced leaves only at base), erect or ascending. Leaves
grayish-tomentose to subglabrous, lacking punctate-glandular hairs;
basal leaves numerous, up to 4.0-4.5 cm long and 1.5 cm wide, on

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long (sometimes longer than lamina) petioles strongly thickened at
base; with more or less pinnately parted oblong or ovate lamina; leaf
lobes 2-5 on each side, in turn 2—5-lobed or pinnate, less often all or
some entire; terminal segments ovate to broadly linear, to 1(1.5) mm
wide, obtuse or subacute. Capitula solitary (but often numerous on a
single plant), on long (4-16 cm long), almost leafless, erect peduncles.
Involucre 7-12 mm in dia, 2-4 mm long, more or less sparsely
tomentose to subglabrous; involucral bracts herbaceous, outer few,
linear-lanceolate, acute, more or less lacking membranous border, inner
usually not more 1.5 times as long, linear-oblong, subobtuse, with
rather wide, light colored, membranous border. Corolla of tubular florets
1.2-1.8 mm long, very short and wide (its teeth usually only 1/3-1/2
as long as tube). Achenes 1.8-2.2 mm long, about 0.6 mm wide, with
5 longitudinal ribs and somewhat truncate 0.8-1.0 mm long corona,
obtusely and irregularly 5-lobed up to middle or more. Flowering June
to August.

Stony, rubbly and clayey slopes up to lower mountain zone.—
Western Siberia: Altai (southwestern part); Soviet Central Asia: Aralo-
Caspian Region (northeastern part of Ulutau Mountains), Lake Balkhash
Region, Dzhungaria-Tarbagatai. General distribution: Mongolia
(western part), Dzhungaria-Kashgaria. Described from southwestern
spurs of Altai. Type and isotypes in Leningrad.

Note. A polymorphic species, possibly comprising several
ecogeographical races, differing in the degree of hairiness, shape of
the leaves and structure of the involucre.

Section 4. Cancrinia.—Perennial, less often biennial(?) 2-10 cm
high plant; capitula solitary (but many on a single plant); involucre
patelliform, 10-17 mm in dia; receptacle weakly convex, glabrous,
punctate-tuberculate, with inconspicuous alveolation; corolla tube rather
short and wide, glabrous. Achenes 1.3-2.0 mm long, more or less
sparsely hairy; corona 2.3—3.0 mm long, not truncate, divided into 8-
12 linear or oblong-linear scales to or almost to base, more or less
irregularly toothed along margin.

Type of section: type of genus.

Note. A few species of this section apparently are the result of
secondary adaptation to severe high-mountain conditions of an ancestor
of the C. discoidea type, which had already been subjected to significant
ephemeralization. The Mongolian-Chinese, relatively low mountain
species C. lasiocarpa Winkl. in its outward appearance (and also in
many other more important characters) is still quite similar to C.
discoidea, whereas species of the C. chrysocephala type are already
true high-mountain perennials.

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300

17. C. chrysocephala Kar. and Kir. in Bull. Soc. Nat. Mosc. XV
(1842) 125; Ldb. Fl. Ross. II, 519; O. and B. Fedtsch. Perech. Rast
Turk. IV, 178.

Perennial. Plants 2-10 cm high, with slender, more or less branched
rhizome, grayish from extensive tomentum of long simple hairs. Stems
strongly reduced, scape-like or almost scape-like, usually caespitose
together with short non-flowering shoots. Basal leaves numerous,
densely grayish- or whitish-tomentose, lacking distinct punctate-
glandular hairs, up to 2-3 cm long and 0.8-1.0 cm wide, on rather
long (often longer than lamina), petiole strongly thickened at base,
with more or less pinnately parted, oblong or ovate lamina; leaf
segments (or lobes) 2—4 on each side, linear-oblong to obovate, up to
1.5(2.0) mm wide, obtuse or subobtuse, usually all or some in turn 2-
4 parted or lobed; cauline leaves 1-2, strongly reduced, undivided.
Capitula solitary (but usually many on a single plant), on long (2-8 cm
long) erect peduncles. Involucre 10-17 mm in dia and 4-6 mm long,
depauperate floccose-tomentose; involucral bracts herbaceous, outer
few, lanceolate and linear-lanceolate, inner 1.5 times, less often 2 times
as long, oblong-linear; all bracts with wide, dark brown or brownish,
membranous border, somewhat broadened at apex. Corolla of tubular
florets 2.4—3.0 mm long. Achenes 1.5-2.2 mm long, about 0.6 mm
wide, more or less sparsely hairy in upper part, with 5—6 longitudinal
ribs and short, 2.3-3.0 mm long corona divided up to or almost to base
into 8-12 linear or oblong-linear subacute, more or less irregularly
toothed scales. Flowering July to August. (Plate XIII, Fig. 1.)

Rubbly and stony slopes, gravel beds above 3,000 m.—Soviet Central
Asia: Dzhungaria-Tarbagatai (Dzhungarian Alatau). General distribution:
Dzhungaria-Kashgaria. Described from Dzhungarian Alatau. Type in
Leningrad.

18. C. tianschanica (Krasch.) Tzvel. comb. nova.—C. chryso-
cephala ssp. tianschanica Krasch. in Bot. Mat. Gerb. Glavn. Bot. Sada,
Wa T922 Sih.

Perennial. Plants 2-10 cm high, with slender, more or less branched
rhizome, grayish from extensive lax tomentum of long simple hairs.
Stems strongly reduced, scape-like or almost scape-like, caespitose
with short nonflowering shoots. Leaves like previous species. Capitula
solitary (but many on a single plant), on long erect peduncles. Involucre
10-17 mm in dia, 4-6 mm long, extensively lax floccose-tomentose;
involucral bracts like previous species, but with somewhat narrower,
light brown, membranous border. Corolla of tubular florets 2.4—3.0
mm long. Achenes 1.5—2.2 mm long, about 0.6 mm wide, sparsely
hairy throughout, with 5—6 longitudinal ribs and 2.3-3.0 mm long

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corona, divided to or almost to base into 8-12 linear or oblong-linear
scales. Flowering July to August.

Rubbly and stony slopes above 3,000 m.—Soviet Central Asia:
Tien Shan (central part). General distribution: Dzhungaria-Kashgaria (Tien
Shan). Described from central Tien Shan. Type and isotype in Leningrad.

Note. The discrete range and the absence of transitional forms
with regard to the achene pubescence provide a basis to consider this
subspecies described by Krascheninnikov as a separate species.

GENUS 1537. Lepidolopsis Poljak. ':”

Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959)
374.—Crossostephium auct. non Less.: Rgl. and Schmalh. in Izv.
Obshch. Lyubit. Estestv. Antrop. Etnogr. XXXIV, 2 44 p. p.

Capitula numerous (more than 50), on up to 0.8 cm long peduncles,
in compact spicate-panicles, homogamous, with large number (25-50)
of bisexual, tubular disk florets. Involucre goblet-shaped, obconical at
base, 4-6 mm in dia, 3-4 mm long; involucral bracts coriaceous-
herbaceous, imbricate, in 4 irregular rows, outer broadly lanceolate,
usually lacking membranous border, inner lanceolate-oblong, 2—2.5
times as long, with appendiculately broadened light colored membranous
border at apex. Receptacle weakly convex, short- and obtusely conical,
glabrous, punctate-tuberculate, with inconspicuous alveolation and
abundant resin secretion, solid. Corolla of tubular disk florets yellow,
1.7-2.0 mm long, with short and wide tube slightly and gradually
broadening in upper part, with 5 (less often 4—6) subobtuse, erect,
deltoid teeth, a sixth to a fifth as long as tube. Filaments thickened in
upper part; anthers lacking basal appendage, with vate or oblong ovate
subobtuse apical appendage; pollen grains roundish, spiny. Style bifid;
branches linear, truncate. All achenes similar, glabrous, 1.0-1.6 mm
long, prismatic, gradually narrowed toward base, with 5 prominent
longitudinal ribs; pappus as corona, 0.3—1.0 mm long, divided almost
to base, into 8-12 oblong unequal lobes, ventrally divided up to or
almost to base, unilateral. Perennial herbs; covered with long and short
simple hairs, usually subglabrous with thick oblique rhizome, erect
stem and alternate leaves with 2—3-pinnatisect lamina.

A monotypic genus.

Note. P.P. Poljakov, who described this genus, included in it a
whole series of species of different affinity. I have referred some of
them (species with homogamous capitula) to the genus Cancrinia Kar.

'Treatment by N.N. Tzvelev.
7From Greek words: lepis—scale, and lopsis—not precisely translatable; named
for the pappus of the achene, which is divided into scaly lobes.

3hl7/

302

and Kir. and some (species with heterogamous capitula) to the genus
Tanacetum L. However, Lepidolopsis turkestanica (Rgl. and Schmalh.)
Poljak., being the type species of the new genus, differs from other
species by its unique inflorescence, very similar to the inflorescence of
many worm-woods, as well as by a series of less important characters,
so that the generic name “Lepidolopsis” can be retained for it. On the
basis of the form of the inflorescence and the structure of the achenes,
Lepidolopsis is very close to the genus Crossostephium Less. but the
only species of the latter, C. sinense (L.) Makino, distributed in China
and Japan, is a grayish tomentose half-shrub with heterogamous capitula
and almost equal involucral bracts.

1. L. turkestanica (Rgl. and Schmalh.) Poljak. in Bot. Mat. Gerb.
Bot. Inst. Akad. Nauk SSSR, XIX (1959) 375.—Crossostephium
turkestanicum Rgl. and Schmalh. in Izv. Obshch. Lyub. Estestv., Antrop.
i E’tnogr. XXXIV, 2 (1882) 44; O. and B. Fedtsch. Perech. Rast.
Turk. IV, 203.—Tanacetum johnstonii Hemsl. in Journ. of Bot. XXI
(1883) 135.—T. turkestanicum (Rgl. and Schmalh.) Poljak. 1. c. XV
(1953) 382.—Artemisia turkestanica (Rgl. and Schmalh.) Franch. in
Ann. Sc. Nat. Ser. 6, XVI (1883) 311.—A. chrysostachya Rech. f. and
Koie, Symb. afghan. II (1955) 52.—Pyrethrum johnstonii (Hemsl.)
Buser in Boiss. Fl. or. Suppl. (1888) 299.—Chrysanthemum
chrysostachys (Rech. f. and K6ie) Kitam. im Acta Phyt. et Geobot.
XVI (1957) 34.—Ic.: Rech. f. and Koie, op. cit. fig. 41.

Perennial. Plants 40-100 cm high, with thick oblique rhizome,
usually lacking short nonflowering shoots, usually covered with long
erect and short, papilliform, simple hairs to more or less glabrous.
Stems solitary or few, erect, extensively leafy, usually branched only
in inflorescence. Leaves sparsely hairy to more or less glabrous with
few distinct punctate-glandular hairs; basal (withering early) and lower
cauline leaves up to 10-15 cm long and 3.5 cm wide, with petioles
thickened at base, usually with short lateral segments almost up to
base, lamina broadly linear or oblong, twice or thrice pinnately cut;
apical segments lanceolate or lanceolate linear, somewhat thick, up to
1 mm wide, with short cartilaginous cusp; upper cauline leaves with
more or less reduced lamina, sessile. Capitula numerous, in compressed
spicate panicle; arising from axils of strongly reduced apical leaves in
clusters of 1-10 on very short (up to 0.8 mm long) slender peduncles.
Involucre 4-6 mm in dia, 3-4 mm long, glabrous or subglabrous, usually
with golden tinge. Corolla of tubular florets yellow, 1.7—2.0 mm long.
Achenes 1.0—1.6 mm long and 0.3-0.5 mm wide; corona divided almost
to base on ventral side into 8-12 longitudinal lobes, 0.3-1.0 mm long.
Flowering June to August. (Plate XII, Fig. 2.)

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303

Stony slopes, steppes; up to lower mountain zone.—Soviet Central
Asia: Tien Shan (western part), Syr-Darya Amu-Darya, Pamiro-Alai
Region (excluding Pamir). General distribution: Iran-Afghanistan (Af-
ghanistan). Described from western Tien Shan. Lectotype (Chirchik
River valley near village of Pakrak, VIII, 1876, A. Regel) and isotype
in Leningrad.

GENUS 1538. Tanacetum L. Emend. Tzvel. !:?

L. Sp. pl. (1753) 843 p. p. and Gen. pl. ed. 5 (1754) 366.

Capitula more or less numerous (2-80), in lax or dense corymbs,
less often solitary, always heterogamous, with outer pistillate florets ©
tubular or Ligulate, 1-30 in one row, and much more numerous,
bisexual, tubular disk florets. Involucre goblet-shaped, 3.5-18.0 mm
in dia and 3-8 mm long; involucral bracts coriaceous-herbaceous or
herbaceous, imbricate, in 3-5 irregular rows, more or less scarious on
margin or entirely lacking border; outer bracts lanceolate to broadly
ovate, inner lanceolate-linear to linear-oblong, 1.5-3 times as long.
Receptacle strongly or weakly convex, less often (section Asterotricha)
almost flat, solid, glabrous, weakly punctate-tuberculate. Corolla of
ligulate florets (if present) yellow, with dorsally compressed but
wingless or almost winged tube 1.2—2.5 mm long and limb of various
shapes (from reniform trilobate to oval-oblong with 2-3 more or less
distinct teeth at apex) up to 11 mm long, ligulate florets usually
gradually changing over to pistillate tubular florets, in many respects
similar to bisexual tubular disk florets, but with strongly reduced
stamens and slightly smaller, more or less dorsally compressed corolla
with 2-5 teeth (sometimes differing in shape and size); corolla of
bisexual tubular disk florets yellow, 1.5—3.5 mm long, with relatively
short tube, usually gradually but slightly broadening in upper half,
with 5 erect or weakly deflexed deltoid or lanceolate-deltoid teeth, 2/
11-1/3 as long as tube. Filaments thickened in upper part; anthers
lacking distinct basal appendage, with lanceolate-ovate subobtuse apical
appendage; pollen grains roundish spiny. Styles bifid; style branches
linear, truncate. All achenes similar, glabrous, more or less prismatic-
terete, narrowed towards base, 1.2-3.5 mm long, 0.5-0.9 mm wide;
with 5-10 (very rarely up to 15) more or less prominent longitudinal

'Treatment by N.N. Tzvelev.
*From the Latin word: tanaceta or tanazita—the medieval name of a plant of
unknown origin, probably, belonging to Tanacetum vulgare L.

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304

ribs, scales more compactly appressed to kernel, lacking mucilaginous
cells and secretory canals; pappus 0.1-0.7 mm long, as corona more
or less toothed or lobed along margin, sometimes divided almost to
base. Perennial herbs (very rarely more or less woody at base), usually
less covered with simple, bifid or stellate hairs, with erect or basally
ascending stem, and alternate leaves with more or less pinnately divided,
very rarely (in T. akinfievii) pinnately lobed lamina.

More than 50 species distributed in almost all nontropical countries
of the Northern Hemisphere. Of these, 34 species are found in the
USSR.

Type of genus: Tanacetum vulgare L.

Note. The border between this and the extremely close genus
Pyrethrum Zinn has always been very undefined; consequently, these
genera have usually been separated either on the basis of the presence
or absence of ligulate florets, or on the basis of the peculiar arrangement
and number of the capitula. Neither of these characters holds up to
serious criticism, and not without basis, therefore, many authors in
recent times, while maintaining the separate generic status of
Chrysanthemum L. and Leucanthemum Mill., have combined Tanacetum
and Pyrethrum into the single genus Tanacetum, as was done even by
K.H. Schultz (Bi-pontinus, Uber die Tanaceteen, 1844). However, here
I have retained Pyrethrum as a separate genus, mainly on the basis of
the different color of the ligulate florets in Tanacetum species. This
division is not just a formal one, as it initially appears, but is fully
justified. Actually, all species of Pyrethrum (in the circumscription
accepted here) have white or pink outer ligulate florets, usually with
long ligules, that are missing only in the two cultivated species, so that
the capitula become homogamous. In Tanacetum species, the ligulate
florets are yellow, almost always with strongly reduced ligules, forming
not homogamous but heterogamous capitula with pistillate tubular florets
as a result of gradual reduction. Based on this, the genus Cancrinia
Kar. and Kir. (in the circumscription accepted here), not the genus
Tanacetum, should be considered the closest to Pyrethrum. There is even
a basis to suppose that Pyrethrum may have given rise to Cancrinia with
its homogamous capitula by the complete reduction of its white or pink
florets, and that Tanacetum species then evolved from species of the
Cancrinia type by modification of the outer bisexual tubular florets into
pistillate ones subsequently modified secondarily into yellow ligulate florets.
From this standpoint, in the genus Tanacetum the species with ligulate
florets are more recent than the species having only tubular florets,
which also is usually confirmed by an analysis of other characters.

The genus is divided into four natural sections, which undoubtedly
are related but are so isolated that they could easily be treated as
separate genera.

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305

Entire plant grayish from dense pubescence of stellate hairs;
receptacle almost flat. (sect. ASterotricha.) .....:cccscccsscsesseserseeeees a

Plant, more or less covered with bifid or simple hairs; sometimes
subglabrous; receptacle strongly or weakly CONVEX ...........::000008 3.
Involucre 3.5—6 mm in dia; ligulate florets 2-8, ligule 1.5—-2.5 mm
Nepi SON 26 ea RG el 33. T. turcomanicum (Krasch.) Tzvel.
Involucre 6-10 mm in dia; ligulate florets 8-16, ligule 3-3.5 mm
OAs BROS ARIAL Rnd. 34. T. Walteri (Winkl.) Tzvel.
Plants 30-150 cm high, with densely leafy stem, almost always
lacking short nonflowering shoots; achenes 1.2—2.5 mm long. (sect.
TARR EIut yO SRF Ris lh Re I ARE BU hats. ce 4.
Plants up to 40 (60) cm high, with relatively sparsely leafy stem,
but always with short nonflowering shoots (as rosettes of basal
leaves); achenes, 85~3:5 iin long «...2..44 CY ORR cae 7.
Leaves pinnately parted or pinnately cut, with pinnately lobed, less
often pinnately parted segments; pubescence mainly of simple hairs.

Widely Gisthibimed: Species 1655. ash dW AG AIR =
Leaves twice or thrice pinnately cut; pubescence mainly of bifid
BRIG RRR oe eet vs haan sc esyessuensdaeesnssaiicagnaneneduas dhanltusntieenscn casey 6.
Involucre 7-13 mm in dia; involucral bracts with wider dark brown
mienibranous BOPder !!...12.... ia, 3. T. boreale Fisch. ex DC.
Involucre 5-8 mm in dia; involucral bracts with narrower light
colored or brownish border .............csseeseeseeeeeeees 2. T. vulgare L.
Leaves thrice pinnately cut into narrow-linear and linear-lanceolate
segments. Transcaucasia .............+ 5. T. abrotanifolium (L.) Druce.

Leaves twice pinnately cut into linear and oblong, often irregularly
lobed or toothed segments. Soviet Central Asia ...........::cssccessseeseeenes
PE MY GING nd. sl gid. shaszawnanseveydedenass caccens 4. T. pseudoachillea Winkl.
Pubescence of long simple hairs extensive. Arctic plants. (sect.
Omalates:) sinrwnnwnvninsirnnoars 1. T. bipinnatum (L.) Sch. Bip.
Pubescence lax or dense but always of shorter bifid (often mixed
with simple) hairs. Plants of steppe and semidesert zone. (sect.
NAPARG SIG ssa: ) EL. KOLAR ha NA REE A ARON 8.
Capitula lacking ligulate florets or with indistinctly ligulate (often
bilabiate) florets, slightly exceeding tubular florets; ligules, if present,

not;more than:1.0:(1.5) mm dAON®. siissececceccnevsess greet Rihncens 9.
Capitula always with ligulate florets, distinctly longer than tubular
florets, ligules more than (1.5)2.0 mm ong ...........cscssseseeseeees 24.

Stem strongly branched only near more or less woody base,
caespitose, very sparsely leafy, more or less virgate; capitula solitary
(but numerous on a single plant), 5-9 mm in dia, at apices. Southern
a rime a GIRO. I nee RA AK. Ua hin tae cies 10.

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306

10.

14.

18.

Stem herbaceous, usually branched above base; capitula few (some-
times numerous), very rarely (in very weak specimens) capitula
Solitary) OM SLEMS»:s55.sbeid act pithiss zstesadetans aphatereatneclelin Bieta Li;
Outer involucral bracts lanceolate-ovate, subobtuse, with wide
membranous border .............. 30. T. tenuissimum (Trautv.) Grossh.
Outer involucral bracts lanceolate from ovate base, acuminate, almost
lacking «membranous» Borden yet «12st fesgede rch ous Rehcs OM vepccbewweawtsnnoneabees
{dhs A dese Me 10. T. tabrisianum (Boiss.) Sosn. and Takht.
Outer involucral bracts broadly ovate to lanceolate-ovate, with rather

wide membranous border (degenerating on drying) ................ 12.
Outer involucral bracts lanceolate, acute or subacute, with or without
narrow membranous, borders). 2.diccteh: sees A hee is deg: ada) oa 15.
Plants usually densely grayish-pubescent, from saline steppes and
semideserts; involucre 4—7 (8) mm in dia ............cccccceeeceeeeseees 13.
Green or dull green, more weakly pubescent plants, from rubbly
and stony steppes and slopes; involucre S—11 mm in dia ......... 14.

Involucre usually pubescent only near base; outer involucral bracts
broadly ovate; with wide membranous border .................::ccesseceenes
wa cdhaais sea nies Sees saeekeh tigaucy eine 26. T. achilleifolium (MB.) Sch. Bip.
Involucre pubescent at base and on dorsal side of bracts; outer
involucral bracts lanceolate-ovate, with narrower membranous border
Seididdeih ated ddtabitia nelttanineths Ate ssrsd cuss areata Mea ctebehies 17. T. santolina Winkl.
Capitula (2)5—15(20) on a single stem, in more or less regular and
relatively:dense,,corymbs. Crimear ess: ivvcicestestucsiectenatvaine keesteny
Sonal eaebaraset av Rat lara ON OM ack antag an 24. T. paczoskii (Zefir.) Tzvel.
Capitula solitary or 2-5 on a single stem, in depauperate, usually
irregular corymbs. Ulutau Mountains .............cccsesesescecesesseseeseesseeee

Capitula in regular and rather dense corymbs; peduncles up to 8—
LOGTS): mmo Gipsb he iE owanys socaiettandoas sacar deptenaane omer leans tae ihe 16.
Capitula in lax, usually irregular corymbs; peduncles 1-15 (20) cm
long; Kazakhstan: 'p02.,saasscscire thcaonyekan hth: acnidia dinad algryondaadtia egnaeans 21.
Receptacle rather strongly convex. Altai, Tarbagatai, Saur,
Dzhungaria, Alatauj. .dvjashcns asueuta donteptvncaid woucteabenyd aileenaaeuabeh inees IP
Receptacle weakly convex, sometimes almost flat. Transcaucasia,
Kopetda, .. ciccrsnsainennensnssnaterdtivaxes- cite ghee eect hin R odeantancipsmes dicen s0it 18.
Capitula (6)8—10(12) mm in dia, on strongly thickened peduncles
in very dense, almost capitate COFYMDS ..........csscssseseesserseeseesessessenssens
schscecgate.- timeenl cet tiene. mesial itis 13. T. crassipes (Stschegl.) Tzvel.
Capitula (5)6—-8(10) mm in dia, on relatively thin peduncles in less
dense Corymbs '.. xis. cauie 002s. Chscesks 12. T. tanacetoides (DC.) Tzvel.
Capitula (2)4—10(15), at stem apices, in very dense, almost capitate
COGAN OS a ccsesciersonteactiansitn On cvainidearere ea cea ee 11. T. canescens DC.

S22

20.

21.

ptm

23.

24.

25.

Capitula in depauperate corymbs, usually more numerous ......... 19.
Capitula 2.0-3.5 mm in dia, (10)20—60(80) on single stem; peduncles
relatively slender, covered with fine appressed tomentum .................
ot gece SNe A LEE SE 6. T. myriophyllum Willd.
Capitula 3.5—6.5 mm in dia, (1)5—15(45) on single stem; peduncles
thicker, extensively covered with lax tomentum .............::cs000 20.
Capitula 3.5-5.5 mm in dia, (5)10-35(45) on single stem. Talysh
ett 1 OS) Oe) A eee en 7. T. duderanum (Boiss.) Tzvel.
Capitula 5.0-6.5 mm in dia. (1)5—15(20) on single stem. Kopetdag
Freee sete enette ee ect Aetascl veakeiede czas 8. T. heterophyllum Boiss.
Involucre 4—7 mm in dia; outer involucral bracts usually with more
or less distinct membranous border. Plants of solonetzic steppes
APIA SERACBEEER rota cdek sc seks h ctecceced eat eceevenvevsite 17. T. santolina Winkl.
Involucre (5)7—10(12) mm in dia; outer involucral bracts usually
lacking membranous border. Plants of rubbly and stony slopes,
(i ah EP LL ae ET Sais CR eR Ome Bey en 8 22.
Leaves twice pinnately cut, usually grayish from dense pubescence
a Avan A st ber Sd TS ee ae 14. T. turlanicum (Pavl.) Tzvel.
Leaves pinnately cut, with more or less pinnately lobed segments,
usually green or dull green, weakly pubescent, more hirsute than in
PASSV IES SPECI ware ctlost siescs uo -c enue bad ai hdaerduaie ante edited iW Aahas otras 2:
Segments of leaves more approximate, with acuminate lobes;
peduncles of capitula usually longer, 2-20 cm long. Karakala
Biouatains tie. Fath Se. .axd 15. T. scopulorum (Krasch.) Tzvel.
Segments of leaves more distant, with acute lobes; peduncles of
capitula usually shorter, 1-8 cm long. Mugodzhary Mountains ...
Di Spee ar EAS AA ALLD et! BP MAIO Lah a8 8 eA 16. T. saxicolum (Krasch.) Tzvel.
Lamina broadly linear, uniformly pinnately lobed or crenate.
TAB CSIAI olicelaseterensereeeetateeseddatetotieuie 32. T. akinfievii (Alex.) Tzvel.
Leaves with pinnately cut or parted lamina ............:.scseseereeeeee 25.
Outer involucral bracts more or less lanceolate, acute or subacute,
with or without very narrow. membranous border .............:0006+ 26.
Outer involucral bracts ovate or lanceolate-ovate; obtuse or
subobtuse, with rather wide (but easily degenerating) membranous
Border se Vai ee RO &, BOE PR Oe tal se hcl OES ROE se cones 30.
Lamina pinnatisect, with undivided or partly irregularly pinnately
lobed (less often divided) segments. Caucasus ..........ccscccesceeseeeeees
BD cee RMS on Labvulthc seth Ranke aatensuarcanreatee 9. T. tamrutense (Sosn.) Sosn.

Lamina twice pinnately Cut) «sisi neciecats eR Gein, al.
Plants ‘sreen, weakly pubescent :.02.1 60s IE Spe sceseest 28.
Plants more or less grayish from dense pubescence ..............++ 29.

Lamina up to 3 cm wide; corona of achene 0.2—0.4 mm long .....
Se sah ut teh Ee 5 sen Lleuwihaiytnsnan ini ann 20. T. uralense (Krasch.) Tzvel.

323

308

29:

30.

Six

32,

335

36.

Lamina up to 2 cm wide, on the average less dissected and thicker;
corona of,,achenes0:4—0:7amm lone Wi ee eA ae
sudden Bkibaatagn cee 21. T. sclerophyllum (Krasch.) Tzvel.
Involucre (5)7—10(12) mm in dia; plants short, to 35 cm high .......
Ji Donte nivvrasd held aerate 19. T. kittaryanum (C.A.M.) Tzvel.
Involucre (8)10—12(15) mm in dia; plants taller, to 60 cm high........
cbecbics tubaeGecsealeehd SOD PRE ERS a tied ms 18. T. Karelinii Tzvel.
Capitula solitary, very rarely 2 on a single stem; involucre 8-12
mm in dia; corolla tube of tubular florets rather abruptly and strongly
broadened in upper half; achenes 2.6—-3.5 mm long. Transcaucasia
inlsel aad eee 31. T. uniflorum (F. and M.) Sch. Bip.
Capitula usually rather numerous, less often ( in weak specimens)
solitary; corolla tube of tubular florets gradually but slightly
broadened in upper half; achenes 1.5—2.8 mm long ................. a,
Leaves grayish from extensive pubescence; terminal segments and
lobes roundish at apex, obtuse, or with very short cusp. Transcaucasia

Lasso di sbdcnvadsdeeuab od buenierecbay RU cabh rauee open eee bs Le oe.
Leaves grayish or green; terminal segments more or less acuminate.
Other. regions. of the US SRevcseeotecd ete al ee ae 34.

Capitula solitary or 2—3 on a single stem, on rather long peduncles
(up to 6-10: cm); involucre 7=9 min dia... 08. ee
mhachhauedatay «.ntigad cme oats: 29. T. oligocephalum (DC.) Sch. Bip.
Capitula (3)5—10(15), very rarely (in very weak plants) solitary,
and then involucre less than 7 mm in dia ...........esceeseeeseeeneeees 33.
Capitula on up to 5—6 cm long peduncles, in rather dense corymbs
aobblvk, -athegilan betscaae Sees se tnebeed 27.T. chiliophyllum (F. and M.) Sch. Bip.
Capitula on up to 12 cm long peduncles, usually arcuately divergent
from stem axis, in quite lax but regular CoryMDS .............0.:ssseeeseeeeees
wes hb RR RS eR A REE 28. T. longipedunculatum (Sosn.) Tzvel.
Leaves green, weakly pubescent (to completely glabrous); involucre
5-9 mm in dia; ligulate florets with ligule 1-2 mm long. Crimea
a cecponadasb edt pebetecee tah: apa iaelia Mi eee d crt vce 24. T. paczoskii (Zefir.) Tzvel.
Leaves more or less grayish from rather dense pubescence ........ 35.
Involucre 4-7 mm in dia; ligules 1-2 mm long; lamina of basal
leaves linear, to 1.3(1.5) cm wide. Plants of solonetzic steppes and
solonetzesiaisnaxt ce tehey: 26. T. achilleifolium (MB.) Sch. Bip.
Involucre 7-12 mm in dia; ligules 1.8-3 mm long; lamina of basal
leaves oblong-linear, to 2 cm wide. Plants of rubbly and stony
steppes and slopes) sss.c:::ccc..sserorgget ayaa Feenstra 36.
Stem and leaves extensively pubescent with semi-erect hairs; capitula
in dense corymb on up to 2 (4) cm long peduncles .................
Wie ER AAD. aed: RA A 22. T. odessanum (Klok.) Tzvel.

324

325

309

+ Stem and leaves appressed (and relatively weakly) hairy; capitula
in more lax corymb on up to 5(8) cm long peduncles .................
Fe RMRTING TA Sec seh Mechs Seiveeh sack weed 23. T. millefolium (L.) Tzvel.

Section 1. Omalotes (DC.) Tzvel. comb. nova.—Omalanthus Less.
Syn. comp. (1832) 260, non Juss. 1824.—Omalotes DC. Prodr. VI
(1837) 83.—Plants 6-50 cm high, more or less villous, with short
nonflowering shoots and leafy stems, mainly in lower half. Capitula
solitary or more numerous, and then aggregated in corymb, with or
without ligulate florets. Involucre broadly cupuliform, 10-18 mm in
dia; outer involucral bracts two-thirds, less often a half, as long as
inner; receptacle strongly convex, slightly punctate-tuberculate, weakly
alveolate. Achenes 2.4-3.5 mm long, with 5-7 prominent longitudinal
veins and undivided, corona more or less irregularly toothed, 0.3-0.8
mm long.

Type of section: Tanacetum camphoratum Less.

Note. Of the three closely related species in this section, the most
“primitive” species on the basis of all characters, T. camphoratum
Less., with numerous large capitula lacking ligulate florets and ar-
ranged in dense corymbs, is found in California; the second species, T.
huronense Nutt., with very short ligulate florets, is distributed mainly
in Canada, and the third, relatively “recent” species, T. bipinnatum
(L.) Sch. Bip., with rather large ligulate florets and usually solitary
capitula, is a widely distributed arctic plant.

1. T. bipinnatum (L.) Sch. Bip. Tanacet. (1844) 48, non Druce,
1914.—T. kotzebuense (Bess.) Bess. in Bull. Soc. Nat. Mosc. IX (1836)
107; DC. Prodr. VI, 131.—Chrysanthemum bipinnatum L. Sp. pl.
(1753) 890.—Matricaria bipinnata (L.) Desr. in Lam. Encycl. meth.
III (1792) 736.—Pyrethrum bipinnatum (L.) Willd. Sp. pl. III (1803)
2160; DC. op. cit. 60; Ldb. Fl. Ross. II, 557; Kryl. Fl. Zap. Sib. XI,
2751.—P. velutinum Fisch. ex Trautv. in Tr. Peterb. Bot. Sada, VIII,
2 (1883) 457.—Artemisia kotzebuensis Bess. in Nouv. Mem. Soc. Nat.
Mosc. III (1834) 80.—Ic.: Gmel. Fl. sib. II (1749) t. 85, fig. 1; Rouy,
Ill. Pl. Eur. rar. XIV (1900) t. 339.—Exs.: GRF No. 3271.

Perennial. Plants 6-40 cm high, with creeping, more or less
branched rhizome, bearing short nonflowering shoots and stems. Stems
solitary or few, erect or ascending at base, leafy mainly below, simple
or with few lateral branches above. Leaves grayish-green or dull green
from more or less extensive pubescence of long flexuous simple hairs,
sometimes subglabrous, with inconspicuous punctate-glandular hairs
or lacking them; basal and lower cauline leaves 25-30 cm long, on
rather long petiole, with twice or thrice pinnately cut oblong to linear-

326

310

oblong lamina; terminal segments of leaves ovate to linear, acute; middle
and upper cauline leaves sessile or sub-sessile, like basal, but smaller
and less dissected. Capitula solitary, less often 2-4 on single stem, on
long, more or less apically thickened peduncles. Involucre 10-18 mm
in dia, 4-8 mm long, usually more or less lanate; involucral bracts
herbaceous, with wide, brown or blackish-brown, membranous border,
outer broadly lanceolate, inner 1.5—2 times as long, lanceolate-oblong.
Ligulate florets 15-30, yellow, corolla tube 2.0-2.5 mm long, and
ligules 3-7 mm long, oblong to almost rotund; corolla of tubular florets
2.5-3.0 mm long. Achenes 2.5-3.5 mm long and about 0.7-0.9 mm
wide, corona 0.3—0.8 mm long. Flowering July to August.

Sandy and gravelly riverbanks, stony slopes.—Arctic: Arctic
Europe, Arctic Siberia, Chukotka, Anadyr; European part: Karelia-
Lapland (northeastern part), Dvina-Pechora (northern part); Western
Siberia: Ob River Area (northern part); Eastern Siberia: Yenisei (north-
ern part), Lena-Kolyma ( in the south up to Vilyui River valley); Far
East: Okhotsk. General distribution: Alaska, Canada (northwestern part).
Described from Siberia. The Gmelin illustration cited above is the type.

Note. Very variable in pubescence and the nature of the leaf di-
vision, but apparently does not form any distinct ecogeographical races.
Distributed very sporadically within the limits of its range.

Section 2. Tanacetum.—Tanacetum sect. Eutanacetum DC. Prodr.
VI (1837) 128.—Gymnocline sect. Anactis C. Koch in Linnaea XXIV
(1851) 340.—Pyrethrum sect. Tanacetum (L.) Boiss. FI. or. III (1875)
351 p. p.—P. sect. Tanacetum subsect. Eutanacetum (DC.) Sosn. in
Tr. Tifl. Bot. Sada, XVII (1915) 51.—Chrysanthemum sect. Tanacetum
(L.) Hoffm. in Pflanzenfam. IV, 5 (1889) 278, p. p.—C. sect.
Eutanacetum (DC.) Kitam. Compos. Japon. II (1940) 341.—Plants
weakly pubescent (from short simple and bifid hairs), 30-150 cm high,
usually lacking short non-flowering shoots, with densely leafy stems.
Capitula (3)8-80(100), at apex of main stem in rather dense corymbs
lacking ligulate florets. Involucre cupuliform, 4-13 mm in dia; outer
involucral bracts a half to two-thirds as long as inner. Receptacle weakly
convex, punctate-tuberculate. Achenes 1.2—2.5 mm long, with 5-7 (less
often to 8) prominent longitudinal ribs and undivided corona 0.1—0.4
mm long, more or less irregularly toothed.

Type of section: Type of genus.

Note. A small section including, besides the four species described
below, apparently a few Mediterranean and Northwest Asia species,
e.g., the Iranian species T. budjnurdense (Rech. f.) Tzvel. comb. nova
(= Chrysanthemum budjnurdense Rech. f. in Oesterr. Bot. Zeitschr.
XLVII (1950) 231), which is close to T. abrotanifolium.

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311

Series 1. Vulgaria Tzvel.—Leaves pinnately parted or cut, with
pinnately lobed, less often pinnately parted segments; pubescence
mainly of simple hairs.

2. T. vulgare L. Sp. pl. (1753) 845; MB. FI. taur.-cauc. II, 289;
DC. Prodr. VI, 128; Ldb. Fl. Ross. II, 601; Turcz. Fl. baic.-dahur. II,
74, p. p.; O. and B. Fedtsch. Perech. Rast. Turk. IV, 205; Grossh.
Opred. Rast. Kavk. 462; Kryl. Fl. Zap. Sib. XI, 2756.—T. umbellatum
Gilib. Fl. lithuan. I (1781) 171.—T. vulgare var. genuinum Trautv. in
Bull. Soc. Nat. Mosc. XXXIX, 1 (1866) 144; B. Fedtsch. Rast. Turk.,
(1915) 738.—C. tanacetum Vis. Fl. Dalm. II (1847) 84; Schmalh. FI.
II, 70.—Pyrethrum vulgare (L.) Boiss. Fl. or. II (1875) 352; Sosn. in
Tr. Tifi. Bot. Sada, XVII, 53; Grossh. Fl. Kavk. IV, 130.—Ic.: Rchb.
Ic. Fl. Germ. XVI (1854) t. 996; Majevsky, FI. Sr. Ross. (1892) Fig.
46; Syreistsch. Ill. Fll. Mosk. Gub. III, 267; B. Fedtsch. and Fler, FI.
Evr. Ross. III, 976, Fig. 977; Monteverde, Bot. Alt., Plate 37, Fig. 5;
Bonnier, Fl. Compl. France, Suisse et Belg. V, t. 292; Hegi, Ill. Fl. VI,
2; t: 263, fig. 5 and Fig. 332; Perfilev, Fl. Ser. Kr. 353, Fig. 104;
Zemlinsky, Lekarstv. Rast. SSSR, ed. 2, 210; Mikhailovskaya, in FI.
Beloruss. V (1959) Plate 33.—Exs.: Fl. Stir. exs. No. 794; Fl. Pol.
exs. No. 535; Pl. Pol. exs. No. 275; Pl. Finl. exs. No. 977 and 1387;
Pl. Belg. exs. No. 197; GRF No. 974 and 974a.

Perennial. Plants 30-150 cm high, with creeping, more or less
branched rhizome. Stems solitary or rather numerous, erect, extensively
leafy, usually more or less branched only at apex. Leaves sparsely
hairy (to glabrous), with short simple hairs, sometimes modified into
bifid, with numerous punctate-glandular hairs; basal leaves withering
early; cauline leaves to 20 cm long and 8—10 cm wide, lower on rather
long petiole thickened at base, upper sessile or subsessile; lamina oblong
or oblong-ovate, pinnately parted or cut up to narrow-winged axis,
with pinnately lobed, less often pinnately parted segments; terminal
segments or lobes ovate to linear-lanceolate, usually up to 5 mm wide,
more or less toothed or entire, short acuminate. Capitula (5) 10—70(100),
in fairly dense, compound corymbs. Involucre 5—8 mm in dia, 4-6 mm
long, more or less hairy to subglabrous; involucral bracts herbaceous,
with rather narrow (but more or less broadening at apex), light colored
or brownish, membranous border, outer ovate-lanceolate, inner oblong-
lanceolate. Corolla of tubular florets 1.5—2.4 mm long, slightly smaller
in outer pistillate florets. Achenes 1.2—1.8 mm long and about 0.5 mm
wide, corona 0.2—0.4 mm long. Flowering July to October.

Meadows, scrubs, steppes, riverbanks, thinned forests, as a weed
of roadsides, along borders of fields, and old fields, up to middle
mountain zone.—Arctic: (rarely, as introduced plant); European part:

32

(All regions); Siberia: (All regions); Far East: (Mainly as introduced
plant); Caucasus: Ciscaucasia, Dagestan, Transcaucasia (rarely); So-
viet Central Asia: Aralo-Caspian Region, Lake Balkhash Region,
Dzhungaria-Tarbagatai, Tien Shan (central part) introduced into other
districts. General distribution: Western Europe, Turkey, China (north-
ern part), Mongolia, Korea, Japan (northern part), North America
(mainly as introduced plant), introduced into other countries. Described
from Western Europe. Type in London.

Note. Known already to Linnaeus, T. vulgare var. crispum Ldb.
(FI. Ross. II, 601), with crimped leaves, was developed under cultivation
and has been spread through botanical gardens, now and then becoming
a naturalized garden variety of this species.

Economic Importance. It is a medicinal plant, long used in folk
medicine. Its stems and capitula contain an insecticidal substance. A
detailed account of its medicinal importance can be found in S.E.
Zemlinsky’s book (pp. 209-211) cited above.

3. T. boreale Fisch. ex DC. Prodr. VI, (1837) 128; Ldb. Fl. Ross.
II, 602; O. and B. Fedtsch. Perech. Rast. Turk. IV, 205.—T. vulgare
auct. non L.: Turcz. Fl. baic.-dahur. II (1845) 74 p. p.—T. vulgare var.
boreale (Fisch. ex DC.) Trautv. and Mey. FI. ochot. (1856) 54; Kryl.
Fl. Zap. Sib. XI, 2757.—Chrysanthemum boreale (Fisch. ex DC.) B.
Fedtsch. Rast. Turk. (1915) 738, non Makino 1909.—C. vulgare var.
boreale (Fisch. ex DC.) Makino in Makino and Nemoto, FI. Jap. (1925)
43.—Ic.: Gmel. Fl. Sib. II (1749) t. 65, fig. 1.

Perennial. Plants 30-100 cm high, with creeping, more or less
branched rhizome; stems solitary or few, erect, usually branched only
at apex. Leaves like previous species, but on the average more hairy;
terminal lobes of pinnately parted segments usually more distant and
narrower, with longer cusp. Capitula (3)5—40(50), in dense, compound
corymbs. Involucre 7-13 mm in dia and 5-7 mm long, more or less
hairy to subglabrous; involucral bracts like previous species, but with
wider dark brown membranous border. Corolla of tubular florets 2—3
mm long, less often modified into ligulate in outer pistillate florets,
usually smaller. Achenes 1.5-2.0 mm long, about 0.6 mm wide, corona
0.1-0.3 mm long. Flowering July to September.

Meadows, banks of lakes and rivers, herb slopes, scrubs, by
roadsides (in southern regions in middle and upper mountain zones).—
Arctic: Arctic Europe(?), Arctic Siberia, Chukotka, Anadyr; Western
Siberia: Ob River Area (northern part), Altai; Eastern Siberia: Yenisei
(northern part), Lena-Kolyma, Angara-Sayans, Dauria; Far East: (All
regions); Soviet Central Asia: Dzhungaria-Tarbagatai, Tien Shan
(central part). General distribution: China (northern part), Mongolia,

329

313

Korea, Japan (northern part), Alaska, Canada. Described from garden
specimens, apparently coming from Far East. Type in Berlin; isotypes
(?) in Leningrad.

Note. This species is connected to T. vulgare by fairly numerous
intermediate forms and differs from it not so much in the degree of
division of the leaves (individuals of T. vulgare with strongly divided,
twice pinnatipartite leaves are found almost throughout its range) as in
larger and fewer capitula and involucral bracts with a broader, dark
brown membranous border. I have not seen completely authentic
specimens of 7. boreale from the European Part of the USSR, and the
available reports refer, apparently, to specimens of T. vulgare with
more divided leaves.

Series 2. Pseudoachillea Tzvel.—Leaves pinnately cut, with
pinnately parted or cut segments; hairs mainly bifid.

4. T. pseudoachillea Winkl. in Tr. Peterb. Bot. Sada, XI, 12 (1891)
373; O. and B. Fedtsch. Perech. Rast. Turk. IV, 205.—T. newesskyanum
Winkl. 1. c. 374; O. and B. Fedtsch. ibid. 205.—Chrysanthemum
pseudoachillea (Winkl.) B. Fedtsch. Rast. Turk. (1915) 738.—C.
newesskyanum (Winkl.) B. Fedtsch. ibid. 738.—Lepidolopsis
pseudoachillea (Winkl.) Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XIX (1959) 375.—Pyrethrum pseudoachillea (Winkl.)
Bornm. Pl. Turkest. No. 565.—Exs.: Bornmuller, Pl. Turkest. No. 565;
GRF No. 3274.

Perennial. Plants 30-120 cm high, with creeping, more or less
branched rhizome; stems solitary or few, erect, extensively leafy, usually
more or less branched only at apex. Leaves sparsely hairy (to
subglabrous) with short bifid hairs, partly simple, and very short
papilliform hairs, as also numerous but not always punctate-glandular
hairs; basal leaves withering early; cauline leaves to 20 cm long and
8 cm wide, lower on rather long petiole, upper sessile or subsessile;
their lamina oblong, pinnatisect to very narrow-winged axis, with
pinnately parted or cut segments; lobes shallow-lobate or irregularly
toothed acuminate. Capitula (10)15-—60(80), in rather dense, compound,
corymb. Involucre 4-6 mm in dia, 3.5-4 mm long, glabrous or
subglabrous. Involucre bracts coriaceous-herbaceous, outer lanceolate,
almost lacking membranous border, inner linear-oblong, with narrow,
appendiculately broadened at apex, light colored or brownish,
membranous border. Corolla of tubular florets 1.5—2.5 mm long, usually
smaller in few outer pistillate florets. Achenes 1.8-2.5 mm long and
about 0.6—0.8 mm wide, corona 0.2—0.4 mm long. Flowering June to
August.

314

Stony slopes, meadows, forest glades, juniper forests; up to upper
mountain zone.—Soviet Central Asia: Tien Shan (western part), Pamiro-
Alai (with the exception of Pamir). General distribution: Afghanistan(?).
Described from western Tien Shan (Chirchik River Valley). Lectotype
(Am Fusse d. Berges Tschimgan am d. Ufer d. Tschirschik, 20. VI-2.
VII, 1881, A. Regel) and isotype in Leningrad.

Note. Probably by mistakes, this species, extremely close to T.
vulgare, was included by P.P. Poljakov (op. cit.) in the genus
Lepidolopsis Poljak. Tanacetum newesskyanum Winkl., which differs
from T. pseudoachillea by having broader leaf lobes and larger capitula,
apparently was described from a shade form of T. pseudoachillea.

Series 3. Abrotanifolia Tzvel.—Leaves thrice pinnately cut, with
linear terminal lobes; hairs mainly bifid.

5. T. abrotanifolium (L.) Druce in Rep. Bot. Exch. Cl. Brit. Isl.
1913, III (1914) 425.—T. setaceum Tausch in Flora, XII (1829) 11,
nom. nov. (for Achillea myriophylla Willd.).—T. millefoliatum F. and
M. in DC. Prodr. VI (1837) 128; Ldb. Fl. Ross. II, 602; Manden. in
Fl. Gruzii, VII, 373.—T. szowitsii (C. Koch) Sosn. and-Takht. in
Takht. and Fler. Fl. Erevana (1945) 311; Grossh. Opred. Rast. Kavk.
462.—Achillea abrotanifolia L. Sp. pl. (1753) 897.—A. myriophylla
Willd. Enum. hort. Berol. II (1809) 916, non Tanacetum myriophyllum
Willd.—Gymnocline szovitsii C. Koch in Linnaea, XXIV (1851) 340.—
Pyrethrum polyphyllum Boiss. Fl. or. II (1875) 351, nom. nov. (for
Tanacetum millefoliatum F. and M.); Sosn. in Tr. Tifl. Bot. Sada,
XVII (1915) 51.—P. szowitsii (C. Koch) Grossh. and Schischk. Pl. or
exs. (1924) 13; Sosn. in Zhurn. Russk. Bot. Obshch. XIV (1929) 84;
Grossh. Fl. Kavk. IV, 130, non Boiss. 1849.—Chrysanthemum szowitsii
(C. Koch) Bordz. in Zap. Kievsk. Obshch. Estestv. XXV (1915) 123.—
Exs.: Pl. or exs. No. 45.

Perennial. Plants 30-100 cm high, with creeping, more or less
branched rhizome; stems solitary or few, erect, extensively leafy, usually
more or less branched at apex. Leaves sparsely covered with short
bifid hairs, partly simple, with inconspicuous punctate-glandular hairs;
basal leaves usually withering early; cauline leaves up to 20 cm long
and 6 cm wide, lower on rather long petiole thickened at base, upper
sessile or sub-sessile; their lamina oblong or linear-oblong, thrice
pinnately cut up to wingless or almost wingless axis; terminal segments
narrow-linear and linear-lanceolate, acute; uppermost leaves twice
pinnately cut, smaller. Capitula 15-80, in rather dense, compound
corymbs. Involucre 4—5 mm in dia and 3.5-4.5 mm long, glabrous or

330 subglabrous; involucral bracts coriaceous-herbaceous, outer lanceolate,

315

almost lacking membranous border, inner linear-oblong, with narrow
but apically broader, light-colored or brownish membranous border.
Corolla of tubular florets 2.4-3.0 mm long, usually smaller in few
outer pistillate florets. Achenes 1.8-2.4 mm long and about 0.5—0.8
mm wide, corona 0.3-0.4 mm long. Flowering June to August.

Stony slopes, scrubs, riverbanks, gravel beds; up to upper mountain
zone.—Caucasus: Southern Transcaucasia. General distribution:
Armenia and Kurdistan. Described from garden specimens, apparently
acquired from Turkey. Type in London.

Note. The species has an extremely complex synonym in which
the oldest names (Achillea abrotanifolia L. and Tanacetum setaceum
Tausch.) were not even mentioned in the later botanical literature,
beginning with “Flora Orientalis” of E. Boissier.

Section 3. Xanthoglossa (DC.) Sch. Bip. Tanacet. (1844) 48.—T.
sect. Xanthogymnocline Sch. Bip. ibid. 47.—Pyrethrum *chrysanthema
MB. FI. taur.-cauc. II (1808) 326, nom. nud.—P. sect. Xanthoglossa
DC. Prodr. VI (1837) 60 p. p.—P. sect. Gymnoclines DC. ibid. 58 p.
p.—P. sect. Chrysoglossa C. Koch in Linnaea, XXIV (1851) 332.—P.
sect. Gymnocline subsect. Xanthogymnocline (Sch. Bip.) Sosn. in Tr.
Tifl. Bot. Sada, XVII (1915) 46.—Gymnocline Cass. in Dict. sc. nat.
XX (1821) 119 p. p.—G. sect. Chrysactis C. Koch in Linnaea, XXIV
(1851) 339.—Chrysanthemum sect. Gymnoclinia Benth. and Hook. Gen.
pl. II (1873) 427; Hoffm. in Pflanzenfam. IV, 5 (1894) 277.—C. sect.
Gymnoclines (DC.) Ling in Contr. Inst. Bot. Nat. Acad. Peiping, III
(1935) 481.—More of less hairy (mainly long or short bifid hairs,
often mixed with simple) plants 10-60 cm high, always with short (as
rosulla of basal leaves) nonflowering shoots, and more sparsely leafy
stems. Capitula (2)4—40(80), in dense or lax, usually irregular corymb,
less often solitary, with or lacking ligulate florets. Involucre cupuliform,
2.5-15.0 mm in dia, outer involucral bracts two-fifths to two-thirds as
long as inner. Receptacle weakly to strongly convex, punctate-
tuberculate. Achenes 1.5-3.5 mm long, with 5-12 prominent
longitudinal ribs and 0.1-0.7 mm long corona: corona undivided but
sometimes irregularly lobed almost to base.

Lectotype of section: T. millefolium (L.) Tzvel.

Note. The section with the largest number of species, particularly
widely distributed in countries of Northwest Asia.

Series 1. Myriophylla Sosn. in Izv. Kavk. Muz. X (1916) 17.—
Capitula lacking true ligulate florets, usually numerous in regular and
rather dense, corymb. Involucre 2.5-6.5 mm in dia; outer involucral

331 bracts lanceolate or ovate-lanceolate, lacking or almost lacking

316

membranous border; capitulum weakly convex. Herbaceous plants with
twice pinnately cut leaves.

6. T. myriophyllum Willd. Achill. (1789) 50 and Sp. pl. III (1803)
1814.—Achillea bipinnata L. Sp. pl. (1763) 1265, non Tanacetum
bipinnatum Sch. Bip. 1844.—Pyrethrum myriophyllum_ var.
subvirescens DC. Prodr. VI (1837) 59; Ldb. Fl. Ross. II, 555.—P.
myriophyllum auct. non C.A.M.; Boiss. Fl. or III, 350 p. p.; Grossh.
Fl. Kavk. IV, 131 and Opred. Rast. Kavk. 459 p. p.; Manden. in FI.
Gruzii, VIII, 371.—P. szowitsii Boiss. Diagn. ser. 1, XI (1849) 25.—
Gymnocline argyrophylla C. Koch in Linnaea, XXIV (1851) 340.—
Chrysanthemum myriophyllum (Willd.) Nabel. in Publ. Fac. Sc. Univ.
Masaryk. Brno, No. 52 (1925) 23.—Ic.: Takht. and Fedor. Fl. Erevana,
Atlas (1946) Plate 110.

Perennial. Plants 20-60 cm high, with thick, more or less branched
rhizome, more or less grayish or dull-green from rather dense appressed
pubescence of bifid hairs mixed with simple. Stems usually rather
numerous, loosely caespitose with short nonflowering shoots, less often
solitary, erect, relatively densely leafy, more or less branched only at
base and apex. Young leaves densely pubescent, later grayish or dull-
green from more or less dense pubescence, with numerous punctate-
glandular hairs; basal leaves up to 20(25) cm long, 3(4) cm wide, on
rather long (but shorter than lamina) petiole; their lamina oblong or
oblong-linear, twice pinnately cut; terminal segments broadly obovate
to oblong, up to 1—-1.5 mm wide, in turn 2—5-lobed, roundish at apex,
but with very short cusp; cauline leaves 3-10, like basal, but smaller,
upper sessile, lower short-petiolate. Capitula (10)20—60(80), in rather
dense, compound, corymb, on more slender (0.5)1.0—8.0(10.0) mm
long peduncles, more or less covered with appressed fine hairs.
Involucre 2.5-3.5 mm in dia, 3.5—4.5 mm long, at base densely
appressed hairy; involucral bracts coriaceous-herbaceous, outer
lanceolate, almost lacking membranous border, inner 1.5—2 times as
long, oblong or linear-oblong, with narrow but apically broader, light-
colored, membranous border; outer pistiliate florets between tubular
and ligulate, sometimes with up to 1.5 mm long deeply 3-parted ligule;
corolla of tubular disk florets 1.8-2.5 mm long. Achenes 1.5—2.0 mm
long and about 0.6 mm wide, with 5—7 longitudinal ribs; corona 0.1-—
0.3 mm long, irregularly lobed up to middle. Flowering June to August.

Stony and rubbly slopes, rocks; up to 2,500 m.—Caucasus: Eastern
Transcaucasia (western and southern parts), Southern Transcaucasia,
Talysh. General distribution: Armenia-Kurdistan. Described from
“Orient,” probably from northeastern Turkey. Type in London.

332

317

Note. “Tanacetum myriophyllum Willd.” is actually a new name
given to Achillea bipinnata L. without any basis. In the botanical lit-
erature on the Caucasus, this species has usually been confused with
Pyrethrum myriophyllum C.A.M., described quite independently.
Apparently, the only specimen of T. myriophyllum at my disposal from
Borzhomi (on the rocks of the ravine between Kazak settlement of
Sakogavi and the village of Sakogavi, 14.VII.1918, V. Kozlovsky)
belongs to a unique ecogeographical race (tentatively named by us as
ssp. borshomicum Tzvel. ssp. nova); it is similar to that species on the
basis of the size of the capitula, but has abundant flocculent-tomentose
pubescence on the peduncles of the capitula and the involucre and also
on the average has longer outer involucral bracts.

7. T. duderanum (Boiss.) Tzvel. comb. nova.—Pyrethrum
myriophyllum C.A.M. Verzeichn. Pfl. Cauc. (1831) 74; DC. Prodr. VI,
59, excl. var.; Ldb. Fl. Ross. II, 554; Boiss. Fl. or. III, 350 p. p.;
Grossh. Fl. Kavk. IV, 131 and Opred. Rast. Kavk. 459, p. p., non
Tanacetum myriophyllum Willd. 1789.—P. duderanum Boiss. Diagn.
ser. 1, XI (1849) 26.—P. myriophyllum var. eriocephalum Boiss. FI.
or. III (1875) 350; Grossh. Fl. Kavk. [V, 132.—Gymnocline myriophylla
(C.A.M.) C. Koch in Linnaea, XXIV (1851) 339.—Chrysanthemum
myriophyllum (C.A.M.) B. Fedtsch. Rast. Turk. (1915) 737, quoad
nomen.—Exs.: Kotschy, Pl. Pers. bor. No. 266.

Perennial. Plants 20-50 cm high, with thick, more or less branched
rhizome, more or less grayish from rather dense, appressed bifid and
simple hairs; stems usually numerous, less often solitary, erect, branched
only at base and apex. Leaves like previous species. Capitula (5)10—
35(45), in dense, compound corymb, on few thick peduncles (1)3-
15(20) mm long and sparsely covered with abundant sparse pubescence.
Involucre 3.5—5.5 mm in dia 4-5 mm long, densely tomentose near
base; involucral bracts coriaceous-herbaceous, outer broadly lanceolate,
almost lacking membranous border, inner linear-oblong, with wider
light colored or brownish membranous border. Outer pistillate florets
and tubular disk florets like previous species; achenes 1.5-2.0 mm
long, about 0.6 mm wide, with 5—7 elongate ribs and 0.1-0.3 mm long,
more or less lobed corona. Flowering June to July.

Stony and rubbly slopes; from 1,000 to 2,500 m.—Caucasus:
Talysh (southern part). General distribution: Iran-Afghanistan (north-
western part). Described from northern Iran (Dudera Mountain). Type
in Geneva; isotype in Leningrad.

Note. It replaces the previous species in the extreme south of
Talysh and in the mountains of northern Iran (Bogrov-Dag and Elburz
ranges).

333

318

8. T. heterophyllum Boiss. Diagn. ser. 1, VI (1845) 90.—T.
kotschyi Boiss. in Kotschy. Pl. Pers. austr. No. 433, nom. nud.; Winkl.
in Tr. Peterb. Bot. Sada, XI, 2 (1889) 126, nom. nud.—Pyrethrum
myriophyllum var. variegatum Boiss. Fl. or. III (1875) 351; O. and B.
Fedtsch. Perech. Rast. Turk. IV (1911) 188.—Chrysanthemum
myriophyllum (C.A.M.) B. Fedtsch. Rast. Turk. (1915) 737, quoad
pl.—Lepidolopsis kotschyi (Boiss.) Poljak. in Fl. Turkm. VII (1960)
108.—Exs.: Kotschy, Pl. Pers. austr. No. 433 (isotypes).

Perennial. Plants 15-30 cm high, with thick, more or less branched
rhizome, grayish from densely appressed bifid and simple hairs. Stems
few or solitary, erect or ascending at base, densely leafy, more or less
branched only at base and apex. Leaves grayish from dense pubescence,
with numerous, inconspicuous punctate-glandular hairs; basal leaves
up to 8(10) cm long and 1.5 cm wide, on rather long petiole; their
lamina oblong-linear, twice pinnately cut; terminal segments obovate
to broadly linear, up to 1 mm wide, in turn often 2—5-lobed or pinnate,
roundish at apex, but often with very short cusp; cauline leaves 3-7,
like basal, but sessile except lowermost. Capitula (1)5—15(20) on single
stem, in dense, simple or compound corymb, on somewhat thick sparsely
hairy, 0.2-1.0 cm long peduncles. Involucre 5.0-6.5 mm in dia, 3.5-
5 mm long, densely appressed hairy at base; involucral bracts
coriaceous-herbaceous, outer ovate-lanceolate, with very narrow
membranous border, inner 1.5—2.5 times as long, oblong-linear, with
narrow but appendiculately broadened at apex, light-colored or
brownish, membranous border. Outer pistillate florets between tubular
and ligulate, sometimes with up to 1.5 mm long, 3-lobed ligules; corolla
of tubular disk florets 2.4—3.0 mm long. Achenes 1.8-2.2 mm long,
about 0.6 mm wide, with 5—8 longitudinal ribs and 0.1-0.3 mm long,
more or less irregularly toothed corona. Flowering June to August.

Stony slopes, steppes; at 1,500-2,500 m.—Soviet Central Asia:
mountainous Turkmenia. General distribution: Iran. Described from
Iran (near the city of Shiraz). Type in Geneva; isotype in Leningrad.

Series 2. Gymnoclinoidea Sosn. in Tr. Tifl. Bot. Sada, XVII (1915)
40.—Capitula with ligulate florets, usually rather numerous, in regular,
rather dense corymb. Involucre 3.5—5.5 mm in dia; outer involucral
bracts lanceolate, lacking or almost lacking membranous border.
Receptacle weakly convex. Herbaceous plants with pinnatisect leaves
(leaf segments undivided or irregularly lobed).

9. T. tamrutense (Sosn.) Sosn. in Takht. and Fed. Fl. Erev. (1945)
312.—Chrysanthemum tamrutense Sosn. in Vestn. Tifl. Bot. Sada,
XXVII (1913) 11.—Pyrethrum tamrutense (Sosn.) Sosn. in Tr. Tifl.

33

mn

319

Bot. Sada, XVII (1915) 41 and in Zhurn. Russk. Bot. Obshch. XIV,
84; Grossh. Opred. Rast. Kavk. 459.—P. woronowii Sosn. in Izv. Kavk.
Muz. X (1916) 6; Grossh. Fl. Kavk. II, 131 and Opred. Rast. Kavk.
459.—Ic.: Sosn. op. cit. (1913) Plate II.

Perennial. Plants 10-40 cm high, with thick, more or less branched
rhizome, grayish-green or dull green from rather dense pubescence of
appressed bifid (mixed with simple) hairs. Stems solitary or few, erect,
rather uniformly but sparsely leafy, more or less branched only at apex
(in inflorescence). Leaves grayish-green or dull green from rather dense
pubescence, with inconspicuous punctate-glandular hairs; basal leaves
up to 12(15) cm long, 2.5(3) cm wide, on long (sometimes exceeding
lamina), basally thickened petiole; their lamina oblong to oblong-linear,
pinnately cut, with 3-8 distant linear or linear-lanceolate segments on
each side, usually narrowed toward base, undivided or partly irregularly
pinnately parted or lobed, with short cartilaginous cusp; cauline leaves
smaller, except lower, sessile, pinnately cut, with undivided segments.
Capitula 5-25, in rather dense, regular corymb, on (2)4—12(15) mm
long peduncles. Involucre 3.5—5.5 mm in dia, 4—5.5 mm long, more or
less appressed hairy almost throughout; involucral bracts coriaceous-
herbaceous, outer fewer, lanceolate or narrow-lanceolate, gray, lacking
membranous border, inner oblong-linear, 1.5 times, less often 2 times
as long as outer, witl narrow, light colored, membranous border,
terminating into small uniformly toothed appendage at apex of bract.
Outer pistillate florets ligulate, 4-8, bright yellow, corolla tube
1.2-1.8 mm long and ligules 3-5.5 mm long and 2.5-4 mm wide,
broadly ovate; corolla of tubular disk florets 2.4-3.0 mm long. Achenes
1.8-2.5 mm long, about 0.5 mm wide, with 5 longitudinal ribs and
undivided 0.2—0.3 mm long irregularly toothed corona. Flowering June
to July.

Stony and clayey, usually solonetzic slopes, rocks; in lower
mountain zone.—Caucasus: Eastern Transcaucasia (reported from the
village of Shordzha), Southern Transcaucasia. General distribution:
Turkey (northeastern part), Iran (northwestern part). Described from
northeastern Turkey (Olta region). Type in Tbilisi.

Note. The species is quite close to T. flavovirens (Boiss.) Tzvel.
comb. nova [= Pyrethrum flavovirens Boiss. Diagn. ser. 1, IV (1844)
10], described from Iran, whose isotype is preserved in the herbarium
of the Botanical Institute, Academy of Sciences of the USSR, differing
from it only by having copious pubescence on the involucral bracts
and usually more dissected leaves. Judging from the isotype of
Pyrethrum woronowii Sosn., in the herbarium of the Botanical Institute,
Academy of Sciences of the USSR, this species is not different from
C. tamrutense.

320

Series 3. Tabrisiana Tzvel.—Capitula lacking true ligulate flo-
rets, solitary. Involucre 5—9 mm in dia; outer involucral bracts ovate-
lanceolate, acuminate, lacking membranous border. Receptacle weakly
convex, almost flat. Plants more or less woody at base, with pinnately
cut leaves (segments pinnately parted or lobed).

10. T. tabrisianum (Boiss.) Sosn. and Takht. in Takht. and Fed.
Fl. Erevana (1945) 310; Grossh. Opred. Rast. Kavk. 462.—Pyrethrum
tabrisianum Boiss. FI. or. III (1875) 356; Grossh. Fl. Kavk. IV, 130.—
Chrysanthemum tabrisianum (Boiss.) Bornm. in Beih. Bot. Centralbl.
LX, 3 (1939) 194, in obs.

Perennial. Plants 10-30 cm high, with thick oblique rhizome, more
or less grayish-green from rather dense pubescence of appressed bifid
(mixed with simple) hairs. Stems usually numerous, caespitose, erect,
sparsely leafy, branched only at more or less woody base. Leaves more
or less grayish-green from rather dense pubescence, young leaves
densely hairy, with inconspicuous punctate-glandular hairs; basal leaves
up to 8 cm long and 1.3 cm wide, on rather long petiole, with pinnately
cut, linear or oblong-linear lamina; leaf segments 8-12 on each side,
more or less pinnately lobed or parted; terminal segments oblong or
obovate, to 1.5 mm wide, roundish at apex, usually with short cusp;
few cauline leaves strongly reduced, almost all short-petiolate, upper
pinnately parted or undivided. Capitula solitary ( but many on a single
plant) on up to 10-15 cm long peduncles. Involucre 5—9 mm in dia
and 6-8 mm long, more or less appressed hairy; involucral bracts
coriaceous-herbaceous, outer ovate-lanceolate, acuminate, lacking
membranous border, inner 1.5—2 times as long, oblong-linear, with
light colored membranous border only in upper part, modified at apex
into oblong, usually sub-acute appendage. Outer pistillate florets tubular,
partly modified into ligulate, few; corolla of tubular disk florets 2.2—
3.0 mm long. Achenes 2.8-3.6 mm long and about 0.6 mm wide, with
7-10 longitudinal ribs and 0.3-0.6 mm long undivided irregularly
toothed corona. Flowering June to August.

Stony slopes, rocks; at 1,000—2,000 m.—Caucasus: Reported from
Southern Transcaucasia. General distribution: Armenia and Kurdistan.
Described from Iran (vicinity of Tabriz). Type in Geneva; isotypes in
Leningrad.

Note. I have not seen specimens of this species from the USSR,
but maybe they are preserved in some herbaria of the Caucasus.

Series 4. Canescentia Tzvel.—Capitula lacking true ligulate flo-
rets, (2)4—10(15), at stem apex in very dense, almost capitate corymb.
Involucre 3.5—5.5 mm in dia; outer involucral bracts lanceolate, acute,

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lacking membranous border. Receptacle weakly convex, almost flat.
Herbaceous plants with pinnately cut leaves (leaf segments pinnately
parted or lobed).

11. T. canescens DC. Prodr. VI (1837) 129; Ldb. FI. Ross. II,’
602; Grossh. Opred. Rast. Kavk. 462.—T. incanum Fisch. and Mey.
ex DC. 1. c. in syn. non Willd. 1803.—Pyrethrum canescens (DC.)
Boiss. Fl. or. III (1875) 353; Sosn. in Tr. Tifl. Bot. Sada, XVII, 54;
Grossh. Fl. Kavk. IV, 130.—Chrysanthemum canescens (DC.) Rech. f.
in Oest. Bot. Zeitschr. XCVII (1950) 231, in obs.—Ic.: Takht. and
Fed. Fl. Erevana, Atlas (1946) Plate 109.

Perennial. Plants 15-50 cm high, with thick oblique rhizome, more
or less grayish-green from rather dense pubescence of appressed bifid
and simple hairs. Stems solitary or few, erect, sparsely leafy, more or
less branched only near very weakly woody base and in inflorescence.
Leaves grayish-green from rather dense pubescence, with numerous
punctate-glandular hairs; basal leaves up to 8 cm long and 1.2(1.5) cm
wide, on rather long petiole, with pinnately cut, oblong or oblong-
linear lamina; leaf segments 5-12 on each side, pinnately parted or
lobed; terminal segment broadly linear to obovate and even almost
semicircular, to 1.5 mm wide, apex roundish, often with short cusp;
cauline leaves reduced and less dissected, middle and upper usually
sessile. Capitula (2)4—10(15), compactly clustered at stem apex in very
dense, almost capitate corymb, on peduncles 1—5(8) mm long and finely
tomentose. Involucre 3.5—5.5 mm in dia, 4-6.5 mm long, poculiform,
basally more or less covered with appressed hairs; involucral bracts
coriaceous-herbaceous, outer lanceolate, acute, lacking membranous
border, inner 1.5—2 times as long as outer, broadly linear, with narrow
and light-colored membranous border in upper part, broadening at apex in
a small discontinuously fimbriate appendage along margin. Outer pistillate
florets tubular, few; corolla of tubular disk florets 2.4-3.0 mm long.
Achenes 2.2—3.0 mm long, about 0.6 mm wide, with 10-15 longitudinal
ribs (of which 3-6 ribs usually more prominent), corona 0.1-0.2 mm
long, irregularly lobed almost to base. Flowering June to July.

Rubbly, stony and clayey slopes; up to lower mountain zone.—
Caucasus: Southern Transcaucasia. General distribution: Iran (north-
western part). Described from Nakhichevan ASSR. Type in Geneva;
isotypes in Leningrad.

Note. Very variable in the size and pubescence of the capitula and
the form of the leaves. The only specimen of this species from Armenia
(vicinity of Yerevan near the village of Dzhirmich, 35.V.1922, A.
Grossheim) in the Herbarium of the Botanical Institute, Academy of
Sciences of the USSR has leaves with fewer segments, which are ir-

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regularly toothed or lobed along the margin and more or less acute at
the apex, and it is possible to treat it as a separate ecogeographical
race, which we tentatively call spp. erivanense Tzvel. ssp. nova. T.
canescens vat. parvifolium DC. (Prodr. VI, 129) from the vicinity of
the city of Nakhichevan, with shorter leaf lobes, apparently is of no
taxonomic importance.

Series 5. Songarica Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, IX (1946) 166.—Capitula (1)3—10(18), lacking true ligulate
florets, in dense and regular corymb. Involucre (5)6—10(12) mm in
dia; outer involucral bracts lanceolate, subacute, lacking membranous
border. Receptacle weakly convex. Herbaceous plants with twice
pinnately cut leaves.

12. T. tanacetoides (DC.) Tzvel. comb. nova.—T. meyerianum
Sch. Bip. Tanacet. (1844) 47.—Pyrethrum millefoliatum auct. non
Willd.: Ldb. Fl. Alt. TV (1833) 120.—P. tanacetoides DC. Prodr. VI
(1837) 59; Ldb. Fl. Ross. II, 555; O. and B. Fedtsch. Perech. Rast.
Turk. IV, 188; Kryl. Fl. Zap. Sib. XI, 2753.—P. millefoliatum var.
tanacetoides (DC.) Trautv. in Bull. Soc. Nat. Mosc. XX XIX, 2 (1866)
347.—Chrysanthemum millefoliatum var. tanacetoides (DC.) Kryl. Fl.
Alt. III (1904) 622.—C. tanacetoides (DC.) B. Fedtsch. Rast. turk.
(1915) 737.—Ic.: Ldb. Icon. pl. fl. Ross. impr. Alt. IV (1833) t. 369.

Perennial. Plants 15-60 cm high, with short creeping, more or less
branched rhizome, usually grayish-green from rather dense pubescence
of appressed bifid and simple hairs. Stems solitary or few, erect,
sparsely leafy, more or less branched only at apex. Leaves grayish-
green, less often dull green, from rather dense pubescence, with
inconspicuous punctate-glandular hairs; basal leaves up to 15-20 cm
long and 3 cm wide, on rather long petiole, with twice pinnately cut,
oblong to oblong-linear lamina; terminal segments linear to ovate, up
to 1.5 mm wide, subacute; cauline leaves smaller, except lower, sessile.
Capitula (1)3-10(15), at stem apex, in dense, regular corymb on
somewhat thick, finely tomentose peduncles up to 1.5—2 cm long.
Involucre (5)6—8(10) mm in dia and 4-5 mm long, more or less
appressed-hairy; involucral bracts coriaceous-herbaceous, outer
lanceolate, lacking membranous border, inner oblong-linear, 1.5—2 times
as long, with narrow but apically broadened, light colored, membranous
border. Outer pistillate florets more or less transitional between tubular
and ligulate, sometimes with ligules up to 1.5 mm long; corolla of
tubular disk florets 2-2.5 mm long. Achenes 2.0—2.5 mm long, about
0.6—-0.8 mm wide, with 6-8 longitudinal ribs corona 0.1-0.3 mm long
more of less irregularly toothed. Flowering July to August.

323

Stony slopes, steppes; up to middle mountain zone.—Western
Siberia: Altai; Soviet Central Asia: Dzhungaria-Tarbagatai. General
distribution: China (Dzhungaria). Described from southern Altai. Type
in Geneva; isotypes in Leningrad.

13. T. crassipes (Stschegl.) Tzvel. comb. nova.—Pyrethrum
crassipes Stschegl. in Bull. Soc. Nat. Mosc. XXVII (1854) 172; O.
and B. Fedtsch. Perech. Rast. Turk. IV (1911) 189.—Chrysanthemum
crassipes (Stschegl.) B. Fedtsch. Rast. Turk. (1915) 737.

Perennial. Plants 25-60 cm high, with short creeping, more or less
branched rhizome, dull green or grayish-green from rather dense
pubescence of appressed bifid (mixed with simple) hairs. Stems solitary
or few, erect, sparsely leafy, branched only at apex (in inflorescence).
Leaves like previous species, but usually less densely hairy. Capitula
(2)3—8(15), at stem apex, in very dense, almost capitate corymb, on
strongly thickened, finely tome.tose, peduncles up to 1(1.5) cm long.
Involucre (6)8—10(12) mm in dia and 4.0-5.5 mm long, more or less
appressed hairy; involucral bracts like previous species. Outer pistillate
florets tubular, sometimes ligulate; corolla of tubular florets 2.0-2.6
mm long. Achenes 2.0-2.5 mm long, about 0.6-0.8 mm wide with 6—
8 longitudinal ribs, corona 0.1-0.3 mm long, more or less irregularly
toothed. Flowering July to August.

Stony slopes, steppes, forest glades; in middle mountain zone.—
Western Siberia: Altai (southern part); Soviet Central Asia: Dzhungaria-
Tarbagatai. General distribution: China (Dzhungaria). Described from
Altai (Narym Range). Type in Moscow(?); isotypes in Leningrad.

Note. In relation to the previous species, it apparently is a higher-
mountain ecogeographical race, connected by intermediate forms.

Series 6. Santolina Tzvel.—Capitula lacking true ligulate florets,
(1)2-10(15), in lax, not always regular, corymbose inflorescence.
Involucres 4-12 mm in dia; outer involucral bracts lanceolate or ovate-
lanceolate, with or lacking narrow membranous border. Receptacle
weakly convex. Herbaceous plants with twice pinnately cut leaves.

14. T. turlanicum (Pavl.) Tzvel. comb. nova.—?T. barclayanum
DC. Prodr. VI (1837) 128; Ldb. Fl. Ross. II, 602.—Pyrethrum
achilleifolium var. discoideum Kar. and Kir. in Bull. Soc. Nat. Mosc.
XV (1842) 382.—P. turlanicum Pavl. in Vestn. Akad. Nauk Kazakhsh.
SSR, No. 3 (1950) 39.—P. mindshelkense B. Fedtsch. in herb.—P.
kasakhstanicum ssp. karataviense Krasch. in herb.—Ic.: Pavl. op. cit.
38, Fig. 13.—Exs.: GRF No. 3269b (P. kasachstanicum Krasch.).

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Plate XIV.
1 — Tanacetum tenuissimum (Trautv.) Grossh., habit of plant, corolla of tubular
floret, and achene; 2 — T. akinfievii (Alexeenko) Tzvel., habit of plant, corolla
of ligulate floret, corolla of tubular floret, and achene.

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325

Perennial. Plants 15-40 cm high, with short creeping, more or less
branched rhizome, usually grayish-green from rather profuse pubes-
cence or appressed bifid and simple hairs. Stems solitary or rather
numerous, erect or ascending at base, sparsely leafy, usually branched
above middle, less often almost from base, sometimes simple. Leaves
grayish-green or dull-green from rather profuse pubescence, with in-
conspicuous punctate glands; basal leaves up to 15—20 cm long and 3
cm wide, on rather long petiole, twice pinnately cut, oblong to oblong-
linear, terminal segments linear-lanceolate to ovate, up to 1.5 mm wide,
short-acuminate; cauline leaves smaller, lower short-petiolate, middle
and upper sessile. Capitula (1)2—5(10) on a single stem, in lax, not
always regular corymbose inflorescence, on peduncles more or less
finely tomentose, 1-10 cm long. Involucres 7-12 mm in dia, 1.5-5 mm
long, more or less appressed hairy; involucral bracts coriaceous-herba-
ceous, outer lanceolate and lanceolate-ovate, usually lacking membra-
nous border, inner oblong, 1.5—2.5 times as long as outer, with narrow,
lighter or brownish, membranous border. Peripheral pistillate florets
tubular but often more or less transitional to ligulate; corolla of tubu-
lar florets of disk 2.2-2.6 cm long and about 0.7 mm wide, with 5-8
longitudinal ribs, corona 0.2-0.3 mm long, more or less irregularly
toothed. Flowering July to August.

Rubbly and stony slopes, rocks; in lower and middle mountain
zones.—Western Siberia: Altai (southern part); Soviet Central Asia:
Lake Balkhash Region (sporadically), Dzhungaria-Tarbagatai, Tien Shan
(Karatau, Talass Alatau) and Ugam ranges. General distribution: China
(Dzhungaria). Described from Karatau. Type in Alma-Ata; isotype in
Moscow.

Note. According to the very brief original description, T.
barclayanum DC. (op. cit.) is very similar to this species. It was
described from cultivated specimens of unknown origin. Unfortunately,
the herbarium of the Botanical Institute of the Academy of Sciences of
the USSR does not have reliably identified specimens of this species.

The higher-mountain (and usually also smaller) specimens of T.
turianicum, with a relatively wider, brownish, membranous border on
the involucral bracts, possibly belong to a separate ecogeographical
race, named in the herbarium by B.A. Fedtschenko as “Pyrethrum
mindshelkense B. Fedtsch” (based on collections from Mynzhilga
Mountain in the Karatau, 15.VI.1908, No. 516).

15. T. scopulorum (Krasch.) Tzvel. comb. nova.—Pyrethrum
scopulorum Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
IX (1946) 164.—Lepidolopsis scopulorum (Krasch.) Poljak. in Bot.
Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 376.—Ic.: Krasch.
op. cit. 167, fig. 6.

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Perennial. Plants 20-50 cm high, with short creeping, more or less
branched rhizome, sometimes with appressed bifid and simple hairs.
Stems solitary or rather numerous, erect, sparsely leafy, usually
branched above middle, less often almost from base, sometimes simple.
Leaves green or dull green, usually pubescent to subglabrous, with
numerous punctate glands; basal leaves up to 8-10 cm long and 1.5 cm
wide, rather long-petiolate, pinnately cut, broadly linear, segments of
the first order pinnately lobed or parted; terminal lobe lanceolate-
ovate to oblong-lanceolate, up to 1.5 mm wide acuminate; cauline
leaves strongly reduced, middle and upper sessile, often pinnately parted
or lobed. Capitula (1)3—8(12) on single stem, in very lax, often irregular,
almost always simple corymbose inflorescence, on sparsely hairy, 2—
20 cm long peduncles. Involucres 7-12 mm in dia, 4.0-6.5 mm long,
more or less appressed hairy; involucral bracts coriaceous-herbaceous,
outer broadly lanceolate, almost lacking membranous border, inner
oblong, 1.5—2.5 times as long, with narrow, light colored, membranous
border. Peripheral pistillate florets tubular, but often more or less
transitional to ligulate; corolla of tubular florets of disk 2.2-2.6 mm
long. Achenes 2.0—2.5 mm long and about 0.6—0.8 mm wide, with 6—-
9 longitudinal ribs, corona more or less irregularly toothed along upper
edge, 0.2-0.3 mm long. Flowering June to August.

Granite rocks, stony slopes, in lower mountain zone.—Soviet
Central Asia: Lake Balkhash Region (Karkarala Mountains). Endemic.
Described from the Karkarala Mountains (Bektau-Ata Mountain). Type
and isotypes in Leningrad.

16. T. saxicolum (Krasch.) Tzvel. comb. nova.—Pyrethrum
kasakhstanicum ssp. saxicola Krasch. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, IX (1946) 162.

Perennial. Plants 20-35 cm high, with short, creeping, more or
less branched rhizome, sometimes with appressed bifid and simple
hairs. Stems solitary, or rather numerous, erect or ascending at base,
sparsely leafy, more or less branched above middle, sometimes simple.
Leaves green or dull green, usually sparsely hairy, with numerous
punctate glands. Basal leaves up to 10-12 cm long and 1.8 cm wide,
rather long-petiolate, pinnately cut, oblong-linear, their lobes more
distant than in previous species, pinnately lobed or irregularly toothed;
terminal lobe and teeth short-acuminate; cauline leaves strongly reduced,
upper and middle sessile, uppermost often pinnately lobed or even
entire. Capitula (1)2—6(10) on single stem, in lax, not always regularly
corymbose inflorescence, on peduncles sparsely hairy, 1-8 cm long.
Involucre 5-12 mm in dia, 4.0-5.5 mm long, more or less appressed
hairy; involucral bracts coriaceous-herbaceous, outer broadly lanceolate,

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with very narrow membranous border, inner oblong, 1.5—2.5 times as
long, with narrow, light-colored membranous border, more or less
broadened at apex. Peripheral pistillate florets tubular, but often more
or less transitional to ligulate; inner tubular florets of disk 2.2-2.6 mm
long. Achenes 2.0-2.5 mm long, about 0.7 mm wide, with 6-9
longitudinal ribs, corona 0.2-0.3 mm long, more or less irregularly
toothed. Flowering June to August.

Rubbly and stony slopes, rocks; up to lower mountain zone.—
Soviet Central Asia: Aralo-Caspian Region (Mugodzhary and much
higher parts of “melkosopochnik” [low, rounded, isolated hill]. Endemic.
Described from the Mugodzhary Mountains. Type (Aktyubinsk Region,
Mugodzhary Mountains, high ridge in region of Baktubai Mountains
near village of Berchogur, 26.VI.1927, No. 587, I. Krascheninnikov)
and isotypes in Leningrad.

Note. The most typical plants of this species are found in the
Mugodzhary Mountains, while the majority of the specimens from the
Kazakh “melkosopochnik” exhibit more or less intermediate features
between this and the other species of the series.

17. T. santolina Winkl. in Tr. Peterb. Bot. Sada, XI, 12 (1891)
375; O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 207.—
Chrysanthemum santolina (Winkl.) B. Fedtsch. Rast. Turk. (1915)
738.—Pyrethrum kasakhstanicum Krasch. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, IX (1946) 160.—Ic.: Krasch. op. cit. 161, fig. 3.—
Exs.: GRF No. 3269a (isotypes of P. kasakhstanicum Krasch.).

Perennial. Plants 10-35 cm high, with short creeping, more or less
branched rhizome, usually grayish from rather profuse pubescence of
appressed bifid and simple hairs. Stems rather numerous, less often
solitary, erect or ascending at base, strongly leafy, more or less branched
above, less often simple. Leaves usually grayish-green from rather
profuse pubescence, with numerous, though not always conspicuous,
punctate glands. Basal leaves up to 8(12) cm long and 1.0(1.5) cm
wide, rather long-petiolate, twice pinnately cut, linear or broadly linear;
terminal lobe ovate to oblong-linear, up to 1.0(1.5) mm wide, short-
acuminate; cauline leaves strongly reduced and less divided, middle
and upper sessile. Capitula (1)3—10(15) on a single stem, in lax, not
always regularly corymbose inflorescence, on more or less tomentose,
peduncles 1-8 cm long. Involucres 4—7 mm in dia; 3.0—-4.5 mm long,
at base and dorsally bracts more or less appressed hairy; involucral
bracts coriaceous-herbaceous, outer broadly lanceolate, with very
narrow membranous border, inner linear-oblong, 1.5 to 2.5 times as
long as outer, with relatively narrow, light colored membranous border,
more or less broadened at apex. Peripheral pistillate florets more or

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less transitional from tubular to ligulate, sometimes with 3 lobed limb
up to 1.0(1.5) mm long; corolla of tubular florets of disk 1.8-2.2 mm
long. Achenes 1.8-2.2 mm long and about 0.6 mm wide, with 6-9
longitudinal ribs, and corona more or less irregularly toothed or lobed
(sometimes almost to base) 0.2—-0.4 mm long. Flowering May to July.

Solonetz, solonetzic steppes, semideserts, mainly on clayey and
sandy soils.—European part: Lower Volga (southern part); Western
Siberia: Upper Tobol (southern part), Irtysh (southern part), Altai
(foothills; rarely); Soviet Central Asia: Aralo-Caspian Region, Lake
Balkhash Region, Kyzyl-Kum (northern part), Syr-Darya (northern part).
General distribution: China (western part). Described from Kzyl-Orda
Region of Kazakh SSR. Type in Leningrad.

Note. The species is described from specimens with the previous
year’s dry stems and capitula. In other respects it is entirely similar to
the type specimens of Pyrethrum kasakhstanicum Krasch.

Series 7. Kittaryana Tzvel——Capitula with more or less developed
ligulate florets, (1)2—6(10), in lax, not always regular corymbose
inflorescence. Involucre (5)7—12(15) mm wide; outer involucral bracts
lanceolate, lacking membranous border; receptacle moderately convex.
Herbaceous plants with twice pinnately cut leaves.

18. T. karelinii Tzvel. nom. nov.—T. saxatile (Kar. and Kir.)
Sch. Bip. Tanacet. (1844) 47, non Lodd. Bot. Cabin. II (1818) t. 126.—
Pyrethrum saxatile Kar. and Kir. in Bull. Soc. Nat. Mosc. XV (1842)
382; Ldb. Fl. Ross. II, 555; O. and B. Fedtsch. Perech. Rast. Turk. IV,
188; Kryl. Fl. Zap. Sib. XI, 2754.—Chrysanthemum saxatile (Kar. and
Kir.) B. Fedtsch. Rast. Turk. (1915) 737.

Perennial. Plants 20-60 cm high, with short creeping, more or less
branched rhizome, usually grayish-green from rather profuse pubescence
of appressed bifid and simple hairs. Stems few or solitary, erect,
sparsely leafy, more or less branched above middle. Leaves usually
grayish-green from profuse pubescence, with inconspicuous punctate
glands; basal leaves up to 15-20 cm long, 3 cm wide, rather long-
petiolate, twice pinnately cut, oblong to oblong-linear; terminal lobe
linear to ovate, up to 1.5 mm wide, short-acuminate; cauline leaves
much smaller and less divided, middle and upper sessile. Capitula
(1)3-6(10) on long (up to 8 cm) peduncles in lax, not always regular,
corymbose inflorescence. Involucres (8)10—12(15) mm in dia. 4.0-5.5
mm long, more or less appressed hairy; involucral bracts coriaceous-
herbaceous, outer lanceolate, lacking membranous border, inner oblong-
linear, 1.5 to 2 times as long as outer, with narrow, light-colored
membranous border, withering at apex. Peripheral pistillate florets

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ligulate, yellow, 10-25, with 1.2-1.6 mm-long corolla tube, limb
2.4—-3.0 mm long and 1.5—2.0 mm wide; inner tubular florets of disk
2.4-3.0 mm long. Achenes 2.4—3.0 mm long, about 0.6 mm wide, with
6-8 longitudinal ribs, and corona 0.3—0.5 mm long, usually irregularly
toothed or lobed along the edge. Flowering July to August.

Stony slopes, rocks from 2,000 m. up—Western Siberia: Altai
(southern part); Soviet Central Asia: Dzhungaria-Tarbagatai
(Dzhungarian Alatau). General distribution: Dzhungaria-Kashgaria.
Described from Dzhungarian. Alatau. Type or isotype(?) in Leningrad.

Note. It is very close to the following species but is a higher
mountain ecogeographical race.

19. T. kittaryanum (C.A.M.) Tzvel. comb. nova.—Pyrethrum
kittaryanum C.A.M. in Kleine Beitr. zur. Nahern Kennth. der FI. Russl.
(1830) 4; Stank. in Stank. and Tal. Opred. Rast. Evrop. Ch. SSSR
(1949) 632.—P. millefoliatum var. macrocephalum Ldb. FI. Ross. II
(1846) 557 p. p.—P. uralense var. tomentosum Krasch. in Bot. Mat.
Gerb. Bot. Inst. Akad. Nauk SSSR, IX (1946) 164.—Chrysanthemum
millefoliatum var. kittaryanum (C.A.M.) Krasch. in Fl. Yugo-Vost.
Evrop. Ch. SSSR, VI (1936) 346. —Exs.: GRF No. 3268a (under
Pyrethrum millefoliatum).

Perennial. Plant (6)10-40(50) cm high, with short creeping more
or less branched rhizome, usually grayish-green from rather profuse
pubescence of bifid and simple hairs. Stems rather numerous, less
often solitary, erect or basally ascending, sparsely leafy, usually more
or less branched above middle, less often simple. Leaves usually
grayish-green from rather profuse pubescence, with inconspicuous
punctate glands; basal and lower cauline leaves up to 10-15 cm long,
3 cm wide, rather long-petiolate, twice pinnately cut, oblong to oblong-
linear; terminal lobes linear to ovate, up to 1.5 mm wide, short
acuminate; cauline leaves much smaller and less divided, middle and
upper sessile. Capitula 1-6(8) on rather long (0.5—10 cm) peduncles,
in lax, not always regular, corymbose-inflorescence. Involucres (5)7-
10(12) mm in dia and 4.0-5.5 mm long, more or less appressed-hairy;
involucral bracts coriaceous-herbaceous, outer bracts lanceolate, lacking
membranous border (or almost without), inner oblong-linear, 1.5—2
times as long as outer, with narrow, light colored membranous border
broadening at apex. Peripheral pistillate florets ligulate, yellow, 15-
40, corolla tube 1.0-1.5 mm long, limb [ligule] 2.5-3.5 mm long and
1.5-2.0 mm wide; corolla of tubular florets of disk 1.8-2.5 mm long.
Achenes 22.6 mm ([sic.] long and about 0.6—0.7 mm wide, with 5—10
longitudinal ribs and more or less irregularly toothed or lobed, corona
0.2-0.3 mm long. Flowering June to August.

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Steppes, stony and rubbly slopes, up to lower mountain zone.—
European part: Volga-Kama (southeastern part), Trans- Volga, Lower
Volga (northern part); Western Siberia: Upper Tobol (southern part),
Irtysh (southern part), Altai (western foothills); Soviet Central Asia:
Aralo-Caspian Region (northern part), Lake Balkhash Region (northern
part). Endemic. Described from southern Urals. Type in Leningrad.

Note. The species is described from very small specimens with
solitary capitula on the stems. In my opinion, they do not constitute a
separate ecogeographical race; hence I combine them with the larger,
grayish-pubescent plants bearing several capitula on a stem (Pyrethrum
uralense var. tomentosum Krasch.).

20. T. uralense (Krasch.) Tzvel. comb. nova.—Pyrethrum
millefoliatum auct. non Willd.: Korsh. Tent. Fl. Ross. or. (1898) 214
p. p.—P. uralense Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, IX (1946) 162.—Chrysanthemum uralense Krasch. in Fl. Yugo-
Vost. Evrop. Ch. SSSR, VI (1936) 347, diagnosis in Russian.—Ic.:
Krascheninn, op. cit. (1946) 163, fig. 4.

Perennial. Plant (10)15—40(60) cm high, with short creeping, more
or less branched rhizome, sometimes pubescent, with bifid and simple
hairs (to subglabrous). Stems rather numerous, less often solitary, erect
or basally ascending, sparsely leafy, usually branched above middle,
less often simple. Leaves like previous species, but green or dull green,
sometimes hairy, often subglabrous (but younger leaves always hairy).
Capitula like previous species but involucres more weakly hairy.
Peripheral pistillate florets ligulate, yellow, 15—40, corolla tube 1.0—
1.5 mm long and limb 2.5-3.5 mm long, 1.5—2.0 mm wide; corolla of
tubular florets of disk 1.8—2.5 mm long. Achenes 2.0—2.6 mm long,
about 0.6 mm wide, with 5—10 longitudinal ribs and more or less
irregularly toothed, corona 0.2—0.3 mm long. Flowering June to August.

Stony and rubbly slopes, rocks up to lower mountain zone.—
European part: Volga-Kama (southeastern part), Trans- Volga, Lower-
Volga (northern part); Western Siberia: Upper-Tobol (southern part),
Irtysh (southern part); Soviet Central Asia: Aralo-Caspian Region
(northern part). Endemic. Described from southern Urals. Type in
Leningrad.

Note. Distinguished from the previous species only by the much
weaker pubescence, this species apparently is a morphologically very
weakly isolated ecogeographical race, confined to the more stony and
rocky habitats.

21. T. solerophyllum (Krasch.) Tzvel. comb. nova.—Pyrethrum
sclerophyllum Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,

348

331

IX (1946) 164.—Ic.: Krasch. ibid. 165, fig. 5—Exs.: GRF No. 3270
(isotype of species).

Perennial. Plant 20-35 cm high, with short, creeping, more or less
branched rhizome, more or less covered with bifid and simple hairs.
Stems few or solitary, erect, sparsely leafy, more or less branched
above middle. Leaves green or dull green, more or less sparsely hairy
to subglabrous (but younger leaves always profusely hairy), lacking
distinct punctate glands; basal leaves up to 10-15 cm long and 1.8—2.0
mm wide, rather long-petiolate, twice pinnately cut, oblong-linear or
broadly linear, hard and on the average less divided than in previous
species; terminal lobe linear-oblong to broadly ovate, to 1.5 mm wide,
short-acuminate; cauline leaves much smaller and less divided, middle
and upper sessile. Capitula (1)2—4(6), on rather long (up to 10 cm)
peduncles, in lax, not always regular, corymbose inflorescence.
Involucres 10-12 mm in dia, 4.5—5.0 mm long, more or less appressed-
hairy; involucral bracts coriaceous-herbaceous, outer lanceolate, almost
lacking membranous border, inner oblong-linear, with narrow, light
colored, membranous border, at apex broadened like appendage.
Peripheral pistillate florets ligulate, yellow, 20-40, corolla tube about
1 mm long and limb 3-4 mm long and 1.5-2.0 mm wide; corolla of
tubular florets of disk 2.2-3.0 mm long. Achenes 2.2—3.0 mm long,
about 0.6-0.7 mm wide with 6-10 longitudinal ribs, and corona 0.4—
0.7 mm long, more or less irregularly toothed along margin. Flowering
June to August.

Stony slopes (chalk and limestone outcrops).—European part:
Volga-Don (eastern part), Trans-Volga (western part), Lower Don
(northeastern part). Endemic. Described from vicinity of Khvalynsk on
Volga. Type in Leningrad.

Note. The most typical plants of this species are found only in the
vicinity of the city of Khvalynsk. Plants from other regions (for example,
from the Zhigulei District), often occupy a more or less intermediate
position between this and the previous species.

Series 8. Millefoliata Sosn. in Izv. Kavk. Muz. X (1916) 17.—
Capitula with more or less developed ligulate florets, sometimes
transitional to tubular, (1)2—15(20), in compact or lax, not always
regular corymbose inflorescence. Involucres (4)5—10(12) mm in dia;
outer involucral bracts lanceolate-ovate, with rather wide (but fragile)
membranous border. Receptacle strongly convex, often obtusely conical.
Herbaceous plants with bipinnatisect leaves.

22. T. odessanum (Klok.) Tzvel. comb. nova.—T. incanum auct.
non Willd.: Georgi, Beschreib Russ. Reich. VI (1800) 1231; Ldb. FI.

332

Ross. II, 604.—Pyrethrum millefoliatum var. macrocephalum Ldb. FI.
Ross. II, 557 p. p.—P. odessanum Klok. in Ind. Sem. Horti Bot.
Charkow. (1926) 7, and in Vizn. Rosl. URSR (1950) 540.

Perennial. Plants 20-40 cm high, with short, creeping, more or
less branched rhizome, grayish-green from profuse appressed and
squarrose bifid and simple hairs. Stems few or solitary, erect, sparsely
leafy, more or less branched at apex, densely hairy all over. Leaves
grayish from profuse pubescence (on prolonged storage of exsiccates,
hairs turn brownish), with numerous punctate glands; basal leaves up
to 15 cm long and 2 cm wide, rather long-petiolate, twice pinnately
cut, oblong-linear, their terminal lobes lanceolate-ovate to narrow-
lanceolate, up to 1.5 mm wide, short acuminate; middle and upper
leaves strongly reduced, sessile. Capitula 3-15, on relatively short
(0.5-4.0 cm long) somewhat thick peduncles in rather compact
corymbose inflorescence. Involucres 7-12 mm in dia and 4-5 mm
long, at base profusely lanate; involucral bracts coriaceous-herbaceous,
outer lanceolate-ovate, subacute, inner 1.5—2 times as long [as outer],
oblong; all bracts with light colored membranous border, much broader
in inner bracts and at apex broadened like appendage. Peripheral
pistillate florets ligulate, yellow, 15-40, with corolla tube 1.0-1.2 mm
long and limb 2.0—2.8 mm long and 1.5-—2.0 mm wide; corolla of
tubular florets of disk 1.8—2.5 mm long. Achenes 2.4—2.8 mm long and
0.6-0.7 mm wide, with 6-10 longitudinal ribs, and more or less
irregularly toothed corona 0.3—0.5 mm long. Flowering June to August.

Stony steppes, rock outcrops.—European part: Middle Dnieper
(southwestern part), Bessarabia, Black Sea Region (right bank of
Dnieper), Crimea(?). General distribution: Romania. Described from
vicinity of Odessa. Type in Kiev.

Note. The plants from Crimea occupy a more or less intermediate
position between this and the next species.

23. T. millefolium (L.) Tzvel. comb. nova. non F. and M. ex
Aitch. in Journ. Linn. Soc. XIX (1882) 169 in syn. sphalm. (=T.
millefoliatum).—T. tauricum Sch. Bip. Tanacet. (1844) 48, nom. nov.—
Anthemis millefolia L. Sp. pl. (1753) 896.—Chrysanthemum
millefoliatum L. Syst. veg. ed. 12 (1767) 563, nom. nov.; Schmalh. FI.
II, 69.—Matricaria millefoliata (L.) Desr. in Lam. Encycl. III (1792)
733.—Pyrethrum millefoliatum (L.) Willd. Sp. pl. III (1803) 2160;
MB. FI. taur.-cauc. II, 326; DC. Prodr. VI, 60; Boiss. Fl. or. III, 349
p. p.; O. and B. Fedtsch. Perech. Rast. Turk. IV, 189; Kryl. Fl. Zap.
Sib. XI, 2752.—P. millefoliatum var. microcephaium Ldb. FI. Ross. II
(1845-1846) 557.—?P. baumanii Stev. in Bull. Soc. Nat. Mosc. XXIX,

349 4 (1856) 383.—Ic.: Gmel. Fl. Sib. II (1749) t. 86, fig. 1; Krascheninn.

333

in Fl. Yugo-Vost. Evrop. Ch. SSSR, VI (1936) 346, fig. 681.—Exs.:
GRF Nos. 881 and 3268b.

Perennial. Plant (10)15—40(50) cm high, with short creeping, more
or less branched rhizome, usually grayish-green form rather profuse
pubescence of appressed bifid and simple hairs. Stems few or solitary,
erect or basally ascending, sparsely leafy, more or less branched above,
more or less appressed-hairy. Leaves usually grayish from rather profuse
pubescence, but on the average weakly pubescent in comparison with
previous species, sometimes dull green, with numerous punctate glands;
basal leaves up to 10-12 cm long, 2 cm wide, rather long-petiolate,
twice pinnately cut, oblong-linear, their terminal lobes lanceolate-ovate
to lanceolate-linear, up to 1.5 mm long, short-acuminate; middle and
upper cauline leaves strongly reduced, sessile. Capitula (2)3—10(15),
on peduncles 0.5—8.0 cm long, in lax, not always regular, corymbose-
inflorescence. Involucres 7-10 mm in dia and 4-5 mm long, at base
more or less lanate to subglabrous; involucral bracts coriaceous-
herbaceous, outer broadly ovate, subacute, inner oblong-lanceolate; all
bracts with rather wide, light-colored membranous border, but much
broader in inner bracts and at apex widened like appendage. Peripheral
pistillate florets ligulate, yellow, 15-30; corolla tube about 1 mm long
and limb 1.8-3.0 mm long and 1.5—2.0 mm wide; corolla of tubular
florets of disk 1.8—-2.5 mm long. Achenes 2.0—2.7 mm long and about
0.6-0.7 mm wide, with 5-8 longitudinal ribs, and more or less
irregularly toothed corona 0.3—0.6 mm long. Flowering June to August.

Steppes, stony and rubbly slopes, chalk and limestone outcrops.—
European part: Middle Dnieper (southern part), Volga-Don (southern
part), Trans-Volga (rarely in Kuibyshev Region), Bessarabia, Black
Sea Region, Lower Don, Lower Volga (northwestern part), Crimea;
Western Siberia: Upper Tobol (southern part, rarely); Caucasus:
Ciscaucasia, Dagestan (northern part); Soviet Central Asia: Aralo-
Caspian Region (northern part, rarely). Endemic. Described from garden
specimen of unknown origin, probably from the south of the European
part of the USSR. Probably the Gmelin drawing cited above is the type
specimen.

Note. Judging from the original description, Pyrethrum baumanii
Stev., described from Crimea (prope Ursuf. Baumann), belongs to this
species; I did not see its type.

24. T. paczoskii (Zefir.) Tzvel. comb. nova.—Pyrethrum
discoideum MB. in herb. non Ldb.—P. paczoskii Zefir. in Bot. Mat.
Gerb. Bot. Inst. Akad. Nauk SSSR, XVII (1957) 255.—P. achilleifolium
ssp. wisockianum Pacz. ex Zefir. |. c. in syn.

350

35

—

334

Perennial. Plant 15-40 cm high, with short, creeping, more or less
branched rhizome, with sparse bifid (mixed with simple) hairs. Stems
few or solitary, erect, sparsely leafy, more or less branched above,
short-appressed-hairy to subglabrous. Leaves green or dull green, more
or less sparsely hairy to completely glabrous (but younger leaves always
hairy), with numerous punctate glands; basal leaves up to 10-12 cm
long and 1.5(2.0) cm wide, rather long-petiolate, twice pinnately cut,
linear, their terminal lobes lanceolate-ovate to linear-lanceolate, up to
1.5 mm long, short-acuminate; middle and upper cauline leaves strongly
reduced, sessile. Capitula (2)5—15(20), on peduncles 0.5—5.0 cm long,
in lax, not always regular, corymbose inflorescence. Involucres 5-9
mm in dia and 4-5 mm long, usually subglabrous; involucral bracts
coriaceous-herbaceous, outer broadly ovate, subacute, inner oblong-
lanceolate, 1.5—2 times as long as outer; all bracts with rather wide,
light-colored, membranous border but wider in inner bracts and
appendiculately broadened at apex. Peripheral pistillate florets ligulate,
yellow, 20-30, but usually fewer, some transitional to tubular,
sometimes all florets tubular; corolla of ligulate florets (if present)
with tube about 1 mm long and ligule 1-2 mm long and 1.0-1.5 mm
wide; corolla of tubular florets of disk 1.8—2.4 mm long. Achenes 1.5-
2.0 mm long and about 0.6 mm wide, with 5-6 longitudinal ribs and
more or less irregularly toothed, corona about 0.3 mm long. Flowering
June to July.

Stony and rubbly slopes, steppes.—European part: Crimea (west-
ern part and northern foothills of Yaila). Endemic. Described from
Crimea. Type in Leningrad.

25. T. ulutavicum Tzvel. sp. nova in Addenda, XXVI, 878.

Perennial. Plant 12-25 cm high, with short, creeping, more or less
branched rhizome, usually dull green from rather sparse bifid (mixed
with simple) hairs. Stems rather numerous, less often solitary, erect or
basally ascending, sparsely leafy, simple or branched above middle.
Leaves dull green from not so dense pubescence, with inconspicuous
punctate glands; basal leaves up to 8 cm long and 1.3 cm wide, rather
long-petiolate, pinnatisect, broadly linear; primary segments 5—10 on
each side, usually pinnately lobed or parted, their terminal segments or
lobes broadly ovate to linear, up to 1.5 mm wide, short-acuminate;
cauline leaves strongly reduced, middle and upper sessile; upper leaves
usually pinnately parted. Capitula solitary or 2-5 on single stem and
then on long (up to 8-10 cm) peduncles, in lax, often irregular,
corymbose inflorescence. Involucres 6-11 mm in dia and 4.5-—6.0 mm
long, weakly hairy; involucral bracts coriaceous-herbaceous, outer
lanceolate-ovate, inner 1.5—2 times as long as outer, oblong-lanceolate;

352

335

all bracts with rather wide, light colored membranous border but in
inner bracts appendiculately broadened at apex. Peripheral pistillate
florets more or less transitional from tubular to ligulate, true ligulate
florets usually absent; corolla of tubular florets of disk 2-3 mm long.
Achenes 1.8—2.5 mm long and about 0.6 mm wide, with 6-9 longitudinal
ribs and irregularly toothed corona about 0.3 mm long. Flowering June
to July.

Rocks, stony slopes.—Soviet Central Asia: Aralo-Caspian Region
(Ulutau Mountains). Endemic. Described from the Ulutau Mountains.
Type (Karaganda Region, central part of Karsakpai District, slope of
the Kazan-Tau Mountain, 12 km south of Ulutavskii, 12. VII.1929, No.
298, N. Shipchinsky) in Leningrad.

Note. Despite some superficial similarity (first of all, the absence
of true ligulate florets) to species of series Santolia, the structure of
the involucres indicates very close affinity of this species with T.
achilleifolium and T. millefolium, in relation to which it is a narrowly
endemic race confined to rocky and stony habitats.

26. T. achilleifolium (MB.) Sch. Bip. Tanacet. (1844) 47.—
Pyrethrum achilleifolium MB. FI. taur.-cauc. II (1808) 327 and III
(1819) 580; DC. Prodr. VI, 58; Ldb. Fl. Ross. Ii, 554; Grossh. FI.
Kavk. IV, 135 and Opred. Rast. Kavk. (1949) 459; Kryl. Fl. Zap, Sib.
XI, 2752.—P. millefoliatum var. achilleifollum (MB.) O. and B.
Fedtsch. Perech. Rast. Turk. IV (1911) 189.—Gymnocline achilleifolia
(MB.) C. Koch in Linnaea, XXIV (1851) 340.—Chrysanthemum
achilleifolium (MB.) Kuntze in Tr. Peterb. Bot. Sada, X (1887) 202;
Krascheninn. in Fl. Yugo-Vost. Evrop. Ch. SSSR, VI (1936) 347.—C.
millefoliatum var. achilleifolium (MB.) Schmalh. Fl. Yugo-Zap. Ross.
(1886) 299 and FI. IT (1897) 70; Kryl. Fl. Alt. III (1904) 622.—Ic.:
Gmel. Fl. Sib. II (1749) t. 86, fig. 2; Krasch. op. cit. 247, Fig. 682.

Perennial. Plant (8)10—35(40) cm high, with short, creeping, more
or less branched rhizome, more or less grayish green from rather profuse
lax pubescence of appressed bifid (mixed with simple) hairs. Stems
rather numerous, less often solitary, erect or basally ascending, sparsely
leafy, more or less branched above middle. Leaves grayish-green from
rather profuse pubescence, with numerous punctate glands; basal leaves
up to 10(15) cm long and 1.3(1.5) cm wide, rather long-petiolate,
twice pinnately cut, linear, their terminal lobes lanceolate-ovate to
lanceolate-linear, up to 1 mm wide, long- or short-acuminate; cauline
leaves reduced, upper and middle sessile. Capitula (1)3—12(15), on
relatively long (0.5—8.0 cm) peduncles, in lax or somewhat lax, not
always regular, corymbose inflorescence. Involucres 4-7 mm in dia
and 3.0-4.5 mm long, sparsely hairy mainly at base; involucral bracts

336

coriaceous-herbaceous, outer lanceolate-ovate, subobtuse, inner 1:.5—2
times as long as outer, oblong-lanceolate; all bracts with rather wide,
light colored, membranous border, in inner bracts more or less
broadened at apex. Peripheral pistillate florets usually ligulate, yellow,
5-20, limb 1-2 mm long and 1-2 mm wide and tube about 0.8-1.0 mm
long, less often transitional from tubular to ligulate; corollas of tubular
florets of disk 1.5—2.2 mm long. Achenes 1.5—2.0 mm long and 0.3-
0.5 mm wide, with 5-8 longitudinal ribs and irregularly toothed, corona
0.3-0.5 mm long. Flowering June to July.

Solonetzic steppes, solonetzes.—European part: Trans-Volga
(southern part), Black Sea Region (southern part), Lower Don, Lower
Volga, Crimea (northern part); Western Siberia: Upper Tobol (south-
ern part), Irtysh (southern part; rarely), Altai (western foothills; rarely);
Caucasus: Ciscaucasia, Dagestan (northern part); Soviet Central Asia:
Aralo-Caspian Region (northern and eastern parts), Lake Balkhash
Region (northern part). General distribution: Romania. Described from
Ciscaucasia. Type in Leningrad.

Note. In relation to T. millefoliatum* it is a very close, more
ecological, than geographical race, confined more or less to “solo-
netzic” [alkaline], clayey or slightly stony parts of the steppe and
semideserts. Specimens intermediate between both these species are
not uncommon under transitional habitat conditions.

Series 9. Chiliophylla Tzvel.—Capitula with developed ligulate
florets, (1)2—10(12), in compact or lax regular corymbose inflorescence.
Involucres 4-9 mm in dia; outer involucral bracts ovate or ovate-
lanceolate, with rather wide, light colored or brownish, membranous
border. Receptacle strongly or moderately convex. Herbaceous plants
with twice pinnately cut leaves having shorter and more obtuse terminal
lobes than in species of previous series.

27. T. chiliophyllum (F. and M.) Sch. Bip. Tanacet. (1844) 47.—
T. kochii Sch. Bip. ibid.—Pyrethrum chiliophyllum F. and M. in DC.
Prodr. VI (1837) 59; Ldb. Fl. Ross. II, 555; Sosn. in Tr. Tifl. Bot.
Sada, XVII (1915) 48, descr. emend.; Grossh. Fl. Kavk. IV, 132 and
Opred. Rast. Kavk. 460.—P. armenum C. Koch in Linnaea. XVII (1843)
46.—P. millefoliatum auct. non Willd.: Boiss. Fl. or. III, 349 p. p.—
P. cheilanthifolium Sosn. in Izv. Kavk. Muz. X (1916) 11; Grossh. FI.
Kavk. IV, 132 and Opred. Rast. Kavk. 460.—P. chiliophyllum var.
oligocephalum (DC.) Sosn. op. cit. (1916) 14, quoad pl.; Grossh. FI.

*Should read T. millefolium.—Sci. Ed.

353

337

Kavk. IV, 132.—P. transcaucasicum Sosn. in Tr. Azerb. Otd. Zakavk.
Fil. Akad. Nauk SSSR, Bot. I (1933) 46; Grossh. Fl. Kavk. IV < 132
and Opred. Rast. Kavk. 459.—Gymnocline chiliophylla (F. and M.) C.
Koch in Linnaea, XXIV (1851) 339.—G. armena (C. Koch) C. Koch
op. cit. (1851) 340.—Chrysanthemum transcaucasicum (Sosn.) Bornm.
in Fedde, Repert. XXXVI (1934) 350.

Perennial. Plant (6)10—35(50) cm high, with short, creeping, more
or less branched rhizome, grayish from profuse fine tomentum of
appressed bifid and simple hairs. Stems few, less often solitary, erect
or basally ascending, sparsely leafy, more or less branched at apex.
Leaves grayish-green from profuse fine tomentum, with inconspicuous
punctate glands; basal leaves up to 10(15) cm long and 2.5(3.0) cm
wide, rather long-petiolate, twice pinnately cut, somewhat oblong to
broadly linear their second order segments often lobed or divided,
terminal lobes and segments almost sub-orbicular to oblong, up to 1.5
mm wide, obtuse but often with very short cusp; cauline leaves much
smaller and less divided, middle and upper sessile. Capitula (1)3-
10(12), on relatively short (0.4-5.0 cm) peduncles, in rather compact,
regularly corymbose inflorescence. Involucres 4.5—9.0 mm in dia and
3.5-5.0 mm long, more or less appressed-hairy; involucral bracts
coriaceous-herbaceous, outer ovate, sub-obtuse, inner oblong-linear,
1.5—2 times as long as outer, all bracts with wide, light-colored or
brownish, membranous border, in inner bracts appendiculately
broadened at apex. Peripheral pistillate florets ligulate, yellow, 10-25,
corolla tube 1.2—1.6 mm long and limb (1.5)2.0-4.0(5.0) mm long 1-
3 mm wide; corolla of tubular florets of disk 1.6—2.4 mm long. Achenes
1.5-2.0 mm long and about 0.5 mm wide, with 5—8 longitudinal veins
and irregularly toothed (or lobed), corona 0.1-0.3 mm long. Flowering
June to August.

Stony and rubbly slopes, steppes to middle mountain zone.—
Caucasus: Dagestan (southern part), Eastern and Southern
Transcaucasia, Talysh. General distribution: Turkey (northeastern part),
Iran-Afghanistan (northwestern part). Described from Southern
Transcaucasia (“Karabakh”). Type and isotypes in Leningrad.

Note. A highly polymorphic species, probably divisible into many
narrow endemic (local) races differentiated by the size of the capitula,
length of their peduncles, size of the limb [ligule] in the ligulate florets
(specimens with particularly large ligulate florets are found in southern
Dagestan and in the region of Lake Sevan), and so on. Nevertheless,
I consider it possible for the time being to retain as synonyms of this
species such later described species as Pyrethrum armenum C. Koch,
P. cheilanthifolium Sosn. and P. transcaucasicum Sosn. The first two
of these were described from northeastern Turkey near its border with

338

the Armenian SSR, and, in my opinion, do not differ significantly from
the type specimens of Tanacetum chiliophyllum. The third species was
described from very small specimens collected by M. Sakhokiya in the
Geokchai District of the Azerbaidzhan SSR, apparently in places
subjected to intense cattle grazing (there are also larger specimens of
T. chiliophyllum from this same area). Specimens with a brownish
membranous border of the involucral bracts are found about equally
often in the mountains of the Lesser Caucasus as in the foothills of the
Greater Caucasus, although it is not ruled out, all the same, that they
form a very weakly differentiated, higher-mountain race of the series
Chiliophylla.

28. T. longipedunculatum (Sosn.) Tzvel. comb. nova.—Pyrethrum
longipedunculatum Sosn. in Zam. po Sist. i. Geogr. Rast. Bot. Inst.
Akad. Nauk Gruz. SSR, XV (1949) 6; Grossh. Opred. Rast. Kavk.
459.

Perennial. Plant 25-50 cm high, with short, creeping, more or less
branched rhizome, grayish from profuse, appressed, bifid and simple
hairs. Stems few, less often solitary, erect, sparsely leafy, more or less
branched at apex. Leaves as in previous species. Capitula (3)5—10(12),
on rather long (1.5—12 cm), arcuate peduncles, in lax but regular
corymbose inflorescence. Involucres 4—5 mm in dia and 3-4 mm long,
at base profusely hairy; involucral bracts coriaceous-herbaceous, outer
ones ovate-lanceolate, with narrow membranous border, inner ones
oblong-linear, 1.5—2 times as long as outer, with wide, light colored,
membranous border, appendiculately broadened at apex. Peripheral
pistillate florets ligulate, yellow, 5-15, corolla tube 1.2-18.0 mm long
and limb 1.8—2.5 mm long and 1-1.5 mm wide; corolla of tubular florets
of disk 2—2.5 mm long. Achenes 1.5—2.2 mm long and about 0.5 mm
wide, with 5—7 longitudinal ribs and corona 0.1—0.2 mm long, obtusely
and irregularly lobed almost up to base. Flowering June to July.

Stony and rubbly slopes of lower mountain zone.—Caucasus:
Southern Transcaucasia (Nakhichevan ASSR). Endemic. Described from
Nakhichevan ASSR. Type in Tbilisi; isotypes in Leningrad.

29. T. oligocephalum (DC.) Sch. Bip. Tanacet. (1844) 48.—
Pyrethrum oligocephalum DC. Prodr. VII (1838) 297.—P. millefoliatum
var. oligocephalum (DC.) Boiss. Fl. or. III (1875) 349.—P.
chiliophyllum var. oligocephalum (DC.) Sosn. in Izv. Kavk. Muz. X
(1916) 14, quoad nom.—P. sosnowskyanum Grossh. Fl. Kavk. IV
(1934) 132, Diagnosis in Russian, and Opred. Rast. Kavk. (1949) 460.

Perennial. Plant 20-40 cm high, with short, creeping, more or less
branched rhizome, grayish-green from profuse pubescence of appressed

355

339

bifid and simple hairs. Stems few, less often solitary, erect, sparsely
leafy, simple or with few lateral branches in upper part. Leaves as in
previous species. Capitulum solitary or 2 or 3 on a single stem, on
rather long, up to 6(10) cm peduncles. Involucres 7-9 mm in dia and
5-6 mm long; involucral bracts as in previous species. Peripheral
pistillate florets ligulate, yellow, corolla tube 1.2-1.8 mm long and
limb 2—4 mm long; corolla of tubular florets of disk 1.5—2.4 mm long.
Achenes 1.5—2.5 mm long and about 0.6 mm wide, with 5-8 longitudinal
veins [ribs] and irregularly toothed, corona about 0.2 mm long.
Flowering June to July.

Stony slopes and rocks in middle mountain zone.—Caucasus:
Southern Transcaucasia (Daralgez). General distribution: Armenia and
Kurdistan. Described from Turkey. Type in Geneva.

Note. Despite the complete similarity of their original descrip-
tions, it is not ruled out that Pyrethrum oligocephalum DC. and P.
sosnowskyanum Sosn. are not entirely identical. I did not see the type
specimens of both these species.

Series 10. Dumosa Tzvel.—Capitula lacking true ligulate florets,
solitary (but numerous on a plant). Involucre 5.5—7.5 mm in dia; outer
involucral bracts lanceolate-ovate, with rather wide, light colored,
membranous border. Receptacle moderately convex. Plants more or
less woody at base, with pinnately cut leaves.

30. T. tenuissimum (Trautv.) Grossh. Opred. Rast. Kavk. (1949)
462.—Pyrethrum tenuissimum Trautv. in Tr. Peterb. Bot. Sada, IX, 1
(1884) 392; Grossh. Fl. Kavk. IV < 129.—P. dumosum auct. non Boiss.:
Lipsky in Tr. Peterb. Bot. Sada, XIII (1894) 306 and Fl. Kavk. (1899)
349; Sosn. in Tr. Tifl. Bot. Sada, XVII (1915) 42 p. p.—P. tabrisianum
auct. non Boiss.: Sosn. in Zhurn. Russk. Bot. Obshch. XIV (1929)
84.—Chrysanthemum tenuissimum (Trautv.) Winkl. in Tr. Peterb. Bot.
Sada, XI (1889) 123; B. Fedtsch. Rast. Turk. 737.

Perennial. Plant 15—40 cm high, with thick, many-branched rhizome,
grayish-green from rather profuse pubescence of bifid and simple hairs.
Stems numerous, branched only at woody base, erect or more or less
arcuately bent, very sparsely leafy (almost virgate). Leaves grayish
from rather profuse pubescence, with rather numerous punctate glands;
basal and lower cauline leaves up to 5(8) cm long and | cm wide, with
rather long petioles thickened at base, pinnately cut, broadly linear,
their primary segments 5—10 on each side, more or less pinnately lobed,
sometimes all or some entire, terminal segments and lobes up to 1.5
mm wide, obtuse but usually with very short cusp; middle and upper
cauline leaves very small, sessile, pinnately lobed to subentire. Capitula

356

340

solitary, but numerous on a plant, on long, slender peduncles. Involu-
cres 5.5—7.5 mm in dia and 4.5—5.5 mm long, more or less appressed-
hairy; involucral bracts coriaceous-herbaceous, outer ones lanceolate-
ovate, inner ones linearly oblong, 2—3 times as long as outer; all bracts
with wide, light colored, membranous border, in inner bracts
appendiculately broadened. Peripheral pistillate florets tubular or tran-
sitional from tubular to ligulate; corolla of tubular florets of disk 1.8-
2.5 mm long. Achenes 1.8—2.3 mm long and about 0.5 mm wide, with
6-10 longitudinal veins [ribs] and corona irregularly sharp toothed,
0.2—0.4 mm long. Flowering June to July (Plate XIV, Fig. 1).

Rocks, stony slopes in lower [mountain] zone.—Caucasus: Southern
Transcaucasia (southeastern part of Nakhichevan ASSR). General
distribution: Iran (northeastern part). Described from “Karabakh,”
probably from Nakhichevan ASSR. Type in Leningrad.

Note. It is distinguished from the very similar Iranian species T.
dumosum Boiss. (Diagn. ser. 1, 6, 1845, p. 89) by cup-shaped (but not
goblet-shaped) involucres with bracts that are quite profusely hairy on
the dorsal side and with a membranous border that is considerably
narrower at the apex.

Series 11. Uniflora Tzvel.—Capitula with developed ligulate
florets, solitary, less often two on single stem. Involucres 8-12 mm in
dia; outer involucral bracts lanceolate-ovate, with rather wide, light
colored, membranous border. Receptacle weakly or moderately convex.
Herbaceous plants with twice pinnately cut leaves.

31. T. uniflorum (F. and M.) Sch. Bip. Tanacet (1844) 48.—
Pyrethrum uniflorum (F. and M.) in DC. Prodr. VI (1837) 60; Boiss.
Fl. or. III, 349; Sosn: in Izv. Kavk. Muz. X, 11; Grossh. Fl. Kavk. IV,
132 and Opred. Rast. Kavk. 459.—P. dumosum auct. non Boiss.: Sosn.
in Tr. Tifl. Bot. Sada, XVII (1915) 42, p. p.

Perennial. Plant 15S—SO cm high, with thick, many-branched rhizome,
grayish-green form rather profuse pubescence of bifid and simple hairs.
Stems few or solitary, erect, sparsely leafy, simple. Leaves grayish-
green from rather profuse pubescence, with inconspicuous punctate
glands; basal and lower cauline leaves up to 10 cm long and 1.5 cm
wide, rather long-petiolate, twice pinnately cut, oblong-linear; terminal
lobe oblong to broadly-ovate, up to 1.5 mm wide, obtuse but with
short cusp; middle and upper cauline leaves strongly reduced and less
divided, sessile. Capitula solitary, less often 2 on stem, on long (up to
15 cm) peduncles. Involucres 8-12 mm in dia and 5.5—9.0 mm long,
basally somewhat cuneate or obtuse, more or less appressed-hairy;
involucral bracts coriaceous-herbaceous, outer ones lanceolate-ovate,

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341

inner ones linearly oblong, 2—3 times as long as outer; all bracts with
rather wide, light-colored, membranous border but at apex
appendiculately broadened in inner bracts. Peripheral pistillate florets
ligulate, yellow, corolla tube 1.8—2.5 mm long and limb 5-10 mm long
and 1.5—3.5 mm wide; tubular florets of disk 2.6—3.5 mm long. Achenes
2.6—3.5 mm long and about 0.6 mm wide, with 8—12 longitudinal ribs
and corona irregularly toothed or lobed, 0.2-0.4 mm long. Flowering
May to July.

Stony and rubbly slopes, rocks in lower mountain zone.—Caucasus:
Southern Transcaucasia (mainly in Nakhichevan ASSR). General dis-
tribution: Iran (northwestern part). Described from Iran. Type and
isotypes in Leningrad.

Note. Apparently, it is divisible into several narrowly endemic,
ecogeographical races differentiated mainly by the size of the capitula
and the length of the ligulate florets. In the USSR, plants with capitula
smaller than in the type specimens predominate, sometimes with two
on the same stem.

Series 12. Integrifolia Sosn. in Tr. Tifl. Bot. Sada, XVII (1915)
39.—Capitula with developed ligulate florets, solitary. Involucres 4.5—
7.0 mm in dia; outer involucral bracts lanceolate-ovate, with very
narrow, brownish, membranous border. Receptacle weakly convex.
Herbaceous plants with pinnately lobed leaves.

32. T. akinfievii (Alex.) Tzvel. comb. nova.—Pyrethrum akinfiewii
Alexenko in Tr. Tifl. Bot. Sada, VI, 1 (1902) 58; Sosn. in Tr. Tifl.
Bot. Sada, XVII, 39; Grossh. Fl. Kavk. IV, 131 and Opred. Rast.
Kavk. 459.

Perennial. Plant 6-30 cm high, with thick, many-branched rhi-
zome, silky-grayish from, fine appressed tomentum of bifid and simple
hairs. Stems usually rather numerous, less often solitary, erect or ba-
sally ascending, sparsely leafy, simple. Leaves silky-grayish from pro-
fuse pubescence, with inconspicuous punctate glands; basal leaves up
to 10 cm long and 8 mm wide, with rather long petiole thickened at
base, uniformly pinnately lobed, broadly linear, lobes 8-20 on each
side, entire, obtuse or subobtuse; cauline leaves strongly reduced,
basally cuneately narrowed, blunt-toothed to entire. Capitula solitary
(but rather numerous on our plant), with very long (up to 15 cm)
peduncles. Involucres 4.5-7.0 mm in dia and 6.0-8.5 mm long, more
or less appressed-hairy; involucral bracts coriaceous-herbaceous, outer
ones lanceolate-ovate, subacute, with very narrow brownish, membra-
nous border, inner ones oblong-linear, 1.5—2.5 times as long as outer,
with wider membranous border, apically appendiculately broadened.
Peripheral pistillate florets ligulate, orange-yellow, 8-20, with corolla

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342

tube about 1.5 mm long and limb 6-11 mm long and 2.5-4.5 mm
wide; corolla of tubular florets of disk 2.4—3.0 mm long. Achenes 2.4—
3.0 mm long and about 0.7—-0.8 mm wide, with 8-12 longitudinal ribs,
corona 0.1-0.2 mm long, irregularly blunt-toothed almost to base.
Flowering July to August (Plate XIV, Fig. 2).

Calcareous rocks at 1,000-2,000 m.—Caucasus: Dagestan.
Endemic. Described from Dagestan ASSR. Type and isotype in
Leningrad.

Note. This narrowly endemic species, so far known only from the
type locality (mear the village of Tsudakhar), does not show close
affinity with the other Caucasian species of the genus and, possibly,
deserves to be separated into its own section. Among the introduced
species, probably the Turkish species Pyrethrum pectinatum Hausskn.
[in Mittheil. Thuring. Bot. Ver. N. Folge, XVI (1901) 18] is close to
it, which, according to the original description also has yellow ligulate
florets and slightly divided leaves.

Section 4. Asterotricha Tzvel. sect. nova in Addenda XXV, 878.—
Profusely pubescent with more or less appressed stellate hairs, plants
5-30 cm high, always with short nonflowering branches and relatively
sparsely leafy stems. Capitula solitary, always with ligulate florets.
Involucres narrow-goblet-shaped, 3.5—10.0 mm in dia; outer involucral
bracts a third to a half as long as inner bracts. Receptacle almost flat,
punctate-tuberculate. Achenes 2.0-3.2 mm long, with 5-6 distinct,
longitudinal ribs and irregularly obtusely lobed, corona 0.2—0.3 mm
long.

Type of section: Tanacetum walteri (Winkl.) Tzvel.

Note. Besides the two species described below, this section also
includes T. khorassanicum (Krasch.) Tzvel. comb. nova [= Pyrethrum
khorassanicum Krasch. in Fedde, Repert. XXVI (1928) 27] from
northeastern Iran. All the species are closely related ecogeographical
races.

33. T. turcomanicum (Krasch.) Tzvel. comb. nova.—Pyrethrum
turcomanicum Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
IX (1946) 158.—Ic.: Krasch. ibid. 159} fies 2.

Perennial. Plant 10-30 cm high, with thick, many-branched rhizome,
grayish-green from profuse pubescence of appressed stellate hairs.
Stems usually rather numerous, less often solitary, erect or basally
ascending, sparsely leafy, more or less branched only at base, less
often with 1-2 branches arising from axil of upper leaves. Leaves
grayish-green from profuse pubescence, with inconspicuous punctate
glands; basal and lower cauline leaves up to 5 cm long and 1 cm wide,

343

with rather long petiole thickened at base, twice pinnately cut, oblong-
linear; their primary segments 5—10 on each side, sometimes pinnately
lobed; terminal lobes lanceolate-ovate to oblong-lanceolate, up to 1
mm wide, somewhat thick, obtuse but usually with short cusp; middle
and upper cauline leaves strongly reduced and less divided, sessile or
subsessile. Capitula solitary (but usually numerous on a plant),
sometimes two or three (but usually numerous on plant) on single
stem; peduncles up to 10(15) cm long. Involucres 3.5-6.0 mm in dia
and 5.0-6.5 mm long, more or less hairy; involucral bracts coriaceous-
herbaceous, outer lanceolate-ovate, subacute, inner broadly linear, 2—
3 times as long as outer, all bracts with relatively narrow, blackish-
brown, membranous border, but apically broadened in inner bracts as
small, usually more or less fimbriate appendage. Peripheral pistillate
florets ligulate, yellow, 2—8 with corolla tube 1.0-1.4 mm long and
limb 1.5—2.5 mm long and about | mm wide; corolla of tubular florets
of disk 2.5-3.5 mm long. Achenes 2.6—3.2 mm long and about 0.9 mm
wide, with 5 less often 6 longitudinal ribs and corona obtusely lobed,
about 0.3 mm long. Flowering June to July.

Stony and rubbly slopes at 1,500—2,000 m.—Soviet Central Asia:
mountainous Turkmenia. General distribution: Iran-Afghanistan
(Kopetdag). Described from Kopetdag (near Ashkhabad). Type and
isotypes in Leningrad.

34. T. walteri (Winkl.) Tzvel. comb. nova.—Chrysanthemum
walteri Winkl. in Tr. Peterb. Bot. Sada, XI (1889) 123.—Pyrethrum
walteri (Winkl.) B. Fedtsch. in O. and B. Fedtsch. Perech. Rast. Turk.
IV (1911) 189.—Ic.: Winkl. op. cit. t. 3 —Exs.: GRF No. 3266.

Perennial. Plant 5—20 cm high, with thick, many-branched rhizome,
grayish-green from rather profuse pubescence of appressed stellate hairs.
Stems usually quite numerous, less often solitary, erect, sparsely leafy,
more or less branched only at base. Leaves as in previous species;
basal leaves up to 4 cm long and 1.2 cm wide, with much shorter (in
comparison with width) lamina, primary segments 3-6 on each side.
Capitula solitary (but usually numerous on a plant), peduncles up to 6—
7 cm long. Involucres 6-10 mm wide and 5.5—7.0 mm long, profusely
hairy; involucral bracts as in previous species. Peripheral pistillate
florets ligulate, yellow, 8—16; corolla tube about 1.2 mm long and limb
3.0-3.5 mm long and 1.2-1.5 mm wide; corolla of tubular floret of
disk 2.4-3.2 mm long. Achenes 2.0—2.6 mm long and about 0.7 mm
wide, with 5 less often 6, longitudinal ribs and irregularly blunt-toothed,
corona 0.2-0.3 mm long. Flowering June to July.

Rocks, stony slopes at 1,800-3,000 m.—Soviet Central Asia:
mountainous Turkmenia. General distribution: Iran-Afghanistan
(Kopetdag). Described from Kopetdag. Type in Leningrad.

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344

Note. In relation to the previous species, it is a higher-mountain,
mainly saxicolous race.

GENUS 1539. Hemipappus C. Koch 1?

C. Koch, in Linnaea, XXIV (1851) 340.

Capitula 5—60, with peduncles up to 6(10) mm long, at apices of
stem in very compact, simple or compound corymbs, homogamous,
with numerous (20-50) bisexual, tubular florets of disk. Involucres
goblet-shaped, 3.5—5.5 mm in dia and 4.0-5.5 mm long; involucral
bracts coriaceous-herbaceous, imbricate, in 3—4 irregular rows; outer
bracts lanceolate, acute, lacking membranous border, inner broadly-
linear, subobtuse, 1.5—2 times as long as outer, with narrow, light-
colored, membranous border apically broadened into small oval
appendage. Receptacle flat, full, more or less covered with sparse hairs
(to subglabrous), weakly punctate-tuberculate. Corolla of disk florets
yellow, 2.5-3.2 mm long, with tube slightly compressed dorsally, weakly
and gradually broadened in upper half, with 5 lanceolate deltoid, curved
teeth, a fifth to a fourth as long as tube; stamen filaments thickened in
upper part; anthers lacking distinct basal appendage, but with lanceolate-
ovate, somewhat obtuse apical appendages, pollen grains globose,
spinulose; style [?stigma] bifid, with linear, truncate branches. All
achenes similar, glabrous, 1.8—2.5 mm long and about 0.6 mm wide,
almost terete, narrowing toward base, with 5 relatively weakly raised
longitudinal ribs, with pappus as corona 1.2—1.6 mm long, cut dorsally
up to or almost to base, unilateral, finely and irregularly toothed
throughout. Herbaceous perennials, more or less covered with appressed
tomentum of bifid hairs, with oblique, thick rhizome, condensed
vegetative shots (as leaf rosettes), relatively sparsely leafy erect stems,
and alternate pinnatipartite or pinnatisect leaves.

Four species of this genus occur in northeastern Turkey and
bordering regions of Iran.

Lectotype of genus: Hemipappus canus C. Koch.

Note. Differs from the genus Tanacetum L. by having homoga-
mous capitula, a sparsely hairy flat receptacle, and a distinctive struc-
ture of the pappus. The genus Hemipappus C. Koch shows very great,
but apparently only superficial, similarity to the genus Hippolytia

‘Treatment by N.N. Tzvelev.
7From the Greek words hemi—half and pappus—tuft; so named because the
pappus of the achenes in the species of this genus seems as if cut in half on one side.

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345

Poljak., having, however, achenes that completely lack a pappus, and
a receptacle that is glabrous and moderately convex.

1. H. canus C. Koch in Linnaea, XXIV (1851) 341.—Pyrethrum
argenteum var. tenuisectum Boiss. Fl. or III (1875) 354; Sosnovskii,
in Tr. Tifl. Bot. Sada, XVII (1915) 54; Grossh. Fl. Kavk. IV, 130.—
Chrysanthemum isabellinum (C. Koch) Rech. f. in Ann. Naturhist.
Mus. Wien. LVII (1950) 92, p. p.

Perennial. Plant 15-40 cm high, with thick, oblique rhizome,
grayish from profuse appressed tomentum of bifid hairs. Stems solitary
or few, erect, simple (but branched in inflorescence), leafy mainly in
lower half. Leaves grayish from profuse tomentum, with inconspicuous
punctate glands; basal leaves on condensed vegetative shoots to 8 cm
long and 3.5 cm wide, with long (exceeding lamina) petiole thickened
at base, more or less ovate to roundish, twice pinnately parted or cut,
terminal lobes linear or linear-lanceolate, up to 1.5(2.0) mm wide,
subobtuse or with very short cusp; cauline leaves like basal but much
smaller, with shorter petiole or sessile. Capitula 10—60, in very compact
compound corymb. Involucres 3.5-5.5 mm in dia and 4.0-5.5 mm
long, almost completely finely tomentose. Corolla of tubular florets
yellow, 2.5—3.2 mm long. Achenes 1.8—2.5 mm long and about 0.6 mm
wide; corona unilateral, 1.2-1.6 mm long, dorsally cut almost to base.
Flowering June to July.

Stony slopes, rocks; up to middle mountain zone. Found in Artvin
District of Turkey near its border with Adzharian ASSR. General
distribution: Northeastern Turkey. Described from Turkey. Type in
Berlin.

Note. In the Armenian SSR we may also find a close species, H.
argenteus (Lam.) Tzvel. comb. nova (=Achillea argentea Lam., Encycl.
meth. I (1783) 29.—Tanacetum argenteum Willd., Achill. (1789) 51.—
Hemipappus isabellinus C. Koch, op. cit. 341.—Pyrethrum argenteum
(Lam.) Boiss. Fl. or. III (1875) 354, excl. var.—Chrysanthemum
argenteum (Lam.) Bornm. in Fedde, Repert. LXXXIX (1944) 345,
non Willd. 1803.—C. isabellinum (C. Koch) Rech. f. op. cit. 92 p. p.),
which is widely distributed in the bordering regions of Turkey. It differs
from H. canus by having pinnatisect leaves with much wider, toothed
or shallow-lobed segments.

Tanacetum argenteum var. glabratum DC. (Prodr. VI, 131),
described from Lower Volga Region, apparently corresponds to one of
the species of section Xanthoglossa (DC) Tzvel. of the genus Tanacetum
L. found there. Unfortunately, the Herbarium of the Botanical Institute
of the Academy of Sciences of the USSR does not have type material
of this variety. The reported occurrence of Tanacetum argenteum Willd.
in the Altai (Ldb. Fl. Ross. II, 604) is also erroneous.

36

i)

346
GENUS 1540. Pseudohandelia Tzvel. Gen. Nov.!:2

Tzvel. in Addenda, XXV, 878.

Capitula 20-100, on up to 1.5 cm long peduncles, at stem apex in
compact umbellate compound corymbs, homogamous, with numerous
(over 100), bisexual, tubular florets of disk. Involucre wide, goblet-
shaped, 6—9 mm in dia and 3—5 mm long; involucral bracts herbaceous;
all bracts almost equal, outer and middle ones oblong, with wide, more
or less erose-toothed, light colored, membranous border, inner/ones
narrower, with considerably narrower membranous border often
exceeding outer florets in form of pales. Receptacle strongly convex,
conical-hemispherical (sometimes exceeding its width), smooth or with
scaly bracts along margin (inner bracts of involucre), punctate-
tuberculate with distinct chamber. Corolla of tubular florets of disk
yellow, 1.8-2.5 mm long, with very narrow tube, usually not
compressed, strongly glandular-papillose, abruptly and strongly
campanulately expanded in upper third, with 5 lanceolate-deltoid curved
teeth, on eighth to a seventh as long as tube; filaments of stamen
apically thickened; anthers lacking distinct basal appendage, but with
ovate-lanceolate, sub-obtuse apical appendage; pollen grains globose,
spinulose. Style [?Stigma] bifid; style branches linear, truncate. All
achenes similar, rather strongly glandular-papillose, arcuate, narrow-
cylindrical (6-8 times as long as wide), 1.8-2.4 mm long and about
0.3 mm wide, with 4-5 very weakly raised longitudinal veins as ribs,
lacking pappus, in pericarp with irregular rows of mucous cells,
mucilagenous during soaking. Biennial or perennial, monocarpic plants
covered with profuse cobwebby tomentum of simple crisped hairs,
with thick tap root, erect simple (branched only in inflorescence) stem,
and alternate bipinnatisect leaves, with narrow-linear lobes.

A monotypic genus

Note. The only species of this genus—Pseudohandelia umbellifera
(Boiss.) Tzvel., on the basis of a whole series of characters cannot be
included in any of the known genera of the tribe Anthemideae. For
example, it is easily distinguished from the genus Tanacetum L. by the
complete absence of a pappus, very narrow (compared to length), ar-
cuate achenes covered with glandular papillae, a corolla campanulately
widened in the upper third with a very long and narrow tube, a distinc-
tive structure of the involucre, homogamous capitula, not to mention
the significant habit differences. These same characters, except the
absence of the pappus, distinguish the genus Pseudohandelia from the

‘Treatment by N.N. Tzvelev.
*From the Greek word pseudos—false, and the generic name Handelia Heimerl.;
named for the considerable morphological similarity with this genus.

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347

genus Ajania Poljak. Perhaps there is the most justification to relate
the genus Pseudohandelia to the genus Cancrinia Kar. and Kir., pri-
marily to the species C. tadshikorum (Kudr.) Tzvel. However, it is
well distinguished even from this genus by the shape of the achenes,
which are entirely devoid of the pappus, and also by the structure of
the corolla, receptacle, and involucre. It is no less close to the genus
Handelia Heimerl., with which its strong similarity is not only external
but extends also to the structure of the involucre (its inner bracts are
like pales and often extend above the receptacle) and the corolla.

In my opinion, the genus Pseudohandelia is, as it were, a connecting
link between the genera of the tribe Anthemideae having a receptacle
that is covered with chaff and genera of this tribe without chaff. This
clearly shows the artificiality of dividing the tribe Anthemideae into
two groups on the basis of this character, which may be accepted only
for the sake of convenience.

1. P. umbellifera (Boiss.) Tzvel. comb. nova.—Tanacetum
umbelliferum Boiss. Diagn. ser. 2, III (1856) 30; O. and B. Fedtsch.
Perech. Rast. Turk. IV, 206.—T. trichophyllum, Rgl. and Schmalh. in
Tr. Peterb. Bot. Sada, V (1877) 255.—Pyrethrum umbelliferum (Boiss.)
Boiss. Fl. or. III (1875) 352.—P. trichophyllum (Rgl. and Schmalh.)
Bornm. and Sint. in Sintenis. Iter transcasp.-pers. 1900-1901, No.
172b.—Chrysanthemum trichophyllum (Rgl. and Schmalh.) O. Ktze.
in Tr. Peterb. Bot. Sada, X (1887) 202.—C. umbelliferum (Boiss.)
Hoffm. in Pauls. Pl. coll. in As. Med. and Pers. (1909) 149; B. Fedtsch.
Rast. Turk. 738.—C. floccosum Kitamura in Acta Phyt. et Geobot.
XVII (1957) 34 and Fl. Afghan. (1960) 400.—Lepidolopsis umbellifera
(Boiss.) Poljak. in Fl. Turkm. VII (1960) 105.—Ic.: Kitam. op. cit.
(1960) 96.—Exs.: Ed. Horti Petrop. No. 96; Sintenis, |. c.; H.F.A.M.
No. 193a, b.

Biennial or perennial. Plant 20-100 cm high, with thick, cordlike
root, more or less covered with profuse cobwebby tomentum and usually
grayish from it. Stem solitary or few, rather thick, erect, branched only
apically (in inflorescence), basally covered with sheaths of old leaves
with profuse tomentum in their axils. Leaves grayish from lax cobwebby
tomentum, sometimes glabrescent, lacking distinct punctate glands;
basal leaves to 25-30 cm long and 5-6 cm wide, withering before
flowering or persisting, linear, toward base gradually narrowed into
short petiole, twice pinnately cut, with reduced intermediate primary
segments, second order segments palmately or pinnately 2-5 parted, at
different level than primary segments, giving an impression of whorled
arrangement of terminal lobes, latter narrow-linear, somewhat thick,
up to 0.6 mm wide, with cartilaginous cusp; cauline leaves like basal,

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348

sessile or with very short petiole, uppermost leaves strongly reduced.
Capitula numerous, in compact umbellate compound corymb. Involucres
6-9 mm in dia and 3—5 mm long. Corolla of tubular florets yellow,
1.8-2.5 mm long. Achenes 1.8—2.4 mm long and about 0.3 mm wide,
lacking corona. Flowering May to July. (Plate XV).

Stony slopes, sands, gravel beds; up to lower mountain zone.—
Soviet Central Asia: Lake Balkhash Region (southern part), Tien-Shan,
Syr-Darya, Amu-Darya, Pamiro Alai Region, mountainous Turkmenia.
General distribution: Iran-Afghanistan, Dzhungaria-Kashgaria. De-
scribed from Afghanistan. Type in Geneva; isotype in Leningrad.

Note. A comparison of the type material of Tanacetum
trichophyllum Rgl. and Schmalh. with the isotype of T. umbelliferum
Boiss. shows a complete lack of differences between these species.

GENUS 1541. Dendranthema (DC.) Des Moul. Emend. Tzvel.!?

Des Moul. in Act. Soc. Linn. Bord. XX (1853) 561, p. p.—Pyre-
thrum sect. Dendranthema DC. Prodr. VI (1837) 62.—Chrysanthe-
mum sect. Pyrethrum subsect. Dendranthema (DC.) Kitam. in Acta
Phyt. et. Geobot. [TV (1935) 36 and Compos. japon. II (1940) 350 p.
p.

Capitula solitary at stem apex and its leafy branches (if present)
or more numerous (up to 20), and then in more or less regular, lax,
corymbose inflorescence, heterogamous, with 10-30 (in cultivated
species often more than 30) pistillate, peripheral, ligulate florets in a
single row (in cultivated species often in several rows) and numerous
bisexual, tubular florets of disk. Involucres saucer-shaped, (6)8—20(40)
mm in dia and 4—10(15) mm long; involucral bracts herbaceous,
imbricate, in 3, less often 2-4, irregular rows; outer bracts oblong-
ovate to narrowly linear, obtuse or subobtuse, inner oblong to nar-
rowly linear, usually not more than 1.5 times, less often almost 2 times
as long as outer; all bracts with more or less wide membranous border.
Receptacle strongly convex, almost hemispherical, often weakly coni-
cal, full, glabrous, less often (section Haplophylla) more or less hairy,
very weakly punctate-tuberculate, often distinctly alveolate. Corollas
of ligulate florets white, pink or different shades of yellow, with more
or less dorsally compressed, narrow-winged or wingless tube, 0.6—3.0
mm long, and oblong or linear limb, 8-30(60) mm long; corolla of
tubular florets of disk yellow, 2-3 mm long, tube rather weakly and

'Treatment by N.N. Tzvelev.
"From the Greek words dendron—tree, and anthemos—flower; named for the
woodiness of the base of stems in some species of this genus.

366

ta

ew

NY Sin nealhi D
wy an Pasay

Plate XV.
Pseudohandelia umbellifera (Boiss.) Tzvel., habit, involucre, corolla of tubular
florets, achene.

349

367

350

gradually (rather abruptly in section Haplophylla) broadened in upper
half, with 5 deltoid or lanceolately deltoid, weakly curved teeth, a fifth
to a fourth as long as tube; filaments of stamens thickened in upper
part; anthers lacking distinct basal appendage, with lanceolate-ovate or
oblong, subobtuse or obtuse (roundish) apical appendage; pollen grains
globose, spinulose; styles [?stigma] bifid, style branches linear, truncate.
All achenes similar (in ligulate florets sometimes not developed),
glabrous, 1.5—3.0 mm long and about 0.5—0.8 mm wide, almost terete,
narrowed toward base, with 6-8 longitudinal veins, very weakly raised
as faint ribs, pericarps lacking secretory canaliculi, but usually with
muciferous cells in irregular rows almost allover surface of achenes;
pappus completely absent.

Perennial herbs, less often half-shrubs, more or less covered with
mixed bifid and simple hairs or only simple hairs, sometimes entirely
glabrous, with erect (rarely ascending), simple or branched stems and
alternate leaves, with more or less incised, lqbed, or rarely entire blades
(section Haplophylla).

The approximately 50 species of this genus are distributed mainly
in China, Japan, Korea, Mongolia, and the regions of the Soviet Union
bordering these countries; a few of them are found in the arctic regions
of all [the northern] continents, and only one species is distributed
very sporadically in the vast Eurasian area, extending far to the west,
reaching the Carpathians. Fifteen species have been recorded in the
USSR; two of these are found only under cultivation.

Type of genus: Dendranthema indicum (L.) Des Moul.

Note. Many species of this genus, including the widely distributed
species D. zawadskii, were referred by De Candolle (Prodromus, VI,
1837) and Ledebour (FI. Ross. II, 1845-1846), and following them by
most of the authors of the later floras and identification manuals for
different regions of the USSR, to the genus Leucanthemum Mill., whose
species also have achenes without any pappus at least in the tubular
florets. However, in all other characters, both morphological and
anatomical, the achenes in the genus Dendranthema are entirely
different from those in Leucanthemum, as quite correctly noted by
Pawlowsky (B. Pawlowsky, Ochronie Przyrody, 1934, p. 14, and FI.
Polon, exs. No. 272), who transferred D. zawadskii to the genus
Tanacetum L. after combining Pyrethrum Zinn. with the latter.
Pawlowsky paid attention also to the considerable differences in the
achenes of D. zawadskii from all other species of the genus Tanacetum
L. (complete absence of pappus, weakly developed ribs, presence of
muciferous cells in pericarp), indicating that it possibly belongs to a
separate, new genus. Moreover, D. zawadskii, with a whole series of
related species, quite closely borders on the widely cultivated

354

ornamental chrysanthemums, D. indicum and D. morifolium
(=D. sinense), which were already segregated by De Candolle, based
on incidental morphological peculiarities partly arising in cultivation,
into the separate section Dendrathema of the genus Pyrethrum, which
was later elevated to the rank of genus. Since D. indicum is the type
species of this genus, we considered it necessary to retain the generic
name Dendranthema for the whole, broad group of species that border
on D. zawadskii and D. indicum, despite considerable changes in the
circumscription of this genus and its characterization. In the accepted
circumscription, this genus occupies a rather independent position in
the tribe Anthemideae, showing quite clear and close affinity only to
the genera Brachanthemum DC., Ajania P. Pol. and Tridactylina (DC.)
Sch. Bip. The extremely interrupted range of the most widely distributed
species, D. zawadskii, and the concentration of most of the species of
the genus in the countries of Southeast Asia, generally rich with relicts,
undoubtedly speak of the considerable, though relative, antiquity of
the genus Dendranthema. The genus is divided into three natural
sections, phylogenetically related to each other, which could, however,
be considered as separate genera.

Economic Importance. Two species of the genus—D. indicum
(L.) Des. Moul. and D. morifolium (Ramat.) Tzvel.—have long been
cultivated as ornamental plants; many other species deserve to be
introduced into cultivation.

1. Leaves undivided and entire, narrow-linear. Arctic plants, to 15
en high (sect: 'Haplophylla). i055. 20081. RRR EER iacssenes
SEE SS TEE ee ee 13. D. integrifolium (Richards.) Tavel.
Leaves more or less lobed, divided, or incised ..........::csssesseeeeees 2.
Apical appendages of anthers oblong, obtuse, broadly roundish;
achenes not becoming slippery on soaking; rhizome thick, fleshy.
Plants of coastal areas (sect. ArctantheMmum) .........ccccseeceeseeseeees 2.
+ Apical appendages of anthers lanceolate-ovate, subobtuse; achenes
becoming slippery on soaking; rhizome not fleshy (sect.
2) De Ae A eS ee ae Pee, eee ee ee eee 5.
3. Plants up to 15(20) cm high; basal leaves at apex 3-, very rarely 5-
lobed with subobtuse lobes; cauline leaves shallow 3-lobed at apex
to undivided and entire; stems simple. Arctic plants ................+
UNRRMREL SEs ob ates aL eiedns 8 cncatnctenpion 12. D. hultenii A. and D. Love.
+ Plants up to 30(35) cm high; basal leaves with 3- to 7-lobed or
divided lamina (if lobes 3, then usually with distinct wide teeth);
cauline leaves undivided and entire; stems simple or branched. Plants
efithe Pacific: Coastal Regina 74 eked RUA ewvivee A.
4. All or most basal leaves reniform to roundish, near base more or
less strongly truncate or weakly cordate and only at very base

2 *

369

352

a

10.

be

cuneately decurrent on petiole; involucral bracts with relatively
narrow, usually brownish membranous border ..................c::0000008+
CT Sicos ALL RA, ee ee a eee ee «1 10. D. kurilense Tzvel.
Basal leaves oblong to almost roundish, always with broad, cuneate
base; involucral bracts with wider, usually blackish-brown
membranous border ...............ccccceeseeeeee 11. D. arcticum (L.) Tzvel.
Ornamental plants, usually with “double” capitula (all or some
tubular florets modified into ligulate florets) and ligulate florets of
Various: COLOTS bein ea Bis Pose Be Bt 6.

Wild plants with white or pink ligulate florets in a single whorl..7.

Capitula 2.5—4.0(5.0) cm wide; involucres 8-20 mm in dia; ligulate
florets usually (in typical form) yellow, less often (in hybrid varieties)
ofiothencolornineat ic. .cacks. balewes *D. indicum (L.) Des Moul.
Capitula 5—15 cm wide; involucres 2-4 cm in dia; ligulate florets
of ‘various Colors) very rately yellow 1013.20.10. 208 cel ace
aed S5 cs AI TAR | RL ee ee *D. morifolium (Ramat.) Tzvel.
Leaves whitish- or grayish-tomentose beneath. Half-shrubs ............
at, cc Be, sale le ee he ds 1. D. sinuatum (Ldb.) Tzvel.
Leaves glabrous or sparsely hairy beneath, sometimes loosely
tomentose on both sides. Herbaceous plants .............ccsseeseeeeeee 8.

Basal and lower cauline leaves usually not more than to middle
(less often slightly more), pinnately or palmately lobed, with broadly
ovate or oblong, more or less toothed lobes ................eeseeeeeeees 9.

Basal and lower cauline leaves to deeper than middle (usually up
to narrowly winged rachis), pinnately or palmately divided or cut,

with narrower, linear or oblong-linear usually more or less incised
SEP IMCMESTY ARE ea ite AE EE och noavauctlsseonnnenensnds oatsneeeeee 10.
Basal and lower cauline leaves reniform to broadly ovate, near base
truncate or weakly cordate and only at very base cuneately decurrent
on petiole; stems near base more or less ascending, less often erect.
waves dad leal tak, A a eee Bee 2. D. erubescens (Stapf) Tzvel.
Basal and lower cauline leaves ovate to suborbicular always with
broad} cuneate base;/stemiusually erect (i270). Soares. cccetuce
EL nach ethee a RUS | Pc MER, aN 3. D. naktongense (Nakai) Tzvel.
Plant up to 20 cm high; stems thick, leafy, almost always strongly
branched near base, with short spreading branches bearing capitula;
leaves somewhat fleshy. Plants of coastal areas .............:c:ceesee
LAR. A a OS BOSE 9. D. littorale (Maekawa) Tzvel.
Usually taller plants, with thinner stem; simple or branched (above
middle); leavesnthin iH a ib Sate ee eee id
Lobes of basal leaves relatively broad (up to 3 mm wide), short-
acuminate; on the average, leaves less incised. Widely distributed
species, absent in the Pacific Coastal Territory and Sakhalin Island
coN SASS aE, ne ee a al eS Oe oo 12.

353

+ Lobes of basal leaves narrower (up to 1.5, less often 2.0 mm wide),
long-acuminate; on the average, leaves more incised. Primorsk
Rermtory and Sakhalin Island (./..0011.00...ccidecs. radacds ates ssactevvesneseves 13.

370 12. Involucral bracts with wide, blackish-brown, membranous border;

—

peduncles more or less tomentose. Arctic and alpine plants .............

Heke Hime Mn Tt. Moutvees Pek. rtadeeLhedlAe 5. D. mongolicum (Ling) Tzvel.

+ Involucral bracts with light colored or brownish membranous border;
peduncles more or less hairy but usually not tomentose. Plants of
plains and lower mountains ............. 4. D. zawadskii (Herb.) Tzvel.

13. Plant grayish from loose tomentum; involucral bracts with wide,
blackish-brown membranous border ............. 7. D. sichotense Tzvel.

+ Plant green, weakly hairy, often subglabrous; involucral bracts with
narrower, brownish membranous border ..............::ccscssesseeeeeeeenees 14.

14. _ Ligulate florets pink or pinkish-white; leaves, on the average, with
broader lobes. Sakhalin ............ 8. D. weyrichii (Maxim.) Tzvel.

+ Ligulate florets white, less often with pinkish shade; leaves, on the
average, with narrower lobes. Pacific Coastal Territory ............

Tis ge) oad ES ee 6. D. maximowiczii (Kom.) Tzvel.

Section 1. Dendranthema.—Apical appendage of anthers
lanceolate-ovate, subobtuse. Receptacle glabrous. Achenes becoming
slippery when soaked, with muciferous cells in pericarp. Plants with
thin or some vhat thick but not fleshy rhizome, and leaves with more
or less lobed or incised lamina; pubescence of bifid and simple hairs.

Type of section: type of genus.

Series 1. Sinuata Tzvel.—Half shrubs with leaves green above
and whitish- or grayish-tomentose beneath; ligulate florets pink or white.

1. D. sinuatum (Ldb.) Tzvel. comb. nova.—Chrysanthemum
sinuatum Ldb. Fl. Alt. [TV (1833) 118.—C. absinthiifolium Fisch. ex
DC. Prodr. VI (1837) 46, in syn.—C. cinerarium Steph. ex Ldb. FI.
Ross. II (1845-1846) 542, in syn.—Leucanthemum sinuatum (Ldb.)
DC. Prodr. VI (1837) 46; Ldb. Fl. Ross. Ii, 542; Kryl. Fl. Zap. Sib.
XI, 2742.—Tanacetum sinuatum (Ldb.) Sch. Bip. Tanacet. (1844) 35.—
Ic.: Ldb. Ic. pl. Fl. Ross. V (1834) t. 494.—Exs.: GRF No. 3272.

Perennial. Semishrubs, 8-35 cm tall, with thick, woody, many-
branched root, more or less covered with fine, appressed tomentum of
bifid and simple hairs. Stems woody and strongly branched at base,
simple or with few lateral branches above, ascending or erect, densely
leafy. Leaves dull green, glabrous or more or less hairy above, with
distinct punctate glands, whitish or grayish beneath from fine tomentum;
basal and lower cauline leaves with rather long petiole, but occasionally

354

longer than lamina; lamina rather variable in size and shape, ovate to
oblong, pinnately parted or cut, with 2-5 pinnately parted or entire,
oblong lobes on each side; terminal lobes oblong or ovate, subobtuse
or short-acuminate; upper cauline leaves smaller, usually less incised,
with shorter petiole, sometimes with lower lateral lobes close to their
base. Capitula solitary at apex of stem and its leafy branches. Involucre
10-18 mm in dia and 5-6 mm long, more or less finely tomentose;
involucral bracts with rather wide, blackish-brown, less often brownish
membranous border. Ligulate florets pinkish, less often white, with
relatively weakly compressed, corolla tube 2-3 mm long and limb 12-—
25 mm long; corolla of tubular florets 2.5-3.0 mm long, with 5
lanceolate-deltoid teeth. Achenes 2.4—2.8 mm long and about 0.6 mm
wide; with 5—6 inconspicuous longitudinal ribs, sometimes extending
up to apex of achene as very small teeth. Flowering July to August.

Stony slopes, rocks; in middle and upper mountain zones.— Western
Siberia: Altai. Endemic. Described from Altai. Type and isotype in
Leningrad.

Note. A very well delineated species, possibly deserving separa-
tion into its own section. It shows a quite clear, though distant, affinity
with Brachanthemum baranovii (Krasch. and Poljak.) Krasch. as well
as with the groups of unique Central Asian shrubby species of the type
of Pyrethrum roylei DC. and Chrysanthemum stoliczkae Clarke.
Possibly, occurs in China and Mongolia; in the east, extends to the
western Sayans.

Economic Importance. Deserves introduction into cultivation as
an ornamental plant.

Series 2. Indica Tzvel.—Herbaceous plants with more or less
deeply pinnately parted leaves, often grayish tomentose beneath; ligulate
florets yellow.

*D. indicum (L.) Des Moul. in Act. Soc. Linn. Bord. XX (1855)
561.—Chrysanthemum indicum L. Sp. pl. (1753) 889; Kom. in Tr.
Peterb. Bot. Sada, XXV (1907) 644; Stank. in Stank. and Tal. Opred.
Rast. Evrop. Ch. SSSR (1949) 628 p. p.; Grossh. Opred. Rast. Kavk.
458 p. p.—Matricaria indica (L.) Desr. in Lam. Encycl. III (1792)
734.—Pyrethrum indicum (L.) Cass. in Dict. Sc. Nat. XLIV (1826)
149; DC. Prodr. VI, 62.—Tanacetum indicum (L.) Sch. Bip. Tanacet.
(1844) 50.

Perennial. Plant 25-100 cm high, with thick, more or less branched
rhizome producing stolon-like underground shoots, dull green or grayish
from profuse pubescence of bifid and shorter simple hairs. Stems erect
or basally ascending, usually more or less branched, with thin, often

372

373

355

flexuous branches, densely leafy. Leaves with numerous, but not always
distinct punctate glands, green above, sparsely and short hairy to
subglabrous, dull- or grayish-green beneath from rather profuse, often
almost tomentose pubescence; basal leaves withering early, cauline
leaves up to 6—7 cm long and 4 cm wide, with rather long (1-2 cm),
almost wingless, petiole, basally lacking stipules or with pseudostipules
(of axillary shoot); their laminas usually oblong-ovate, at base truncate
or weakly cordate, deeply pinnatipartite, with larger, obtuse terminal
lobe and basally 2 pairs of oblong lateral lobes, more or less lobed or
toothed; upper cauline leaves smaller, with shorter petiole, often
undivided. Capitula 2.5-4.0(5.0) cm in dia, rather numerous, (1)5—
30(50), in lax, compound corymb or corymbose panicle, on thin, more
or less hairy, 0.3-5.0 cm long peduncles. Involucres 8-20 mm in dia,
and 5-6 mm long, more or less hairy to subglabrous; involucral bracts
with rather wide, light-colored or brownish, membranous border,
broadened at apex as large round or oval appendage in inner bracts.
Ligulate florets yellow, in cultivated varieties often of other colors
(pink, white, and others), corolla tube 2.0-2.5 mm long and limb 11-
15 mm long; corolla of tubular florets about 3 mm long. Achenes 1.5-
2.5 mm long and about 0.5—0.6 mm wide, with 5 very weakly raised,
often inconspicuous longitudinal ribs, lacking corona. Flowering August
to September.

Cultivated as an ornamental plant, in the south in gardens and
parks, in northern regions indoors and in greenhouses.—General dis-
tribution: China (eastern part), Korea, Japan. Described from garden-
grown specimens originating probably from China. Type in London.

Economic Importance. The numerous cultivars of this species
widely cultivated under the name of small-flowered or Indian chrysan-
themums are mostly hybrids of D. indicum with other white- or pink-
flowered species. These include many “double” cultivars with all or
part of the tubular florets modified into ligulate florets, whose capitula
often have a receptacle with scarious bracts along the periphery and
strongly reduced inner involucral bracts. The typical wild form of this
species with yellow, one-whorled, ligulate florets is found in cultivation
relatively rarely.

The “Korean chrysanthemum” also of hybrid origin, with a stem
that becomes woody at the base and ligulate florets of various colors
from golden-yellow to red and pink, occupies a more or less
intermediate position between this species and D. morifolium.

Series 3. Sinensia Tzvel.—Herbaceous plants; leaves usually not
deeper than to middle (less often slightly more) pinnately or palmately
lobed, with broadly ovate or oblong lobes, more or less toothed along
margin; ligulate florets white or pink of various shades.

374

356

*D. morifolium (Ramat.) Tzvel. comb. n.—Chrysanthemum
morifolium Ramat. in Journ. Hist. Nat. II (1792) 240; Kitam. Compos.
japon. II, 373.—C. indicum auct. non L.: Thunb. FI. jap. (1784) 320;
Stank. in Stank. and Tal. Opred. Rast. Evrop. Ch. SSSR. (1949) 628
p. p.; Grossh. Opred. Rast. Kavk. (1949) 458 p. p.—C. sinense Sabine
in Trans. Linn. Soc. XIV (1823) 142.—Pyrethrum sinense (Sabine)
DC. Prodr. VI (1837) 62.—Tanacetum sinense (Sabine) Sch. Bip.
Tanacet. (1844) 50.—T. morifolium (Ramat.) Kitam. op. cit. (1940)
373, nom. alt.—Dendranthema sinense (Sabine) Des Moul. in Acta.
Soc. Linn. Bord. XX (1855) 562.

Perennial. Plant 25-100 cm high, with thick, sometimes almost
woody rhizome producing stonon-like underground shoots, dull green
or grayish from profuse pubescence of bifid and shorter simple hairs.
Stem erect, more or less branched, woody at base, densely leafy. Leaves
with numerous but not always distinct punctate glands, usually dull
green above from appressed hairs to subglabrous, more or less grayish
beneath from profuse, often almost tomentose pubescence; basal leaves
withering early; cauline leaves up to 15 cm long and 8 cm wide, with
short or somewhat long petiole, their lamina very variable in shape
and extent of incision, oblong-ovate to almost round, pinnately parted
or lobed, with more or less approximate, wide segments or lobes.
Capitula 5—15 cm in dia, solitary or 2-10 at apex of each leafy branches,
on peduncles to 10(15) cm long. Involucres 2-4 cm in dia and 8-15
mm long; outer involucral bracts with relatively narrow, membranous
border, inner ones with light-colored or brownish membranous border,
apically more or less broadened like appendage. Ligulate florets white,
pink, red of various shades, sometimes yellow, with corolla tube 2-4
mm long and limb (15)20—50(60) mm long; tubular florets usually all
or some modified in ligulate florets. Achenes sometimes developed
(multiplication usually through cuttings).

A widely distributed ornamental plant; in the north, grown as an
indoor or greenhouse plant; in the south, also in open ground. Not
known in the wild state. Described from garden specimens originating
in China or Japan. Type in Paris.

Economic Importance. Numerous cultivars of this species, known
as large-flowered Japanese or Chinese chrysanthemums, have been
developed through centuries of cultivation along with the selection and
hybridization (both natural and induced) of a whole series of Japanese
and Chinese species of the genus Dendranthema. In coatrast to the
small-flowered chrysanthemums, here there is no wild species that could
be considered as the main ancestral type; but it is absolutely clear that
the main ancestors of the large-flowered chrysanthemums should, in
addition to D. indicum, be the species with the much larger (larger

3735

357

than in D. indicum), white or pink ligulate florets. Thus, according to
O. Stapf (Curtis’s Bot. Mag. CLVI, 1933. sub tab. 9330), the probable
ancestral species of D. morifolium, treated by him under the generic
name Chrysanthemum L., are in China: C. indicum L., C. vestitum
(Hemsl.) Stapf, C. erubescens Stapf, and in Japan: C. indicum L., C.
ornatum Hemsl., C. japonense Nakai, C. makinoi Matsum. and Nakai.
Essentially these same species are cited also in the works of many
other authors, for example, S. Kitamura, Compositae Japon., ll, 1940,
po 375:

Numerous works have been devoted to the special features of
cultivation and the varietal diversity of the small- and large-flowered
chrysanthemums, not to mention the rather detailed information about
them available in the different manuals on floriculture (for example, in
the book of G.E. Kiselev—Tsvetovodstvo [Floriculture], II, 1959, p.
702).

2. D. erubescens (Stapf) Tzvel. comb. nova.—Pyrethrum sinense.
Q. sinense Maxim. in Bull. Acad. Sc. Pétersb. XVII (1872) 425.—
Chrysanthemum sinense auct. non Sabine: Kom. in Tr. Peterb. Bot.
Sada, XXV (1907) 644; Kom. and Alis. Opred. Rast. Dalnevost. Kr.
II, 1031.—C. erubescens Stapf in Curtis’s Bot. Mag. CLVI (1933) sub
tab. 9330, nom. nov.—C. maximoviczianum Ling in Centr. Inst. Bot.
Ac. Peip. III (1935) 459 p. p. nom. nov.—C. naktongense auct. non
Nakai: Hand.-Mazz. in Acta Horti Gotoburg. XII (1938) 255 p. p.—
C. zawadskii var. latilobum (Maxim.) Kitam. in Acta Phyt. et Geobot.
VII (1938) 210.

Perennial. Plant 10-50 cm high, with thin, more of less branched
rhizome, more or less covered with crisped simple and appressed bifid
hairs. Stems usually few, erect or ascending, densely leafy, strongly
branched above middle, very rarely simple. Leaves with numerous
punctate glands, more or less covered with short crisped hairs to
subglabrous, green above, pale green beneath; basal leaves up to 10
cm long and 5 cm wide, with rather long petiole thickened at base,
their lamina reniform to broadly ovate, near base straight truncate or
weakly cordate, cuneately decurrent on petiole only at base, subobtuse,
3-lobed or pinnately lobed (or pinnately parted) up to a half, less often
beyond, with 2-3 approximate lateral lobes on each side; segments or
lobes oblong to broadly ovate, 5~12 mm wide, irregularly toothed or
shallow lobed; cauline leaves smaller, with shorter petiole, and ovate
or almost round lamina; uppermost leaves many, more or less toothed
or lobed, cuneately narrowed into petiole. Capitula 2-15 (very rarely
solitary), one on each branch surrounded by reduced bracteal leaves,
in lax compound corymb. Involucre 6-10 mm in dia and 2—4 mm long,

376

358

more or less hairy; outer involucral bracts narrow-linear, with brown-
ish membranous border near apex, oblong-linear, or linear, throughout
with rather wide membranous border. Ligulate florets white or pink-
ish-white, with compressed but almost wingless corolla tube 1.2—1.5
mm long and ligule 8-15 mm long and 2-3 mm wide; corolla of tu-
bular florets 1.8-2.4 mm long. Achenes 1.3-1.8 mm long, lacking
corona. Flowering September to October.

Rocks, stony slopes; up to lower mountain zone.— Far East: Ussuri
(Posjet Bay). General distribution: Northeastern China, Korea. De-
scribed from Far East. Type in London; isotypes in Leningrad.

Note. Both Chrysanthemum erubescens Stapf and C. maximo-
viczianum Ling. are new names for the variety Pyrethrum sinense Q.
sinense described by Maximowicz, which he considered the basionym
for Chrysanthemum sinense Sabine, described from cultivated speci-
mens. In fact, it is a synonym of Dendranthema morifolium. As type
specimens of D. erubescens, I am selecting the specimens collected by
Maximowicz from the area of Posjet Bay and distributed by him to
important herbaria of the world.

The systematic position of the species Chrysanthemum reflexum
Ueki (in Suigengakuho, XXI, 2 (1928) 13), described from Korea, is
not clear to me; it either is D. erubescens (in which case the name of
the latter species must be changed according to the rules of priority),
or it is a synonym of the next species.

3. D. nektongense (Nakai) Tzvel. comb. nova.—Leucanthemum
sibiricum var. latilobum (Maxim.) Primit. fl. amur. (1859) 156.—Chry-
santhemum sibiricum var. latilobum (Maxim.) Kom. in Tr. Peterb.
Bot. Sada, XXV (1907) 642; Kom. and Alis. Opred. Rast. Dalnevost.
Kr. II (1932) 1031, tabl. 307.—C. naktongense Nakai in Bot. Mag.
Tokyo, XXIII (1909) 186 and Fl. Kor. II (1911) 26, t. VIII.—C.
zawadskii var. latilobum (Maxim.) Kitam. in Acta Phyt. et Geobot.
VII (1938) 210 p. p.—C. zawadskii ssp. latilobum Kitag. Lin. fl. Mansh.
(1939) 444.—Ic.: Nakai, loc. cit.; Kom. and Allis. loc. cit.

Perennial. Plant 15—70 cm high, with thin, more or less branched
rhizome, more or less covered with crisped simple and appressed bifid
hairs. Stems usually solitary or few, erect, densely leafy, simple or
branched above middle. Leaves with numerous punctate glands, on
both sides more or less covered with short, crisped simple hairs or
subglabrous; basal leaves up to 10 cm long and 5 cm wide, with rather
long (often exceeding lamina) petiole thickened at base, oblong-ovate
to almost round, usually not deeper than middle pinnately or palmately
3-9 lobed with oblong or ovate, more or less toothed or shallow lobed

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359

lobes, basally round or truncate but always cuneately narrowed into
petiole; cauline leaves like basal but with shorter petiole and usually
with short axillary branches, their lamina oblong-ovate, cuneately nar-
rowed toward base. Capitula solitary or 2-5 (sometimes to 10), soli-
tary at apices of leafy branches of stem, sometimes forming lax irregu-
lar corymb. Involucres 10-16 mm in dia and 4-6 mm long, more or
less hairy; outer involucral bracts narrow-linear, usually with membra-
nous border only at apex; inner bracts oblong-linear, basally somewhat
narrow, with wide, light-colored or brownish, membranous border.
Ligulate florets white or light-pink, with corolla tube 1.4—-2.0 mm long
and ligule 12-26 mm long and 3.0-5.5 mm wide; corolla of tubular
florets 2.3-3.0 mm long. Achenes 1.5—2.0 mm long, lacking corona.
Flowering August to October.

Thinned pine and oak forests, scrubs, stony slopes; up to lower
mountain zone.—Far East: Ussuri (southern and southwestern part).
General distribution: China (northeastern part), Korea. Described from
Korea. Type in Tokyo.

Note. Morphologically, this species is completely different from
the widely distributed D. zawadskii, to which it has often been ap-
pended as a variety, but is quite close to D. erubescens, being a more
northern ecogeographical race linked mainly to the forest groups.

Series 4. Zawadskiana Tzvel.—Herbaceous plants; leaves (at least
basal and lower cauline) considerably deeper than middle, often up to
rachis pinnately or palmately divided or incised, with narrow-linear or
oblong-linear segments or lobes, in turn often cut or divided; ligulate
florets white or pink of various shades.

4. D. zawadskii (Herb.) Tzvel. comb. nova.—Chrysanthemum
zawadskii Herb. Addit. ad Fl. Galic. (1831) 44, t. 1; DC. Prodr. VI,
67; Trautv. in Tr. Peterb. Bot. Sada, VIII, 2 (1883) 458.—C. arcticum
auct. non L.: Ldb. FI. Alt. IV (1833) 115.—C. sibiricum Turcz. ex DC.
Prodr. VI (1837) 46, in syn.—C. gmelinii Ldb. ex Turcz. in Bull. Soc.
Nat. Mosc. I (1838) 94, nom. nud.—C. sibiricum Fisch. ex Turcz. Fl.
baic.-dahur. II (1856) 42, in syn.—C. arcticum ssp. alaunicum K.-Pol.
in sb. “25 Let Nauchno.pedag. i Obshch. Deyat. B.A. Kellera”* (1931)
318.—C. sibiricum var. acutilobum (DC.) Kom. and Alis. Opred. Rast.
Dalnevost. Kr. II (1932) 1031.—?C. hwangshanense Ling. in Contr.
Inst. Bot. Acad. Peip. III (1935) 472, t. 49.—C. kozo-poljanskii Golitz.

*In the collection of articles “25 years of Scientific-Pedagogic and Social Work
of B.A. Keller.”—Translator.

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360

in Byull. Obshch. Estestv. pri Voronezhsk. Univ. V (1949) 21, nom.
nud.—Leucanthemum sibiricum var. acutilobum DC. op. cit. 46; Turcz.
op. cit. (1856) 42.—L. sibiricum auct. non DC.: Ldb. FI. Ross. II, 541;
Kryl. Fl. Zap. Sib. XI, 2741.—L. alaunicum (K.-Pol.) Golitz, et ssp.
Kozo-Poljanskii Golitz. in Majevsky, Fl. Izd. 8 (1954) 581.—
Tanacetum gmelinii Sch. Bip. Tanacet. (1844) 35 p. p. nom. nov.—T.
alaunicum K.-Pol. in Delect. sem. horti bot. univ. Voroneg. II (1930)
30.—T. zawadskii (Herb.) B. Pawl. Ochron. Przyr. (1934) 14 and Pl.
Polon. exs. No. 272.—Pyrethrum zawadskii (Herb.) Nym. Syll. fl. Eur.
(1854-1855) 11.—P. sibiricum (DC.) Stank. in Stank. and Tal. Opred.
Rast. Evrop. Ch. SSSR (1949) 630.—Ic.: Herb. loc. cit.; Jav. Csap. Ic.
Fl. Hung. (1933) 523; Ling, loc. cit.—Exs.: Fl. exs. Austro-Hung. No.
2673; Fl. exs. reip. Bohem.-Sloven. No. 1098; Fl. Hung. exs. No. 82;
Fl. Polon. exs. No. 182; Pl. Polon. exs. No. 272; GRF No. 419.

Perennial. Plant 15-50 cm high, with thin, more or less branched
rhizome, more or less covered with short bifid and simple hairs. Stems
solitary or few, erect or at base ascending, rather densely leafy, simple
or with few branches mainly in upper half. Leaves with numerous
punctate glands, sometimes also appressed-hairy to subglabrous; basal
and lower cauline leaves to 8 cm long and 4 cm wide, with rather long
(often 1.5—2 times as long as lamina), narrow-winged petiole, ovate to
almost roundish, cuneately narrowed at base, usually twice (rarely once)
pinnately cut or divided, with 1-3 pairs of segments or lobes of first
order, often coalesced at base, terminal lobes linear to linear-oblong,
usually 2-3 mm wide, at apex short and abruptly acuminate; middle
cauline leaves with shorter, often broad-winged petiole, sometimes with
reduced lateral segments approximate to their base, usually pinnately
parted with longer, acuminate, terminal lobe; uppermost leaves entire
or pinnately lobed. Capitula solitary or 2—5, singly at apices of stem
and its leafy lateral branches, usually not forming corymb, peduncles
relatively sparsely hairy, sometimes finely tomentose. Involucres 10—
20 mm in dia and about 4.5—6.0 mm long, more or less hairy to com-
pletely glabrous; involucral bracts with rather wide, light-colored or
brownish membranous border, outer bracts broadly linear to oblong,
inner linear to oblong-linear. Ligulate florets white, pink, pinkish-vio-
let, of various shades, corolla tube 1.8—3.0 mm long and ligule 12—26
mm long and 3-6 mm wide; corolla of tubular florets about 2.5—3.0
mm long. Achenes 1.8-2.5 mm long and about 0.5—0.6 mm wide,
lacking corona. Flowering July to September.

Stony slopes, forest glades, pine and larch forests, rocks, mainly
on calcareous and sandy soils; up to middle mountain zone.—Euro-
pean part: Dvina-Pechora (Urals and along Pinega River at its
confluence with Northern Dvina), Volga-Kama, Volga-Don (limestones

361

along Don and Oskol rivers), Trans-Volga (eastern part); Western
Siberia: Ob River Area (southern part), Upper Tobol, Irtysh, Altai;
Eastern Siberia: Yenisei (southern part), Angara-Sayans, Lena-Kolyma,
Dauria; Far East: Zeya-Bureya, Ussuri (Bureyi Range). General dis-
tribution: Central Europe (Carpathians), Mongolia (northern part), China
(northeastern part). Described from Carpathians. Type in Vienna;
isotype in Leningrad.

Note. A highly polymorphic species forming throughout its wide
range numerous more or less distinctive populations distinguished
mainly by the color of the ligulate florets and the shape of the leaves.
However, comparison of the extensive material of this species from
the Carpathians, Urals, and the Baikal Region in the Herbarium of the
Botanical Institute of the Academy of Sciences of the USSR shows a
complete absence of differences among the Carpathian, Ural and Baikal
materials as a whole, given the considerable variation of populations
within each of these three regions. The segregation of plants from
geographically isolated habitats of this species in the Central Russian
Plains into separate ecogeographical races (Tanacetum alaunicum K.-
Pol. from the upper reaches of the Oskol River and Chrysanthemum
kozo-poljanskii Golitz. from the Don River in the vicinity of the village
of Plyushchan) is based more on the absence of comparable material
from other regions than on morphological differences between them
and specimens of D. zawadskii from Siberia and the Carpathians.

Economic Importance. Deserves to be introduced into cultivation
as an ornamental plant.

5. D. mongolicum (Ling) Tzvel. comb. nova.—Leucanthemum
sibiricum var. peleiolepis Trautv. in Middendorff’s Sibir. Reise I, 2
(1847) 37.—L. sibiricum var. alpina F. Schmidt, Reisen in Amur-
Lande und Sach. (1863) 49.—Chrysanthemum mongolicum Ling in
Contr. Inst. Bot. Acad. Peip. III (1935) 463, fig. 2 and t. 46.—
Tridactylina turczaninovii Krasch. in herb.—Ic.: Ling, loc. cit.

Perennial. Plant 5—25(30) cm high, with thin, more or less branched
rhizome, more or less covered with appressed, simple and bifid hairs,
often covered with fine tomentum. Stems solitary or few, erect or at
base ascending, rather densely leafy, simple or with few lateral branches
almost from base. Leaves with not always distinct punctate glands,
more or less appressed-hairy to completely glabrous; basal and lower
cauline leaves like previous species, but, on the average less incised;
middle cauline leaves usually pinnately parted, or lobed; terminal lobe
of all leaves often with relatively long cusp. Capitula solitary or 2-5,
singly at apices of stem and lateral branches, peduncles, particularly in
upper part, rather profusely tomentose. Involucres 10-18 mm in dia

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362

and 4-6 mm long, with more or less loose tomentum at base, often on
dorsal side; involucral bracts with rather wide, blackish-brown mem-
branous border. Ligulate florets white or pink, of various shades, co-
rolla tube 1.2-2.5 mm long and ligule 12-25 mm long and 3-5 mm
wide, often sterile because of a partial reduction of the pistil; corolla
of tubular florets 2.5—3.0 mm long. Achenes 1.5—2.3 mm long, lacking
corona. Flowering August to September.

Stony and rubbly slopes, gravel beds, sometimes in thinned for-
ests; in the north in stony tundra, in the south mainly on bald moun-
tains in middle and upper mountain zones.—Arctic: Arctic Siberia
(Taimyr, lower reaches of Lena and other rivers); Eastern Siberia:
Yenisei, Lena-Kolyma, Angar-Sayans, Dauria; Far East: Okhotsk, Uda
River Area, Zeya-Bureya, Ussuri (Bureya Range). General distribu-
tion: Mongolia (northern part), China (northeastern part). Described
from China. Type in Peking.

Note. In Yakutia and the Far East, plants are found with narrower
leaf lobes and in this respect seemingly transitional to D. maximowiczii.
In relation to the previous species, it is a more alpine and arctic race,
linked with it by specimens with intermediate characters.

6. D. maximowiczii (Kom.) Tzvel. comb. nova.—Chrysanthemum
maximowiczii Kom. in Izv. Peterb. Bot. Sada, XVI (1916) 179; Kom.
and Alis. Opred. Rast. Dalnevost. Kr. II (1932) 1028.—C.
maximovicziianum var. dissectum Ling in Contr. Inst. Bot. Akad. Peip.
III (1935) 461, fig. la-e.—C. nactongense var. dissectum (Ling) Hand.-
Mazz. in Acta Horti Gotoburg. XII (1938) 256.—C. zawadskii ssp.
acutilobum (DC.) Kitag. Lin. fl. Mansh. (1939) 444 p. p.—Ic.: Ling,
be Gs

Biennial. Plant 8-40 cm high, with thin, more or less branched
rhizome, covered with short bifid and simple hairs, sometimes finely
cobwebby tomentum in upper part. Stems solitary or few, erect, simple
or with few branches. Leaves with inconspicuous punctate glands, more
or less sparsely hairy to completely glabrous; basal and lower cauline
leaves 8-10 cm long, and 4 cm wide, with rather long petiole, often
longer than lamina, ovate to almost roundish, twice pinnately cut, with
1-3 pairs of first order segments, terminal lobe narrow-linear to lin-
ear-lanceolate, up to 1.5(2.0) mm wide, relatively long-acuminate;
middle cauline leaves usually less incised, but also with terminal lobe
narrower than in previous species. Capitula usually 2-5 on stem, soli-
tary at apices of leafy branches, often forming lax irregular corymbose
inflorescence, peduncles more or less hairy to finely tomentose. In-
volucres 10-18 mm in dia and about 4.5—5.5 mm long, more or less
hairy; involucral bracts with rather wide, light colored or brownish

380

383

363

membranous border. Ligulate florets white, less often pinkish-white,
with corolla tube 1.8—2.5 mm long and ligule 12-24 mm long and 3-
5 mm wide; corolla of tubular florets 2.5—-3.0 mm long. Achenes 1.5—
2.5 mm long, lacking corona. Flowering August to September.

Rocks, stony slopes; up to middle mountain zone.—Far East:
Ussuri. General distribution: China (northeastern part), Korea. De-
scribed from Far East (southern Sikhote-Alin). Type and isotype in
Leningrad.

Note. Possibly, Chrysanthemum coreanum (Lev. and Vant.) Nakai
ex Mori (Enum. Cur. Pl. (1922) 352 = Matricaria coreana Lev. and
Vant. in Fedde, Repert. VIII (1910) 169), a species described from
Korea, belongs to this species (and in relation to it is a priority name).
It is difficult to say whether it belongs to any other species or should
be separate, as there is not reliably identified herbarium material of
this species.

7. D. sichotense Tzvel. sp. nova in Addenda, XXV, 879.—Chry-
santhemum sibiricum var. alpinum Nakai in Bot. Mag. Tokyo, XXXI
(1917) 109.—C. zawadskii var. alpinum (Nakai) Kitam. in Acta Phyt.
et Geobot. VII (1938) 210.

Perennial. Plant 10-25 cm high, with thin, more or less branched
rhizome, grayish-tomentose, mixed with long, simple and bifid hairs.
Stems solitary or few, erect, densely leafy, simple or with (1-2) lateral
branches in upper part. Leaves grayish-green from lax tomentum, with
inconspicuous punctate glands; basal and lower cauline leaves
numerous, up to 4 cm long and 2 cm wide, with rather long (often
longer than lamina) petiole, reniform to almost roundish, twice pinnately
cut, with 2-3 pairs of lobes; terminal lobes narrow-linear (to 1 mm
wide), with long- and gradually acuminate apex; middle and upper
cauline leaves smaller, with shorter petiole or sessile, uppermost leaves
with oblong lamina, pinnately parted, with 3-5 pairs of entire or more
or less divided lobes. Capitula usually solitary, less often 2-3, on
single stem, on relatively short peduncles. Involucres 12-18 mm in dia
and 4.5—6.0 mm long, dorsally lanate-tomentose; involucral bracts with
wide, blackish-brown membranous border. Ligulate florets pink or
violet-pink, corolla tube 2.0-2.5 mm long and ligule 12-24 mm long;
corolla of tubular florets about 3 mm long. Achenes 1.5—2.0 mm long,
lacking corona. Flowering July to August (Plate XVI, Fig.1).

Rubbly and stony slopes, balds, above 1,200 m.—Far East: Ussuri
(southern Sikhote-Alin). General distribution: Korea. Described from
southern Sikhote-Alin. Type (Snezhnaya Mountain in southern Sikhote
Alin, at about 1,500 m 15.VII.1930, No. 767, I. Schischkin) and isotype
in Leningrad.

381 Plate XVI.
1 — Dendranthema sichotense Tzvel., habit, corolla of ligulate floret, corolla
of tubular floret, anther tube, achene; 2 — D. integrifolium (Richards.) Tzvel.,
habit; 3 — D. hultenii (A. and D. Léve) Tzvel., habit, corolla of ligulate floret,
corolla of tubular floret, anther tube.

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365

Note. In relation to the previous species, it is a higher mountain
(bald mountain) ecogeographical race, differing from the former by
the abundant tomentose pubescence of the whole plant, color of the
ligulate florets, blackish-brown membranous border of the involucral
bracts, and the form of the basal leaves.

Economic Importance. Deserves to be introduced into cultivation
as an ornamental plant.

8. D. weyrichii (Maxim.) Tzvel. comb. nova.—Leucanthemum
weyrichii Maxim. Prim. fl. amur. (1859) 156.—Chrysanthemum
hakusanense Makino in Bot. Mag. Tokyo, XXIV (1910) 302.—C.
lucidum Nakai in Bot. Mag. Tokyo, XXXII (1918) 110.—C. weyrichii
(Maxim.) Miyabe and Miyake, Fl. Saghal. (1915) 251.—Tanacetum
weyrichii (Maxim.) Kitam. Compos. Japon. II (1940) 348, nom. alt.—
Ic.: Makino, op. cit. 303, fig. XXI.

Perennial. Plant 20-50 cm high, with relatively thin, more or less
branched rhizome, more or less covered with short bifid and simple
hairs. Stems solitary or few, erect or at base ascending, usually branched
above middle, less often simple, relatively sparsely leafy. Leaves
glabrous or subglabrous, with inconspicuous punctate glands; basal
and lower cauline leaves to 8-10 cm long and 4 cm wide, with rather
long petiole, ovate or broadly ovate, twice pinnately cut (or divided),
with 1-3 pairs of primary segments, usually more or less approximate;
terminal lobe lanceolate to lanceolate-linear, usually 1-2 mm wide, on
the average wider than in D. maximowiczii, but narrower than in D.
zawadskii, rather long-acuminate; middle cauline leaves usually less
incised, with rather shorter petiole to subsessile. Capitula usually 2-8
on single stem (solitary at apices of its leafy lateral branches), often
forming lax corymbose inflorescence, less often solitary; peduncles
usually hairy to finely tomentose. Involucres 10-15 mm in dia and
4.0—5.5 mm long, more or less hairy to completely glabrous; involucral
bracts with rather wide, brownish membranous border. Ligulate florets
pink or pinkish-white, with corolla tube 1.0-1.8 mm long and ligule
12-18 mm long; corolla of tubular florets 2.4—2.8 mm long. Achenes
1.5-2.2 mm long, with somewhat acute, raised upper border but lacking
corona. Flowering August to September.

Stony slopes, rocks; up to lower mountain zone.—Far East:
Sakhalin. General distribution: Japan (northern part), Korea(?).
Described from Sakhalin. Type in Leningrad.

Note. Very similar to D. zawadskii and D. maximowiczii; it is an
ecogeographical race occupying an intermediate position between them
with respect to the nature of the leaf incisions.

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9. D. littorale (Maekawa) Tzvel. comb. nova.—Chrysanthemum
littorale Maekawa in Trans. Sapporo Nat. Hist. Soc. VIII (1921) 15,
t. 1, fig. 14-22.—C. weyrichii var. littorale (Maekawa) Kudo, Contr.
Knowl. North Saghal. (1923) 58.—Ic.: Maekawa, I. c.

Perennial. Plant 10-20 cm high, with relatively thin, more or less
branched rhizome, more or less covered with short bifid hairs to
subglabrous. Stems solitary or few, somewhat thick, almost always
branched near base, with short divergent branches, densely leafy
(particularly in lower part), leaves glabrous or subglabrous, with
numerous punctate glands, somewhat thick; basal and lower cauline
leaves to 10 cm long and 5.5 cm wide, with long (usually longer than
lamina), narrow-winged petiole thickened at base, reniform to ovate,
with narrow-cuneate base, twice pinnately parted (or divided), with 1—
3 approximate pairs of lateral lobes, terminal lobes linear to broadly
lanceolate, sometimes some shallow-lobed or toothed, 1-3 mm wide,
short-acuminate; middle cauline leaves smaller, cuneately narrowed
into short winged petiole, ovate or oblong, often pinnately parted,
uppermost leaves some undivided and entire. Capitula usually 2-8,
solitary, at apices of stem and its leafy lateral branches on relatively
short (1-3 cm long), more or less hairy peduncles, less often single.
Involucre 12-18 mm in dia and 4-6 mm long, glabrous or subglabrous;
outer involucral bracts usually almost lacking membranous border, inner
with wide, light colored or brownish membranous border. Ligulate florets
pink or pinkish-white, corolla tube 1.5—2.0 mm long and limb 16-25 mm
long and 3.0-5.5 mm wide; corolla of tubular florets 2.5—3.0 mm long.
Achenes 1.5—2.2 mm long, with somewhat acute raised upper border,
but lacking corona. Flowering August to September.

Rocks and stony slopes of seacoasts.—Far East: Sakhalin, Kuril
Islands. General distribution: Japan (northern part). Described from
Japan (Hokkaido Island). Type in Sapporo.

Note. This species usually has been united with the previous spe-
cies despite quite consistent habit differences and a very definite eco-
logical cohesiveness. On Sakhalin Island, both species apparently are
separated geographically as well. In habit, it resembles the species of
the next section, to which it is also ecologically close.

Section 2. Arctanthemum Tzvel. sect. nova in Addenda XXV,
879.—Apical appendage of anthers oblong, obtuse, broadly round;
receptacle glabrous; achenes not becoming slippery when soaked. Plants
with thick, fleshy rhizome and leaves with more or less lobed or divided,
somewhat thick lamina; hairs long and simple, often entirely lacking.

Type of section: Dendranthema arcticum (L.) Tzvel.

Note. S. Kitamura (S. Kitamura, Compos. Japon. II, 1940, 345—-
349), based only on the achenes not becoming slippery when soaked,

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367

refers species of this section, as well as D. weyrichii from the previous
section, to his subsection “Leucanthemum’” of the genus Chrysanthemum
L. sensu lato, but D. zawadskii, with the group of closely related spe-
cies to subsection “Dendranthema.” However, the character—slipperi-
ness of the fruit wall hardly deserves such importance, particularly
when it is not accompanied by other more or less substantial charac-
ters. Moreover, the slippery fruit wall is an adaptive character linked
with a relatively hot and dry climate, and it is not surprising that in
species confined to more humid and much colder regions the mucifer-
ous cells of the fruit wall disappear entirely. Regarding other charac-
ters (structure of achenes, corolla, receptacle, involucres, and others),
the species of section Arctanthemum show incomparably great close-
ness to the genus Dendranthema, to which I refer them, rather than to
the genus Leucanthemum Mill.

All three species of the section are closely related ecogeographical
races of a single series.

10. D. kurilense Tzvel. nom. nov.—Chrysanthemum arcticum var.
yezoense Maekawa in Trans. Sapporo Nat. Hist. Soc. VIII (1921) 14,
t. 1, fig. 1-4, non Chrysanthemum yezoense Maekawa, ibid. 12.—Ic.:
Maekawa, Ibid.

Perennial. Plant 10-35 cm high, with thick, fleshy rhizome,
subglabrous. Stems solitary or few, erect, simple or with few lateral
branches, leafy mainly in lower half. Leaves glabrous or subglabrous
(cobwebby pubescence disappearing later), lacking distinct punctate
glands; basal and lower cauline leaves numerous, to 10 cm long and
5 cm wide, with long petiole thickened at base, reniform to rotund,
near base more or less truncate or weakly cordate and only at base
cuneately decurrent on petiole, with 3-5 palmate lobes usually not
divided to more than middle, lobes obtusely and irregularly toothed or
shallow lobed; middle cauline leaves strongly reduced, with narrow
winged petiole or subsessile, oblong, more or less pinnately lobed
lobes, with short cartilaginous cusp. Capitula solitary or 2—5 on single
stem, on rather long (to 8 cm) peduncles. Involucres 12—18 mm in dia
and 5—7 mm long, glabrous or at base more or less cobwebby lanate.
Involucral bracts with brownish membranous border. Ligulate florets
white, corolla tube 0.8—1.5 mm long and lobe 10-22 mm long and 3-
5 mm wide; corolla of tubular florets 2.3—3.0 mm long. Achenes 1.8-
2.6 mm long and about 0.5 mm wide, at apex lacking corona but with
indistinctly raised, somewhat acute upper border. Flowering July to
September.

Coastal rocks, gravel beds.—Far East: Kuril Islands (Iturup Island).
General Distribution: Japan (northern part). Described from Japan
(Hokkaido Island). Type in Sapporo.

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368

Note. Chrysanthemum yezoense Maekawa (=C. arcticum ssp.
maekawanum Kitam. Compos. japon. II, 1940, p. 347) from the Island
of Hokkaido, with anthers having lanceolate-ovate, subacute, apical
appendages, apparently belongs to the preceding section, differing from
D. littorale only by the less divided leaves with wider lobes.

11. D. arcticum (L.) Tzvel. comb. nova.—Chrysanthemum
arcticum L. Sp. pl. (1753) 889.—C. adustum Fisch. ex Trautv. in Tr.
Peterb. Bot. Sada, VIII, 2 (1883) 457.—C. arcticum ssp. gmelinii (Ldb.)
Kitam. in Acta Phyt. et Geo-bot. IV (1935) 37.—Matricaria arctica
Desr. in Lam. Encycl. III (1792) 735.—Leucanthemum arcticum (L.)
DC. Prodr. VI (1837) 45, quoad nomen.; Kom. and Alis. Opred. Rast.
Dalnevost. Kr. II, 1031.—L. sibiricum DC. op. cit. 46, excl. var.—L.
gmelinii Ldb. Fl. Ross. II (1845-1846) 541.—Tanacetum arcticum
(L.) Sch. Bip. Tanacet. (1844) 35.—T. gmelinii Sch. Bip. ibid. 35, p.
p., nom. novum (=Leucanthemum sibiricum DC.).—Pyrethrum arcticum
(L.) Stank. in Stank. and Tal. Opred. Rast. Evrop. Ch. SSSR (1949)
630, quoad nomen.—Ic.: Gmel. Fl. Sib. II (1749) 203, t. 84; Hult. FI.
Kamtch. IV (1930) 173.

Perennial. Plant 10-30 cm high, with thick, fleshy, creeping
rhizome, glabrous or subglabrous (with cobwebby pubescence
disappearing later). Stems solitary or few, erect or basally ascending,
simple or with few lateral branches, leafy mainly in lower half. Leaves
glabrous or subglabrous, lacking punctate glands; basal and lower
cauline leaves rather numerous, to 9-12 cm long and 3-4 cm wide,
with long petiole thickened at base, oblong to almost round, always
with broad cuneate base, palmately or pinnately lobed usually not more
than to middle, with 3—7 obtuse or subobtuse lobes, large and coarsely
toothed or entire; middle cauline leaves strongly reduced, with short
winged petiole or subsessile, pinnately or palmately lobed, uppermost
leaves linear, entire. Capitula solitary or 2—5 on single stem, on rather
long peduncle. Involucres 12—22 mm in dia and 4—7 mm long, glabrous
or at base more or less cobwebby-woolly; involucral bracts with rather
wide, blackish-brown, less often brownish membranous border. Ligulate
florets white, corolla tube 0.8—1.5 mm long and limb 10-22 mm long
and 3-5 mm wide; corolla of tubular florets 2.3-3.0 mm long. Achenes
1.8—2.6 mm long and about 0.5 mm wide, apically lacking corona, but
with indistinctly raised, subacute border. Flowering July to September.

Stony and sandy places, gravel-beds, grassy patches along coast.—
Far East: Kamchatka, Comander Islands, Okhotsk, Uda River Area,
Sakhalin, Kuril Islands, Ussuri. General distribution: Pacific Coast of
North America (south up to U.S.A.). Described from Kamchatka.
Gmelin’s drawing cited above is the type specimen.

38

oo

369

Note. Although Gmelin’s drawing is the type for this species, both
De Candolle and Ledebour regarded the following species [D. hultenii]
as Chrysanthemum arcticum L. and on the basis of Gmelin’s drawing
described two new species (Leucanthemum sibiricum DC. and L.
gmelinii Ldb.), which are more recent synonyms of Chrysanthemum
arcticum L.

12. D. hultenii (A. and D. Léve) Tzvel. comb. nova.—Leucanthe-
mum arcticum auct.: DC. Prodr. VI (1837) 45, quoad. plant.; Ldb. FI.
Ross. II, 541; Kryl. Fl. Zap. Sib. XI, 2741, non Chrysanthemum
arcticum L.— L. hultenii A. and D. Love in Bot. Notis. (1961) 44.—
Pyrethrum arcticum (L.) Stank. in Stank. and Tal. Opred. Rast. Evrop.
Ch. SSSR (1949) 630, quoad plant.—Chrysanthemum arcticum ssp.
polaris Hult. in Sv. Bot. Tidskr. XLIII (1949) 776.—Ic.: Fedtsch. and
Fler. Evrop. Ross. (1911) 975.

Perennial. Plant 6—20 cm high, with thick, fleshy, creeping rhizome,
glabrous, less often subglabrous. Stems solitary or few, erect or basally
ascending, simple, leafy mainly in lower half. Leaves glabrous, lacking
punctate glands; basal and lower cauline leaves rather numerous, to 8
cm long and 3 cm wide, with long (sometimes 1.5—2 times as long as
lamina), more or less winged petiole, oblong to almost round, cuneately
narrowed toward base, in upper part with 3, less often 5 shallow,
obtuse lobes or teeth; middle cauline leaves strongly reduced, usually
entire or subentire. Capitula solitary. Involucres 12-18 mm in dia and
4—6 mm long, glabrous; involucral bracts with wide, blackish-brown,
membranous border. Ligulate florets white, corolla tube 0.6-1.5 mm
long and limb 8-15 mm long and 2-4 mm wide; corolla of tubular
florets 2.5—-3.0 mm long. Achenes 1.5—2.2 mm long and about 0.5 mm
wide, lacking corona. Flowering July to August. (Plate XVI, Fig. 3).

Stony and sandy places, gravel beds, grassy patches along coast.—
Arctic: Arctic Europe, Novaya Zemlya, Arctic Siberia (rarely),
Chukotka, Anadyr. General distribution: Arctic. Described from
European Arctic. Type in Stockholm.

Note. Specimens intermediate between this and the previous spe-
cies are common on the Bering Sea Coast.

Section 3. Haplophylla Tzvel. sect. nova in Addenda, XXV, 880.—
Apical appendages of anthers oblong, obtuse, widely rounded;
receptacle less often hairy; achenes slippery when soaked. Plants with
relatively thin, more or less branched rhizome and entire, narrow-
linear leaves; pubescence of long and simple hairs.

A monotypic section.

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370

13. D. integrifolium (Richards.) Tzvel. comb. nova.—Chrysanthe-
mum integrifolium Richards. in Frankl. Narr. Journ. Bot. Appl (1823)
33.—C. scapigerum C.A.M. in herb.—Leucanthemum integrifolium
(Richards.) DC. Prodr. VI (1837) 45; Ldb. Fl. Ross. II, 540.—L.
algidum Fisch. ex Trautv. in Tr. Peterb. Bot. Sada, VIII, 2 (1883)
452.—Tanacetum integrifolium (Richards.) Sch. Bip. Tanacet. (1844)
35.—Ic.: Hook. Fl. bor.-amer. (1833) t. 109.

Perennial. Plant 3-15 cm high, with strongly branched rhizome,
usually forming dense mats, more or less covered with long simple
hairs. Stems erect or at base ascending, simple, with long, distant
hairs, with 1-3 (rarely 5) reduced leaves above base. Leaves subglabrous
to grayish from profuse pubescence of long simple hairs, lacking
punctate glands; basal leaves numerous, narrow-linear, to 2.0—2.5 cm
long and 1.5 mm wide, entire, obtuse or subacute; cauline leaves few,
similar to basal leaves. Capitula solitary on long, usually more or less
with apically woolly-cobwebby peduncles. Involucres 6—12 mm in dia
and 3-5 mm long, at base more or less woolly-pubescent; involucral
bracts with wide, blackish-brown, membranous border. Ligulate florets
white, corolla tube 1.0-1.5 mm long and limb 9-14 mm long and 2-
3 mm wide; corolla of tubular florets 2.0-2.5 mm long. Achenes 1.3—
2.0 mm long and about 0.5 mm wide, with slightly raised, subacute
upper border, but lacking corona. Flowering July to August. (Plate
XVI, Fig. 2).

Stony slopes, relatively dry parts of tundra, usually along coast.—
Arctic: Chukotka, Anadyr. General distribution: Arctic America (south
to British Columbia, east to Hudson Bay). Described from Arctic
America. Type in London.

Note. I did not see mature achenes of this species; the ovaries are
strongly flattened dorsally with prominent lateral ribs which often are
broadly winged in the outer florets of the capitulum; however, these
peculiarities apparently resulted from the deformation of the ovaries
during drying and are hardly evident in the mature achenes.

GENUS 1542. Tridactylina (DC.) Sch. Bip.!:?

Sch. Bip. Tanacet. (1844) 48.—Pyrethrum sect. Tridactylina DC.
Prodr. VI (1837) 61.

Capitula single or 2—30, solitary at apices of stem and its lateral
branches, sometimes forming compound corymb, heterogamous, with

'Treatment by N.N. Tzvelev.
From the Greek words treis—three, and dactylos—finger; named for the palmately
three-lobed leaves.

390

371

10-25 sterile (with rudiment of pistil), peripheral ligulate florets in
single row and numerous bisexual tubular disk florets. Involucres
saucer-shaped, 12-25 mm in dia and 5-7 mm long; involucral bracts
herbaceous, imbricate, in 3 irregular rows, lanceolate or lanceolate-
linear, subobtuse, with wide, blackish-brown membranous border; inner
bracts usually not more than 1.5 times as long as outer bracts.
Receptacle strongly and somewhat conically convex, plump, glabrous,
finely and weakly punctate-tuberculate, distinctly alveolar. Corolla of
ligulate florets yellow or yellowish-white, with strongly flattened, but
narrow-winged tube 1.5—2.0 mm long and linear or oblong-linear, limb
7-16 mm long; corolla of tubular disk florets yellow, 2.4-3.0 mm
long, with tube distinctly and rather abruptly widened in upper half,
with 5 lanceolate-deltoid, weakly bent teeth, a fifth to a fourth as long
as tube. Filaments of stamens apically thickened, anthers lacking distinct
basal appendages, but with lanceolate, subobtuse apical appendages;
pollen grains globose, spinulose. Style[stigma] bifid, style branches
linear, truncate. All achenes similar (but not developing in ligulate
florets), glabrous, 2.4—3.0 mm long and about 0.7 mm wide, more or
less terete, narrowed toward base, with (4) 5(6) indistinctly longitudinal
ribs rising at apex as very short (to 0.2 mm long), subobtuse teeth,
with very loose pericarp. Weakly hairy (pubescence of short, simple
and bifid hairs), annual plants with tap root, erect, simple or branched
stem, and alternate trilobate or tripartite leaves.

A monotypic genus.

Note. The genus Tridactylina shows a clear and rather close affin-
ity with the genus Dendranthema (DC.) Des Moul., being, by compari-
son with the latter, considerably more advanced in its evolution (tran-
sition to annual life cycle, reduction of pistils in the ligulate florets),
but at the same time retaining relatively prominent longitudinal ribs on
the achenes (though this character may be secondary).

1. T. kirilowii (Turcz.) Sch. Bip. Tanacet. (1844) 49; P. Popov,
in Sb. Akad. Sukachevu (1956) 450.—Pyrethrum kirilowii Turcz. in
DC. Prodr. VI (1837) 61 and in Bull. Soc. Nat. Mosc. I (1838) 94;
Ldb. FI. Ross. Ii, 558; Turcz. Fl. baic.-dahur. II, 47.

Annual. Relatively weakly hairy plant, 6-35 cm high, with short
tap root and erect; stem simple or almost from base more or less
branched. Leaves dull-green, less often hairy (to subglabrous), punctate-
glandular, more or less deeply, 3 (sometimes some 5)-lobed or divided
in upper part, gradually narrowed toward base into short, broadly
winged petiole, lacking stipules; basal leaves withering early, with
longer petiole; cauline leaves numerous, to 5 cm long, uppermost often
undivided and entire. Capitula solitary or 2—30, solitary at apices of

372

Plate XVII.
Tridactylina kirilowii (Turcz.) Sch. Bip., habit, corolla of ligulate florets,

391

1

corolla of tubular floret, achene; 2 — Ajania kokanica (Krasch.) Tzvel., habit,

corolla of tubular floret, achene.

393

373

stem and its leafy branches, sometimes forming lax irregular corymb.
Involucres 12—25 mm in dia and 5—7 mm long; involucral bracts with
rather wide, blackish-brown, membranous border; inner bracts 1.5 times,
less often 2 times as long as outer bracts. Corolla of ligulate florets
yellow or yellowish-white(?), tube and 1.5—2.0 mm long and limb 7-
16 mm long; corolla of tubular florets 2.4—3.0 mm long. Achenes 2.4—
3.0 mm long, lacking true corona, with very short (to 0.2 mm long),
subobtuse teeth as many as ribs. Flowering July to September. (Plate
XVII, Fig. 1).

Gravelly and sandy banks of rivers and lakes, sometimes as weed
along roads, old fields; up to middle mountain zone.—Eastern Siberia:
Angara-Sayans (eastern part), Dauria. General distribution: Probably
found in northern Mongolia. Described from area of Lake Baikal. Type
in Leningrad.

GENUS 1543. Brachanthemum DC.!:?

DC. Prodr. VI (1837) 44.—Chrysanthemum sect. Argyranthemum
(Webb ex Sch. Bip.) Benth. and Hook. f. Gen. pl. (1876) 426 p. p.;
Pflanzenfam. IV, 5 (1889) 278 p. p. — C. sect. Argyranthemum subsect.
Brachanthemum (DC.) Ling in Contr. Inst. Bot. Nat. Ac. Peip. III
(1935) 476.—C. sect. Brachanthemum (DC.) B. Fedtsch. Rast. Turk.
(1915) 736.

Capitula solitary, at apices of stem and its leafy branches (if present)
or rather numerous (to 20) and then often in more or less compact, not
always regular corymbose inflorescence, heterogamous, with 1-15
pistillate peripheral ligulate florets in single row, very rarely entirely
lacking (and then capitula heterogamous, with few pistillate tubular
florets) and numerous bisexual tubular disk florets. Involucres goblet-
shaped or narrow goblet-shaped (cup-shaped), 4-7 mm in dia and 4—
6 mm long; involucral bracts coriaceous-herbaceous, imbricate, in 4 or
5 irregular rows; outer bracts broadly ovate to oblong-ovate, obtuse,
with narrow, light colored or brownish membranous border, inner
oblong or oblong-linear, 2 or 3 times as long as outer bracts, with wide
(particularly in upper part) membranous border. Receptacle very weakly
punctate-tuberculate, distinctly alveolate, strongly convex, obtuse-
conical, glabrous (rarely subglabrous) or almost flat, covered with short
flexuous hairs. Corolla of ligulate florets yellow or white, with dorsally

'Treatment by N.N. Tzvelev.
2From the Greek words brachys—short, and anthemos—flower; for the relatively
short ligulate florets.

394

374

strongly flattened, but not winged, tube 1.2-1.8 mm long and limb
oval or oblong, 1.2—8.0 mm long; corolla of tubular disk florets yellow,
2-3 mm long, with narrow tube in lower half quite strongly and abruptly
broadened in upper half, with 5 more or less curved, lanceolate-
triangular teeth 2/9-1/3 as long tube. Filaments of stamen thickened in
upper part; anthers lacking distinct basal appendages, but with lanceolate
or ovate-lanceolate, subobtuse apical appendages; pollen grains globose,
spinulose. Style[Stigma] bifid; style branches linear, truncate. All
achenes similar, glabrous, 1.4—2.5 mm long and about 0.5—0.6 mm
wide, almost terete; narrowed toward base, with 5 (rarely to 7) veins,
indistinct and almost not raised as ribs (according to M.I. Savchenko
in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XI, 1949, p. p. 201-
207—only when dry longitudinal folds of pericarp as though ribs are
formed on the surface of achenes), with pericarp becoming slippery
when soaked (muciferous cells in irregular rows allover the surface of
achenes); pappus entirely absent. Semishrubs, more or less covered
with simple and bifid hairs (usually as fine tomentum), with stems
woody to a considerable height and alternate, pinnately or palmately
cut leaves.

Out of the eight species of the genus occurring in the stony steppes
and semideserts of Soviet Central Asia, from the Altai in the north to
the Tien Shan in the south, five are found in the USSR.

Type of genus: Brachanthemum fruticulosum (Ldb.) DC.

Note. Brachanthemum is very closely linked with the genus Den-
dranthema (DC.) Des Moul., through section Dendranthemopsis
(Tzvel.), which may also be treated as a separate genus. Probably
other intermediates between these genera will be found in the future in
the vast territory of China, but at present, particularly in the USSR,
both these genera can easily be delimited.

1. Corolla of ligulate florets white, with oblong ligule 6-8 mm
long; receptacle almost flat, short-hairy, (Altai) (sect.
Deridrantheriopsis AR YO. 226 A Ea
INI Ree 1. B. baranovii (Krasch. and Poljak.) Krasch.

+ Corolla of ligulate florets yellowish-white or yellow, with oval or
broadly oval ligule 1.2-3.0 mm long; receptacle strongly convex,

glabrous or sub-glabrous (sect. Brachanthemum) .............1:0c000008 2
2. Most leaves palmately cut into 3 linear-subulate lobes, less often 4
or 5 lobes, at base relatively approximate ..............::ccssessseeseeeees 3h
+ Most leaves pinnatisect into 4—7 linear-subulate lobes, only
uppermost leaves palmately 3-parted :n0.. 2.150 epce.-scheynceetecesnaguare 4.

3. Ligulate florets 1-6, ligule 1.2-2.0 mm long, sometimes entirely
absent; capitula solitary, less often 2 or 3 on stem ..............0
dein ddalp sus Bgated wheat ah gales @ sca seen salam ae naEn 5. B. kasakhorum Krasch.

395

375

+ Ligulate florets 3-12, ligule 2.2-3.0 mm long; capitula 1-20 on
stem, usually aggregated in COryMDS .............::ccssccsssceesceeseeeeeeeeteeees
Ree) ESE Ee eee eres 4. B. fruticulosum (Ldb.) DC.

4. Ligulate florets 5-15; involucres 5—7 mm in dia; capitula solitary,
less often 2-5 on stem, on long upright peduncles ...........
ie ee tar keel. cltion eis 3. B. kirghisorum Krasch.

+ Ligulate florets 1-8; involucres 4—5 mm in dia; capitula (1)3—10(15)
on stem, 2-4 capitula usually crowded at stem, apex, rest on long
Supe ge) coal) (0) i as es nee mee 2. B. titovii Krasch.

Section 1. Dendranthemopsis Tzvel. sect. nova in Addenda, XXV,
880.—Brachanthemum ser. Pyrethroides Krasch. in Bot. Mat. Gerb.
Bot. Inst. Akad. Nauk SSSR, XI (1949) 196, diagnosis in Russian.—
Involucres goblet-shaped, outer involucral bracts a third to a half as
long as inner; receptacle almost flat, covered with short flexuous hairs;
ligulate florets white, ligules 6-8 mm long; apical appendages of anthers
ovate-lanceolate, subobtuse.

A monotypic section.

1. B. baranovii (Krasch. and Poljak.) Krasch. op. cit. 196; Kryl.
Fl. Zap. Sib. XI, 2756.—Pyrethrum baranovii Krasch. and Poljak. in
Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, I (1933) 175.—Ic.: Krasch.
op. cit. 187, fig. 2.

Perennial. Semishrub, 10-35 cm high, with thick, woody, many-
headed root, grayish from fine tomentum of appressed simple and bifid
hairs. Stems usually numerous, woody to considerable height, erect,
densely leafy, branched mainly in upper half. Leaves grayish-green
from fine tomentum, with inconspicuous punctate glands, 2.0-2.5 cm
long and 1.0—1.5 cm wide, with rather long (almost as long as lamina),
very narrow-winged petiole, at base lacking stipules but often with 2
narrow-linear stipules in axillary buds; broadly ovate, pinnately cut up
to rather narrow winged rachis, with (1)2—3(4) narrow-linear (to 2 mm
wide), obtuse, lateral lobes on each side. Capitula usually numerous,
solitary on rather long (to 5-6 cm) peduncle at apices of stem and
branches, usually in compound corymb or corymbose-paniculate
inflorescence. Involucres 5-7 mm in dia and 5-6 mm long, more or
less grayish-tomentose; involucral bracts with rather wide brownish
membranous border. Ligulate florets 4-10, white, pistillate, but not
always fertile, corolla tube about 1.5 mm long and ligule oblong 6-8
mm long; corolla of disk florets 2.4-3.0 mm long. Achenes 1.4—1.8
mm long. Flowering August to September.

Stony slopes, up to middle mountain zone.—Western Siberia: Altai
(near mouth of Chuya River). Endemic. Described from Altai. Type in
Leningrad.

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376

Section 2. Brachanthemum—Brachanthemum Ser. Eubranchan-
thema Krasch. op. cit. 196 and ser. Procumbentia Krasch. ibid. 200;
diagnosis in Russian.—Involucres narrow-goblet-shaped, outer
involucral bracts 2/5—1/3 as long as inner; receptacle strongly convex,
obtusely conicle, glabrous, less often with occasional hairs; ligulate
florets yellowish-white or yellow, with ligule 1.2-3.0 mm long; apical
appendages of anthers lanceolate, subobtuse.

Type of section: type of genus.

Note. All species of this section are closely related and replace
each other with ecogeographical races of the same series.

2. B. titovii Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XI (1949) 196.—Ic.: Krasch. Ibid. 193, fig. 5.

Perennial. Semishrubs, 10—35 cm high, with many-headed woody
root, finely grayish-tomentose with appressed simple and bifid hairs.
Stems usually several, arising from base of plant, woody to considerable
height, erect, densely leafy, branched mainly in upper half. Leaves
grayish-green from fine tomentum, with inconspicuous punctate glands,
to 2.0—-2.5 cm long and 1.5 cm wide, with rather long petiole thickened
at base, lacking stipules but sometimes with pseudostipules (of axillary
bud), broadly ovate, pinnately cut up to very narrow-winged rachis,
with (1)2—3(4) linear, subulate, thick lobes on each side having short
cartilaginous cusp; upper leaves often palmately 3-parted, uppermost
undivided. Capitula (1)3—10(18) on single stem, of which 2—4 usually
crowded on short peduncles at stem apex, rest at apices of elongated
lateral branches in lax irregular corymb. Involucres 4-5 mm in dia and
4.5-6.0 mm long, more or less finely tomentose; involucral bracts with
rather wide, light-colored or brownish, membranous border. Ligulate
florets 1-8, yellowish-white, corolla tube 1.2—-1.8 mm long and ligule
2.2—3.0 mm !ong, broadly oval; corolla of disk florets 2-3 mm long.
Achenes 2.0-—2.5 mm long. Flowering July to August.

Stony and rubbly slopes (particularly on red conglomerates) of
middle mountain zone.—Soviet Central Asia: Tien Shan, Ketmen,
Trans-Ili Alatau ranges. General distribution: Dzhungaria-Kashgaria
(Chinese Dzhungaria). Described from Trans-Ili Alatau. Type and
isotypes in Leningrad.

3. B. kirghisorum Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, IX (1946) 171.—Ic.: Krasch. ibid. XI (1949) 191, fig. 4.
Perennial. Semishrubs, 5-25 cm high, with many-headed woody
root, finely grayish-tomentose with appressed simple and bifid hairs.
Stems rather numerous, mainly ascending at base, woody to considerable

397

377

height, simple or with few (1-5) elongated lateral branches. Leaves
like preceding species. Capitula solitary, less often 2—5 on single stem,
on long upright peduncles. Involucres 5—7 mm in dia and 4.5-6.5 mm
long, more or less finely tomentose; involucral bracts with rather wide
membranous border. Ligulate florets 5—12(15), yellowish-white(?),
corolla tube 1.2—-1.8 mm long and ligule 2.2-3.0 mm long, broadly
oval; corolla of disk florets 2-3 mm long. Achenes 1.8—2.4 mm long.
Flowering August to September.

Rubbly and stony slopes, gravel beds above 1,000 m. —Soviet
Central Asia: Tien Shan (central part). General distribution:
Dzhungaria-Kashgaria (eastern Tien Shan). Described from Tien Shan.
Type in Leningrad.

4. B. fruticulosum (Ldb.) DC. Prodr. VI (1837) 44; Ldb. FI.
Ross. II, 540; O. and B. Fedtsch. Perech. Rast. Turk. IV, 183; Kryl.
Fl. Zap. Sib. XI, 2755.—Chrysanthemum fruticulosum Ldb. Fl. Alt.
IV (1833) 117; B. Fedtsch. Rast. Turk. 736.—Ic.: Ldb. Ic. pl. FI.
Ross. impr. Alt. V (1834) t. 495; Krasch. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XI, 189, Fig. 3.

Perennial. Semishrub, 5-20 cm high, with many-headed woody
root, finely grayish-tomentose with appressed simple and bifid hairs.
Stems usually rather numerous, ascending at base or erect, woody to
a considerable height, branched mainly in upper half. Leaves grayish-
green from fine tomentum, with indistinct punctate glands, to 1.5(2.0)
cm long and 1.5 cm wide, with rather long petiole thickened at base,
lacking stipules, but sometimes with pseudostipules (of axillary bud),
almost circular, palmately 3-parted, with linear-subulate, somewhat thick
lobes having short cartilaginous cusp at apex, sometimes some leaves
pinnately 4—S-parted (and then bases of lobes usually approximate);
uppermost leaves undivided, linear. Capitula (1)3—15(20) on single
stem, aggregated in lax, not always regular, simple or compound
corymbs. Involucres 5-6 mm in dia and 4.5-6.0 mm long, more or
less finely tomentose; involucral bracts with light or brownish
membranous border, considerably wider in inner bracts. Ligulate florets
(3)4—10(12), yellowish-white as in preceding species; corolla of disk
florets 2-3 mm long. Achenes 1.8—2.2 mm long. Flowering August to
September.

Rubbly and stony slopes; up to middle mountain zone.—Western
Siberia: Altai (southern part); Soviet Central Asia: Lake Balkhash
Region (eastern part). General distribution: Dzhungaria-Kashgaria,
Mongolia (western part). Described from eastern Kazakhstan (Chingiz
Mountains). Type and isotypes in Leningrad.

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378

5. B. kasakhorum Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, IX (1946) 170.—Ic.: Krasch. ibid. 171, fig. 8.

Perennial. Semishrub, 5-15 cm high, with many-headed woody
root, finely grayish-tomentose with appressed simple and bifid hairs,
sometimes weakly tomentose. Stems usually rather numerous, ascending,
simple, less often with 1-2 branches. Leaves dull- or grayish-green
from fine tomentum, with indistinct punctate glands, to 1(1.5) cm long
and 1.3 cm wide, with rather long petiole thickened at base, almost
circular; all leaves (except uppermost undivided) palmately 3-parted,
with somewhat shorter than in preceding species, but linear-subulate
lobes. Capitula solitary, less often 2-3 on single stem (but numerous
on same plant). Involucres 4.5—6.0 mm in dia and 4-5 mm long, more
or less finely tomentose, sometimes subglabrous; involucral bracts with
lighter or brownish membranous border considerably wider in inner
bracts. Ligulate florets 1-6, sometimes entirely absent (replaced by
pistillate tubular florets), yellow, corolla tube about 0.8-1.0 mm long
and broadly ovate ligule 1.2-2.0 mm long; corolla of disk florets about
2.5 mm long. Achenes 1.6—2.0 mm long. Flowering August to
September.

Stony slopes, steppes; up to lower mountain zone.—Western
Siberia: Upper Tobol (southeastern part); Soviet Central Asia: Aralo-
Caspian Region (northeastern part); Lake Balkhash Region
(northwestern part). Endemic. Described from Akmolinsk [now
Tselinograd] Region (Kazakh Melkosopochnik [area of low, rounded,
isolated hills]). Type and isotype in Leningrad.

GENUS 1544. Ajania Poljak.!?

Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVII
(1955) 419.—Chrysanthemum sect. Pyrethrum sub-sect. Dendranthema
(DC.) Kitam. in Acta Phyt. et Geobot. IV (1935) 477 and Compos.
japon. II (1940) 350 p. p.—Tanacetum auct. p. p. non L.

Capitula usually numerous (2-60) aggregated at apex of stem in
lax or compact corymbs, very rarely (in species of series Tibeticae)
solitary, heterogamous but lacking ligulate florets, with relatively fewer
(1-15) pistillate peripheral tubular florets, in one row, and considerably
large number (15-80) of bisexual, tubular disk florets. Involucres
goblet-shaped, 2.5—-10.0 mm in dia and 3—7 mm long; involucral bracts

'Treatment by N.N. Tzvelev.
*Named after the Ayan port on the Sea of Okhotsk, the neighborhood from which
the species Ajania pallasiana was described.

399

oe

herbaceous or coriaceous-herbaceous, imbricate, in 3—4 irregular rows;
outer bracts linear-lanceolate to broadly ovate, obtuse or subacute,
inner oblong-linear to oblong-ovate, obtuse or subobtuse, |.5—3 times
as long as outer bracts; all bracts with rather wide, membranous border,
often (in inner bracts) apically broadened like appendage. Receptacle
strongly or moderately convex, often obtusely conical, glabrous, very
weakly punctate-tuberculate, usually distinctly alveolar, more or less
filled. Corolla of pistillate tubular florets mainly similar to that of disk
florets but usually smaller, more or less dorsally compressed, with
2-4 stiff teeth; corolla of disk florets yellow, 1.5—3.0 mm long, their
tube quite strongly and abruptly broadened in upper half, with 5
deltoid, more or less curved teeth a sixth to a fifth as long as tube.
Filaments of stamens thickened in upper part, anthers lacking
distinct basal appendage, but with broadly lanceolate, acute or
subobtuse apical appendage; pollen grains globose, spinulose.
Style[stigma?] bifid; style branches linear, truncate. Achenes similar,
glabrous, 1.0-2.2 mm long and 0.4-1.0 mm wide, almost terete,
narrowed toward base, with 4—6 longitudinal veins as very indistinct,
not always prominent ribs, pericarp lacking secretory canals but usually
with muciferous cells in irregular rows almost allover achene; pappus
entirely absent.

Perennial herbs or semishrubs, more or less covered with appressed
bifid hairs, sometimes mixed with simple hairs, lacking telescoped (as
rosettes of basal leaves) vegetative shoots; stem erect but basally often
ascending, branched usually only at base and in inflorescence, densely
leafy with alternate, usually incised, rarely (as in A. kokanica) entire
leaves.

The genus contains about 25 species distributed in China, Japan,
Mongolia, Korea, Afghanistan, and the northern part of India; of these,
nine species are found in parts of the Soviet Union bordering these
countries.

Type of genus: Ajania pallasiana (Fisch. ex Bess.) Poljak.

Note. The group of species belonging to this genus is very closely
linked with the genus Dendranthema (DC.) Des. Moul. and is
distinguished from the latter only by the absence of peripheral ligulate
florets and by smaller capitula on the average. It is quite probable that
in treating the numerous species belonging to these two genera over
the vast territory of China, they generally cannot be distinguished as
separate genera or the basic character distinguishing them will have to
be, not the presence or absence, but the color of the ligulate
florets, (analogous to the genera Tanacetum L. and Pyrethrum Zinn.).
At present, however, having neither the time nor sufficient material for
such a treatment, I prefer to retain both genera, which are well

400

380

distinguished over the territory of the USSR, as separate entities.
Apparently, the affinity between the genera Ajania and Brachanthemum
DC. is just as close, but this, too, can be examined more thoroughly
only by studying materials of these two genera from China and
Mongolia.

The evolution of the genus Ajania from ancestral forms similar to
the genus Dendranthema in the direction of adaptation to habitats in
vast expanse of steppes and deserts of Central Asia has resulted in a
considerable morphological similarity of many species of Ajania with
species of the genus Artemisia L. (possibly representing another line
of evolution from these same ancestral forms of the Dendranthema
type). It was not by accident that P.P. Poljakov (op. cit.), the author
of the genus Ajania, mistakenly referred six species of arctic
wormwoods (Artemisia glomerata Ldb., A. senjavensis Bess., etc.) to
this genus. Despite their superficial resemblance to Ajania pallasiana,
their compound inflorescence is transitional from capitate to racemose
but not to corymbiform; the appendages of the anthers in most
species are attenuate-acuminate (which is not so in species of the
genus Ajania; and the pollen grains of all these species (according to
A.A. Kuprianova) differ so sharply from those of Ajania species,
not only in the absence of spinules on the surface but also in
the structure of the tectum, that the similarity of the indicated
wormwoods to the genus Ajania undoubtedly should just be considered
convergence.

The genus Ajania is divided into several entirely independent series
containing very close ecogeographical races, but can hardly be divided
into any larger taxa such as sub-genera or sections.

1. Leaves undivided, with sharp teeth throughout .................eeeeseseeees
vaeseslaed, bed ae? er Se 6. A. kokanica (Krasch.) Tzvel.

+ Leaves more or less divided, less often 3—5-lobed .................... ai
2. Leaves green and glabrous or weakly hairy above, whitish- or
grayish-tomentose beneath. Far East ............:cssseeseceeesneeeeneeeeenees oF
+ Leaves concolorous on both sides, grayish-green from profuse
pubescence. Altai and Soviet Central Asia .............eeseeeeeeeeeeee 4.

3. Plants herbaceous, leaves tomentose beneath from appressed bifid
hairs; outer involucral bracts linear-lanceolate ..............:::cccccccceeeeseens
Hitesesdhsaets. Gah POPU ea 2. A. pallasiana (Fisch. ex Bess.) Poljak.

+ Stem woody at base; leaves tomentose beneath from flexuous simple
hairs; outer involucral bracts broadly Ovate ............cssesessseseeesens
nsfiduni ope h tated op Aas: cnale anon ate teh. een 1. A. manshurica Poljak.

4. Plants (10)15-60(90) cm high, with relatively small (2.5-4.5 mm
wide) capitula aggregated in compound corymDs .............0:.see++ee 5.

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381

+ Plants 2-15 cm high, with larger (S—10 cm wide) capitula solitary
or 2-8 aggregated in COMpact COTYMDS ...........ccccessceseseeerseeeeeees i.
5. Plants 10-30 cm high; stems many from base, with thick woody
caudex; capitula 4—25 on single stem, relatively large (3.0-4.5 mm
in dia and 4-5 mm long). Altai, Tarbagatai, Kazakh
alkeonarochitihe”” v.2i.s;..kvisil, obs 4. A. fruticulosa (Ldb.) Poljak.
+ Plants 15-90 cm high; capitula 20-80 on single stem not large
(2.54.0 mm in dia and 3.04.5 mm long), Tien Shan, Pamiro-Alai
er Re RU) SOR ICOV ENT Rhy AMOR Seen Tpit sie YG eee eu Een ye See area PE Eee 6.
6. Stem strongly branched at base, woody to considerable height,
middle and lower cauline leaves usually twice ternately divided.
Pattie. eies cack 5. A. gracilis (Hook. f. and Thoms.) Poljak.
+ Stem weakly branched at base (often solitary), almost not woody;
middle and lower leaves usually bipinnate, with 5-7 lobes................+.
Pee ee eens cevsiusinevataesoinena 3. A. fastigiata (Winkl.) Poljak.
7. Middle and lower cauline leaves usually pinnately cut (or parted)
into 5-9 lobes, in turn, all or some 3—5-parted or lobed. Pamir ..
Kebividsdssvnased RU ISEGAT SS. 9. A. tibetica (Hook. f. and Thoms.) Tzvel.
+ Middle and lower cauline leaves palmately or almost 3-5 pinnately
divided (or lobed); their lobes undivided or some 2—3-parted. Central
Bien SheeGl. sab. 302 ud alie. eoetwanl.ianade. geet SOL. 8.
8. Plants 2-7 cm high, with thick, woody caudex; capitula 7-10 mm
Bpidiaaitem salitatyrdss nie bles. obi age ek...
Sh te . Kn 7. A. scharnhorstii (Rgl. and Schmalh.) Tzvel.
+ Plants 5-15 cm high, with thinner, not always woody root; capitula
5-8 mm in dia, always aggregated in COryMDS ............::csccesseeseeeeeeees
alinwede sugia, 2uciiialy .botid.2. gnieiai 8. A. trilobata Poljak.

Series 1. Variifoliae Tzvel_—Stem woody at base; leaves pinnately
parted, glabrous above, densely whitish-tomentose beneath with flexuous
simple hairs; capitula 10-25, aggregated in compact compound corymb.
Involucres 3.5—5.0 mm in dia; outer involucral bracts a half to two-
thirds as long as inner bracts.

1. A. manshurica Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVII (1955) 420.—Chrysanthemum variifolium var.
ramosum Chang in Sinensia V (1934) 163.—Ic.: Poljak. op. cit. 421,
fig. 2.

Perennial. Plant 15-60 cm high, base semishrubby, with thick
woody root. Stem branched in lower woody part, erect, densely leafy,
more or less covered with appressed bifid and simple hairs. Leaves
thick, subcoriaceous, weakly revolute, lacking distinct punctate glands,
glabrous and green, densely whitish-tomentose above with flexuous

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382

simple hairs beneath; lower and middle cualine leaves to 4 cm long
and 2.3 cm wide, with rather long, winged petiole, lacking auricles,
more or less rhombic, 3-5 pinnately parted, with lanceolate or
lanceolate-linear, 1.2-3.0 mm wide lobes, cuneately narrowed toward
base; upper cauline leaves smaller, usually ternate. Capitula 10-25,
with 20-40 florets, aggregated at stem apex in compound corymbs,
peduncles 0.5—6.0 mm long, more or less hairy. Involucres 3.5—5.0
mm in dia and 4.0-5.5 mm long, usually hairy near base to subglabrous;
involucral bracts coriaceous-herbaceous, outer broadly ovate, two-fifths
to a half as long as oblong-ovate inner bracts; all bracts with rather
wide, brownish membranous border. Corolla of tubular disk florets
2.2—2.6 mm long; peripheral pistillate florets usually smaller, 3-8 on
each capitulum. Achenes 1.8-2.2 mm long, with unevenly raised
subobtuse upper border, lacking corona. Flowering August to
September.

Rocks, stony slopes; up to upper mountain zone.—Far East: Ussuri
(basin of Suifun River). General distribution: China (northeastern part),
Korea. Described from Far East. Type in Leningrad.

Note. It is distinguished from the very similar species A. variifolia
(Chang) Tzvel. comb. nova (=Chrysanthemum variifolium Chang, op.
cit. 161) from Shansi Province only by the wider lobes of the leaves
and the stems that usually branch near the base.

It belongs to the group of Ajania species that are very close to the
genus Brachanthemum DC.

Series 2. Pallasianae Tzvel.—Stems herbaceous; leaves pinnately
parted or ternate, sometimes 3-lobed, glabrous above, less often hairy,
densely tomentose beneath with appressed bifid hairs; capitula (3)5-
50(60), aggregated in rather compact, compound corymb. Involucres
4-7 mm wide; outer involucral bracts not less than two-thirds as long
as inner bracts.

2. A. pallasiana (Fisch. ex Bess.) Poljak. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XVII (1955) 420.—Artemisia pallasiana Fisch.
ex Bess. in Nouv. Mém. Soc. Nat. Mosc. III (1834) 61; DC. Prodr. VI,
116, non A. pallasii Spreng. 1826.—Tanacetum pallasianum (Fisch.
ex Bess.) Trautv. and Meyer, Fl. ochot. (1856) 55, t. 27; Maxim. Prim.
Fl. Amur. (1859) 163.—Pyrethrum pallasianum (Fisch. ex Bess.)
Maxim. in Mel. biol. VIII, 2 (1872) 514.—Chrysanthemum pallasianum
(Fisch. ex Bess.) Kom. in Tr. Peterb. Bot. Sada, XXV, 1 (1907) 645;
Kom. and Alis. Opred. Rast. Dalnevost. Kr. II (1932) 1028.—Ic.:
Trautv. and Meyer, loc. cit.; Poljak. op. cit. 421, fig. 1.—Exs.: GRF
No. 3273.

383

403 Plate XVIII.
Habit, corolla of tubular floret, achenes: 1 — Ajania scharnhorstii (Rgl. and
Schmalh.) Tzvel.; 2— Hippolytia darvasica (Winkl.) Poljak.

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384

Perennial. Plant 15—60 cm high, with thick, more or less branched
rhizome, more or less covered with appressed bifid hairs. Stems erect,
solitary or few, densely leafy, branched only in inflorescence. Leaves
not always with distinct punctate glands, glabrous above or less often
hairy, grayish- or whitish-tomentose beneath with appressed bifid hairs;
lower and middle cauline leaves to 8 cm long and 4 cm wide, with
rather long (but always shorter than lamina), winged petiole, at base
often with oblong or linear lobes of axillary shoot but often adnate
with base of petiole, oblong to more or less rhombic, ternate or pinnately
parted, sometimes pinnately lobed, with few linear or linear-lanceolate
pinnate or lobes, often in turn more or less lobed or irregularly toothed,
uppermost cauline leaves strongly reduced and less divided (often 3-
lobed or even undivided). Capitula (3)5-50(60) with 30-80 florets,
aggregated at stem apex in rather compact, compound (less often simple)
corymb, their peduncles 2-15 mm long, usually with strongly reduced
subtending leaves, appressed-hairy. Involucres 4-7 mm in dia and 4—
6 mm long, more or less hairy to subglabrous; involucral bracts
herbaceous, outer bracts linear-lanceolate, not less than two-thirds as
long as oblong and linear-oblong inner bracts; inner bracts with rather
wide, light-colored or brownish membranous border. Corolla of tubular
disk florets with 2.5-3.2 mm long; few peripheral pistillate florets
usually smaller. Achenes 1.2-1.8 mm long and about 0.8 mm wide,
with somewhat raised obtuse upper border, lacking corona. Flowering
July to September. (Plate XXXI, Fig. 2).

Stony slopes, rocks; up to middle mountain zone.—Far East:
Okhotsk (near Ayan), Kamchatka(?), Uda River Area, Zeya-Bureya
(eastern part), Ussuri, Sakhalin. General distribution: China
(northeastern part), Korea. Described from Okhotsk seacoast. Type in
Leningrad.

Note. A highly polymorphic species, apparently breaking up into
several ecogeographical races, distinguished mainly by the shape of
the leaves and, to a lesser extent, by the size of the capitula and the
shape of the involucral bracts. For example, in the northern Sikhote-
Alin District near the city of Sovetskaya Gavan we find specimens of
A. pallasiana with wide leaves, which are trilobate only in the upper
third and have well developed auricles at the base of the petioles,
while from other areas we know specimens with bipinnate leaves without
auricles.

Series 3. Fruticulosae Tzvel.—Stem woody at base; leaves more
or less divided or incised, concolorous on both sides, grayish from
profuse pubescence of appressed bifid hairs; capitula (4)10-—60(80) on
single stem, aggregated in lax or compact compound corymb. Involucres

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385

2.5-4.5 mm in dia; outer involucral bracts a third to a half as long as
inner bracts.

3. A. fastigiata (Winkl.) Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk, XVII (1955) 428.—Artemisia fastigiata Winkl. in Tr.
Peterb. Bot. Sada, XI, 12 (1891) 373; O. and B. Fedtsch. Perech. Rast.
Turk. IV, 200.—Tanacetum fastigiatum (Winkl.) Krasch. in Spisok
Rast. Gerb. Fl. SSSR, XI (1949) 40.—Exs.: GRF No. 3275.

Perennial. Plant 30-90 cm high, with thick, woody root, grayish
from profuse pubescence of appressed bifid hairs. Stems at base almost
not woody, usually few (sometimes solitary), erect, densely leafy,
usually more or less branched only above. Leaves grayish-green, with
numerous punctate glands; basal leaves withering early; cauline leaves
to 4-5 cm long and 3-4 cm wide, with very narrow-winged petiole and
linear auricles; wide deltoid-oval, bipinnate (or pinnately cut) up to
narrow-winged rachis, with 5-7, less often 3 lobes; terminal lobes
linear or lanceolate, to 2 mm wide, obtuse or subacute; upper cauline
leaves reduced and less divided, often subsessile. Capitula with 15-25
florets, 20—80 on single stem, aggregated in rather compact, compound
corymbose inflorescence with multiple branched peduncles, peduncles
(terminal branches of corymb) to 8 mm long, usuaily with very small
linear subtending leaves. Involucres 2.5-4.0 mm in dia and 3.0--4.5
mm long, more or less tomentose near base; involucral bracts
herbaceous, outer broadly lanceolate and ovate, with narrow
membranous border, inner bracts 2—3 times as long as outer bracts,
oblong-obovate, with wide, light-colored or brownish, membranous
border appendiculately broadened at apex. Corolla of tubular disk florets
1.8-2.5 mm long; few peripheral pistillate florets usually smaller.
Achenes 1.0-1.5 mm long and about 0.4 mm wide, lacking corona.
Flowering August to September.

Stony slopes, steppes, scrubs, at 1,500 to 3,000 m. — Soviet Central
Asia: Dzhungaria-Tarbagatai (Dzhungarian Alatau), Tien shan, Pamiro-
Alai Region (near Pamir). General distribution: China (western part),
Afghanistan. Described from area between Alai and Peter the First
ranges. Type in Leningrad.

4. A. fruticulosa (Ldb.) Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVII (1955) 428.—Tanacetum fruticulosum Ldb.
Ic. pl. Fl. Ross. I (1829) 10, t. 38; DC. Prodr. VI, 129; Ldb. Fl. Ross.
II, 603; O. and B. Fedtsch. Perech. Rast. Turk. IV, 204 p. p.; Kryl. Fl.
Zap. Sib. XI, 2758.—Artemisia athanasia Bess. in Bull. Soc. Nat.
Mosc. VII (1834) 24.—Pyrethrum athanasia (Bess.) Boiss. Fl. or. III
(1875) 353, quoad. nomen.—Chrysanthemum fruticulosum (Ldb.) B.

407

386

Fedtsch. Rast. Turk. (1915) 737. non Ldb. 1829.—C. neofruticulosum
Ling in Contr. Inst. Bot. Ac. Sc. Peip. III (1935) 482, nom. nov.—C.
athanasia (Bess.) B. Fedtsch. in herb.—Ic.: Ldb. 1. c.

Perennial. Plant 10-30 cm high, with thick, woody root, grayish
from profuse pubescence of appressed bifid hairs. Stems woody at
base, usually rather numerous, erect or basally ascending, densely leafy,
weakly branched only above. Leaves grayish-green, with numerous
punctate glands; basal leaves withering early; cauline leaves to 2-3 cm
long and 2.5 cm wide, with rather long (sometimes as long as lamina),
almost not winged petiole, usually with linear auricles, usually deltoid
roundish, ternate (or divided) up to quite narrow-winged rachis, less
often pinnately parted, lobes usually divided into 3—5 secondary lobes,
terminal lobe narrow-linear, to 0.8—1.0 mm wide, obtuse or subacute;
upper cauline leaves reduced and less divided, usually sessile or
subsessile. Capitula with 15-30 florets, 4-25 on single stem, aggregated
in compact, compound, corymbose inflorescence, peduncles to 10 mm
long, usually with linear subtending leaves at base. Involucres 3.0—4.5
mm in dia and 4—5 mm long, glabrous or subglabrous; involucral bracts
herbaceous, outer broadly lanceolate and ovate, a third to a half as
long as oblong-obovate inner bracts; all bracts with wide light or
brownish membranous border, appendiculately broadened at apex of
inner bracts. Corolla of tubular florets with 1.8-2.8 mm long; few
peripheral florets pistillate. Achenes 1.2-1.5 mm long and about 0.4
mm wide, lacking corona. Flowering August to September.

Stony and rubbly slopes, steppes; up to middle mountain zone.—
Western Siberia: Irtysh (southern part), Altai (southern part); Eastern
Siberia: Angara-Sayans (southwestern part); Soviet Central Asia: Lake
Balkhash Region, Dzhungaria-Tarbagatai (Tarbagatai). General
distribution: China (western part), Mongolia. Described from Altai.
Type and isotype in Leningrad.

5. A. gracilis (Hook. f. and Thoms.) Poljak. comb. nova. in litt.—
Tanacetum gracile Hook. f. and Thoms. FI. Brit. Ind. III (1882) 318.—
Chrysanthemum gracile (Hook. f. and Thoms.) B. Fedtsch. Rast. Turk.
(1915) 737, non Masf. 1881.—C. hookeri Kitamura in Acta Phyt. et
Geobot. XVII (1957) 34, nom. nov.

Perennial. Plant 15-45 cm high, with thick, woody root, grayish
from abundant pubescence of appressed bifid hairs; stems strongly
branched at base and woody to considerable height, usually rather
numerous, erect, densely leafy. Leaves grayish-green, with numerous
punctate glands; basal leaves withering early; cauline leaves to 2-3 cm
long and 2.5 cm wide, with rather long, almost wingless petiole, at
base with linear auricles or lacking them; their lamina usually deltoid

408

387

roundish, twice ternate up to narrowly winged rachis, terminal lobe
oblong to linear, to 1.5 mm wide, subobtuse; upper cauline leaves
reduced, subsessile, usually 3—5 lobed. Capitula with 15-25 flowers,
20-60 on single stem, aggregated in compact compound corymbs,
peduncles usually 1-5 mm, less often to 8 mm long, often with linear
subtending leaves. Involucres 2.5-3.0 mm in dia and 3-4 mm long,
subglabrous; involucral bracts herbaceous, with wide, light or brownish
membranous border appendiculately broadened at apex; outer bracts a
third to a half as long as inner bracts. Tubular florets with 1.5-2.0 mm
long corolla, few peripheral florets pistillate. Achenes 1.2-1.5 mm
long and about 0.5 mm wide, lacking corona. Flowering August to
September.

Rocks, stony slopes; above 3,000 m.—Soviet Central Asia: Pamiro-
Alai Region (Pamir). General distribution: India (northern part), China
(Tibet). Described from western Tibet. Type in London.

Note. Because of the absence of reliably identified material of this
species, I am not entirely convinced of the complete identity of the
Pamir specimens with the typical Tibetan specimens.

Series 4. Sedifoliae Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk. SSSR, IX (1946) 170.—Stems herbaceous; leaves undivided,
sharp-toothed, dull- or grayish-green from rather profuse pubescence
of appressed bifid hairs; capitula (1)2—10(12), aggregated in very
compact, simple or compound corymb. Involucres 4-6 mm in dia;
outer bracts usually not less than 2/3 as long as inner ones.

6. A. kokanica (Krasch.) Tzvel. comb. nova.—Tanacetum
kokanicum Krasch. in Bot. Mat. Gerb. Glavn. Bot. Sada, IV (1923)
7.—T. aphanassievi Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, IX (1946) 168.—Hippolytia aphanassievii (Krasch.) Poljak. in
Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVIII (1957) 290.

Perennial. Plant 15-40 cm high, with thick, woody root, dull- or
almost grayish-green from rather profuse pubescence of appressed bifid
hairs. Stems numerous, at base usually not woody, erect or basally
ascending; densely leafy, branched only in inflorescence. Leaves dull-
or grayish-green from rather profuse pubescence, with numerous
punctate glands, sessile, lacking auricles or with small auricles at base,
oblong-linear or oblong-lanceolate, up to 3 cm long and 0.8 cm wide,
long-attenuate and acuminate, sharp-toothed; basal and lower cauline
leaves withering early, subtending leaves strongly reduced, often up to
inflorescence. Capitula many-flowered, (1)2—10(12) on single stem,
aggregated in very compact, often globose, simple or compound corymb,
peduncles 0.5—15(20) mm long. Involucres 4-6 mm in dia and 4-5 mm

388

long, at base more or less hairy; involucral bracts herbaceous, with
relatively narrow, brownish or blackish-brown membranous border,
outer bracts lanceolate, acute, fewer, inner bracts usually not more
than 1.5 (rarely almost 2) times as long as outer bracts, linear-oblong,
subobtuse. Corolla of tubular florets with 1.8-2.4 mm long, few (1-5)
peripheral florets usually pistillate. Achenes about 1.6 mm long and
0.6 mm wide, lacking corona: Flowering August to September. (Plate
XVII, Fig. 2).

Rubbly and stony slopes, talus; above 2,500 m.— Soviet Central
Asia: Pamiro-Alai Region (Turkestan Range and western part of Alai
Range). Endemic. Described from Turkestan Range. Type and isotype
in Leningrad.

Note. Tanacetum aphanassievii Krasch., also described from the
Turkestan Range, hardly deserves to be treated as a separate species,
as its differences (somewhat weaker pubescence, slightly wider leaves
with much longer teeth, and, on the average, more compact
inflorescence) apparently are not hereditary but individual features
without any taxonomic significance, related to more or less shady or
more humid habitats.

Series 5. Tibeticae Tzvel.—Stems at base almost not woody; leaves
more or less pinnatipartite or 3-5 lobed, grayish from profuse
pubescence of appressed bifid hairs; capitula solitary at apices of stems
or 2—8 aggregated in rather compact, not always regular corymbs.
Involucres 5-12 mm in dia; outer involucral bracts a half to two-thirds
as long as inner bracts.

7. A. schanhorstii (Rgl. and Schmalh.) Tzvel. comb. nova.—
Tanacetum scharnhorstii Rgl. and Schmalh. in Tr. Peterb. Bot. Sada,
V, 2 (1878) 620; O. and B. Fedtsch. Perech. Rast. Turk. IV, 206.—T.
grigorjewii Krassn. in Scripta bot. Hort. Univ. Petrop. II, 1 (1887-
1888) 16; O. and B. Fedtsch. op. cit. 206.—Chrysanthemum
scharnhorstii (Rgl. and Schmalh.) B. Fedtsch. Rast. Turk. (1915) 738.—
C. grigorjewii (Krassn.) B. Fedtsch. op. cit. 738—Hippolytia
scharnhorstii (Rgl. and Schmalh.) Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVIII (1957) 289.

Perennial. Plants 2-7 cm high, with thick, woody, many-headed
root, grayish from profuse pubescent of appressed bifid hairs. Stems
numerous from base, herbaceous, erect to almost prostrate, densely
leafy, simple or branched only inflorescence. Leaves grayish or whitish
from profuse pubescence, with numerous punctate glands less distinct
because of pubescence; basal leaves withering early; lower and middle
cauline leaves to 1.3 cm long and 1.0 cm wide, with rather long (but

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usually shorter than lamina), narrow-winged petiole, lacking auricle at
their base, broadly ovate to reniform, usually cuneately narrowed toward
base, 3-5 palmately or pinnately parted, their terminal lobe ovate to
broadly linear, often distinctly broadened in upper half, obtuse and
often divided into 2-3 secondary lobes; upper cauline leaves usually
ternate or three-lobed, smaller. Capitula solitary, on more or less
tomentose, to 1 cm long peduncles, or 2-4 on single stem, aggregated
in compact, not always regular corymb. Involucres 7-10 mm in dia
and 6-7 mm long, finely tomentose near base; involucral bracts
herbaceous, outer bracts broadly lanceolate or lanceolate-ovate, with
narrow membranous border, inner bracts 1.5—2 times as long as outer
bracts, oblong or ovate, with very wide, brownish, less often light-
colored membranous border appendiculately broadened at apex. Corolla
of tubular florets 2.2-3.0 mm long, few (1-6) peripheral pistillate
florets. Achenes 1.6—2.2 mm long and about 0.6-0.8 mm wide, lacking
corona. Flowering July to August. (Plate XVIII, Fig. 1).

Stony and rubbly slopes, gravel beds; above 3,000 m.—Soviet
Central Asia: Tien Shan (central part). General distribution: China
(Tien Shan). Described from Central Tien Shan. Type in Leningrad.

Note. Like the following two species, this species is very closely
linked to the species of the series Fasciculatae Tzvel., being their
high-mountain derivative. Apparently, it was erroneously referred to
the genus Hippolytia Poljak. by P.P. Poljakov (op. cit.).

8. A. trilobata Poljak. sp. nova in Addenda, XXV, 880.

Plant 5—15 cm high, with root thinner than in previous species, not
always woody, grayish from profuse pubescence of appressed bifid
hairs. Stems numerous from base, herbaceous, erect or ascending,
branched only in inflorescence. Leaves like previous species but, on
the average larger and less dense. Capitula (1)3—8(10) on single stem,
aggregated in compact simple or compound corymb. Involucres (4.5)5—
7(8) mm in dia and 5.0—-6.5 mm long, more or less tomentose near
base; involucral bracts like in previous species. Corolla of tubular
florets 2.2-3.0 mm long, few (1-8) peripheral florets pistillate. Achenes
like in previous species. Flowering July to August.

Gravel beds, stony slopes, above 2,500 m.—Soviet Central Asia:
Tien Shan (central part). General distribution: China (Tien Shan).
Described from Central Tien Shan. Type (basin of Sarydzhas River,
slopes below Nansen Peak, height 2,950 m, 29.VII.1955, M.
Grudzinskii) in Leningrad.

Note. In relation to the previous species, it is, on the average, a
much lower mountain ecogeographical race confined to more wet
(usually gravelly) habitats and is linked to it by specimens having
capitula that are intermediate in size and number.

41

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390

9. A. tibetica (Hook. f. and Thoms.) Tzvel. comb. nova.—
Tanacetum tibeticum Hook. f. and Thoms. in Clarke, Compos. Ind.
(1876) 154; O. and B. Fedtsch. Perech. Rast. Turk. IV, 204.—T. nanum
auct. non Clarke: O. and B. Fedtsch. ibid. 204.—T. tenuifolium auct.
non Jacq.: O. and B. Fedtsch. ibid. 206.—T. xylorrizum Krasch. ex
Stanjuk. Rast. Pokr. Vost. Pamira (1949) 53, diagnosis in Russian—
Chrysanthemum tibeticum (Hook. f. and Thoms.) Hoffm. in Pauls. PI.
coll. in Asia med. and Pers. (1903) 149; B. Fedtsch. Rast. Turk. 737.—
C. nanum auct.: B. Fedtsch. ibid., non Tanacetum nanum Clarke.—C.
tenuifolium auct.: B. Fedtsch. ibid. 738, non Tanacetum tenuifolium
Jacq.—Ic.: Pampanini, Fl. Caracorum (1930) t. VII, fig. 1; Stanjukovicz,
op. cit. 98.

Perennial. Plant 2-10 cm high, with thick, woody, many-headed
root, grayish from profuse pubescence of appressed hairs; stems
numerous from base, usually herbaceous, erect or ascending, densely
leafy, simple or apically branched. Leaves grayish from profuse
pubescence, with punctate glands less distinct because of pubescence;
basal leaves withering early, lower and middle cauline leaves to 1.5
cm long and 1.2 cm wide, with rather long, narrow-winged petiole,
often with linear auricles, ovate to reniform, more or less narrowed
toward base, usually pinnately parted in 5—9 lobes, of which all or
some 3-5 lobed or divided, terminal lobe oblong to almost semicircular,
on the average shorter than in previous species, obtuse; upper cauline
leaves like others but smaller, usually not palmately 3-lobed or
palmately 3-parted. Capitula solitary on up to 1.5 cm long peduncles,
or 2-8 on single stem, aggregated in compact, simple or compound
corymb. Involucres 5-9 mm in dia and 5-6 mm long, more or less
tomentose; involucral bracts herbaceous, outer bracts lanceolate-ovate,
inner bracts ovate, 1.5—2 times as long as outer bracts; all bracts with
wide, brownish, less often light-colored membranous border. Corolla
of tubular florets 2.2-3.0 mm long, few peripheral florets pistillate.
Achenes 1.6—2.2 mm long and about 0.6—0.8 mm wide, lacking corona.
Flowering July to August.

Stony and rubbly slopes, gravel beds, above 3,000 m.—Soviet
Central Asia: Pamiro-Alai Region (Pamir and Trans-Alai ranges).
General distribution: China (Tibet), India (northern part), Afghanistan
(northeastern part). Described from Tibet. Type in London; isotype in
Leningrad.

Note. In our circumscription, A. tibetica is quite distinctly divided
into two ecological forms linked through intermediate plants, but
possibly they are separate ecogeographical races (analogous to A.
scharnhorstii and A. trilobata); the typical form, associated with the
stony and rubbly slopes of river valleys, is small, with a thick woody

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root and larger, often solitary, capitula, and the form of river gravel
beds has a thinner, often herbaceous root, taller stem and smaller
capitula, always aggregated in corymbs. In habit, the latter form
approaches the closely related Himalayan species A. tenuifolia (Jacq.)
Tzvel. comb. nova (Tanacetum tenuifolium Jacq. in DC. Prodr. VI,
129) but is, nevertheless, distinguished from it by wider leaf lobes and
also, on the average, by shorter plants.

GENUS 1545. Hippolytia Poljak.'?

Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVIII
(1957) 288 p. p.—Tanacetum auct. p. p. non L.

Capitula usually numerous (2-15), aggregated at stem apex in
compact or lax corymbs, less often solitary, homogamous, with
numerous bisexual, tubular disk florets. Involucres goblet-shaped, 6—
12 mm in dia and 5-8 mm long; involucral bracts herbaceous or
coriaceous-herbaceous, imbricate, in 3—4 irregular rows, outer broadly
lanceolate to lanceolate-ovate with relatively narrow, membranous
border, inner 1.5—2 times as long as outer bracts, oblong-linear to
oblong-ovate, with wider membranous border appendiculately
broadened at apex; innermost bracts sometimes reduced, linear, almost
lacking border. Receptacle moderately convex, filled glabrous, very
weakly punctate-tuberculate, usually distinctly alveolate. Corolla of
tubular disk florets yellow, 2.5-3.6 mm long, tube strongly and abruptly
broadened in upper half, with 5 lanceolate-triangular teeth, a fifth to
a fourth as long as tube; anther filaments thickened in upper part;
anthers lacking distinct basal appendage, but with lanceolate-ovate,
subobtuse apical appendage; pollen grains globose, spinulose;
style[stigma] bifid, style branches linear, truncate. All achenes similar,
glabrous, 2.0-3.2 mm long and 0.6-0.8 mm wide, almost terete,
narrowed toward base, with 4—7 very inconspicuous longitudinal veins
as indistinct ribs and very loose pericarp; pappus entirely lacking but
upper border of achenes often raised in as obtuse, annular ridge.

Perennial herbs, more or less covered with appressed bifid hairs,
sometimes transitional to simple hairs, with condensed nonflowering
shoots (as rosette of basal leaves); stems erect or ascending, more or
less branched only at base and apex, sparsely leafy, leaves alternate
with divided lamina.

‘Treatment by N.N. Tzvelev.
2Named after the renowned botanist Ippolit Mikhailovich Krascheninnikov, who
worked on the systematics of the tribe Anthemideae.

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392

The 12-15 species of this genus are distributed in the alpine re-
gions of Central and Soviet Central Asia, from the Dzhungarian Alatau
in the north to the Himalayan Mountains in the south; four species are
found in the USSR.

Type of genus: Hippolytia darvasica (Winkl.) Poljak.

Note. Of the nine species proposed by P.P. Poljakov (op. cit.), the
author of this genus, in his original description, three, viz. H.
scharnhorstii (Rgl. and Schmalh.) Poljak., H. trifida (Turcz.) Poljak.,
and H. aphanassievii (Krasch.) Poljak., which with heterogamous
capitula and an entirely different habit (densely leafy stems without
condensed vegetative shoots) are referred by me to the genus Ajania
Poljak., one. more very distinct species—H. kaschgarica (Krasch.)
Poljak., whose systematic position is not yet clear to me, should also
be removed from the genus Hippolytia. Many closely related species
from Tibet and the Himalayan Mountains (Tanacetum senecionis (Jacq.
ex Bess.) J. Gay, T. nanum Clarke, T. longifolium Wall., etc.)
undoubtedly belonging to the genus Hippolytia, although not included
in P.P. Poljakov’s list, must also be added to the remaining five species.
In such a somewhat modified circumscription Hippolytia is a rather
separate genus and shows a much closer affinity with the genera
Dendranthema and Ajania, occupying a place analogous to that occu-
pied by the genus Cancrinia Kar. and Kir. in relation to the genera
Pyrethrum Zinn. and Tanacetum L.

1. Involucral bracts thicker, coriaceous-herbaceous, outer lanceolate-

ovate, inner oblong-ovate. Western Pamir ...............::cesscseseeeeeeeees 2}
+ Involucral bracts herbaceous, outer broadly lanceolate, inner oblong-
linear. Tien Shan, Alai, Dzhungarian Alatau ............ sees eeeeeeeee Bi

2. Leaves usually dull green, bipinnate into relatively broad (1-3 mm
wide) lobes; capitula solitary, less often 2-3 on single stem; border
of involucral bracts usually light-colored 0.0.0.0... cs ceseseeeeeeseeeeees
tee eee press eee seers O tee VY 4. H. schugnanica (Winkl.) Poljak.

+ Leaves grayish from profuse pubescence, bipinnate (or divided)
into narrower (0.5—2.0 mm wide) lobes; capitula (2)5—10(15) on
single stem, aggregated into corymb; border of involucral bracts
usually brownish, less often light-colored ..................sceseeseeseees
Shon akh CR a NE aE ekg 3. H. darvasica (Winkl.) Poljak.

3. Leaves dull green, on the average more divided. Western Tien
SAAN 20:2, eee ee 1. H. megacephala (Rupr.) Poljak.

+ Leaves grayish from profuse pubescence, on the average less divided
spas cna Walaa uc catia Seen 2. H. herderi (Rgl. and Schmalh.) Poljak.

414

393

1. H. megacephala (Rupr.) Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVIII (1957) 289.— Artemisia megacephala Rupr.
in Osten-Sacken and Rupr. Sertum tiansch. (1869) 52; O. and B.
Fedtsch. Perech. Rast. Turk. IV, 198.—Tanacetum capusii Franch. PI.
turkest. (1883) 85, t. 16; O. and B. Fedtsch. Rast. Turk. (1913) 738.—
C. glomeratum Saposhn. in herb.—Ic.: Franch. op. cit.

Perennial. Plant 10-35 cm high, with thick woody rhizome
producing stems and condensed nonflowering shoots, more or less
covered with appressed bifid hairs. Stems few or solitary, ascending or
erect, weakly leafy, branched only at apex (in inflorescence). Leaves
dull green, relatively sparsely hairy (to subglabrous), with indistinct
punctate glands; basal leaves to 8-12 cm long and 4 cm wide, with
rather long (often longer than lamina) petiole strongly thickened at
base, broadly obovate to almost round, twice pinnately cut (or divided)
up to narrowly winged rachis with 2-4 pairs of approximate lateral
lobes; terminal lobes oblong to linear, usually to 1.5 mm, in plants
from shady places up to 3 mm wide, subobtuse or subacute; cauline
leaves few, reduced, with short petiole or sessile, usually pinnately
parted or pinnately lobed; uppermost leaves undivided. Capitula (1)3-
12(15), aggregated at stem apex into compact, often subglobose, simple
or compound corymb, with up to 1 cm, less often up to 2 cm long
peduncles. Involucre 7-11 mm in dia and S—7 mm long, usually finely
tomentose near base; involucral bracts herbaceous, outer broadly
lanceolate with narrow, dark brown or brownish, membranous border;
inner oblong-linear, with wide border of same color, appendiculately
broadened at apex. Corolla of tubular florets 2.8-3.6 mm long. Achenes
2.5—3.2 mm long and about 0.6—0.8 mm wide, lacking corona. Flowering
July to August.

Rocks, stony slopes, above 2,000 m.—Soviet Central Asia: Tien
Shan (Chatkal, Pskem, Ugam, Talass Alatau, Kirgiz, Fergana ranges).
Endemic. Described from Fergana Range. Type in Leningrad.

Note. This species is very similar to the Himalayan species H.
senecionis (Jacq. ex Bess.) Poljak. comb. nova in litt. (=Artemisia
senecionis Jacq. ex Bess. in Bull. Soc. Nat. Mosc. IX (1836) 75;
Tanacetum senecionis (Jacq. ex Bess.) J. Gay in DC. Prodr. VI (1837)
129). However, the two specimens of the latter species (including an
isotype or very badly preserved type specimen) in the Herbarium of
the Botanical Institute are not entirely identical with the herbarium
specimens of H. megacephala, a fact which, given the considerable
gap between the range of these two species, can serve as a basis for
retaining H. megacephala as a separate species, at least until we have
more abundant comparative material of the similar Himalayan species.

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394

2. H. herderi (Rgl. and Schmalh.) Poljak. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR XVIII (1957) 289.—H. leucophylla (Rgl.)
Poljak. ibid.—Tanacetum herderi Rgl. and Schmalh. in Tr. Peterb.
Bot. Sada, V, 2 (1878) 619; O. and B. Fedtsch. Perech. Rast. Turk. IV,
206.—T. leucophyllum Rgl. in Tr. Peterb. Bot. Sada, VII, 2 (1881)
551; O. and B. Fedtsch. op. cit. 206.—T. tomentosum auct. non DC.:
Rgl. and Herd. in Bull. Soc. Nat. Mosc. KL, 3 (1867) 128.—
Chrysanthemum herderi (Rgl. and Schmalh.) Minkw. and Knorr. Rast.
Chimk. Uezda (1910) 112; B. Fedtsch. Rast. Turk. (1915) 738.—C.
leucophyllum (Rgl.) B. Fedtsch. ibid. Ic.: Rgl. in Garten-flora, XXX
(1881) 358, t. 1064.

Perennial. Plant 10-30 cm high, with thick woody rhizome
producing stems and condensed vegetative shoots, grayish from fine
lax tomentum of bifid hairs, in some plants transitional to simple hairs.
Stems usually few, ascending or erect, sparsely leafy, more or less
branched only at apex. Leaves grayish from profuse pubescence, with
inconspicuous punctate glands, leaves as in previous species, but on
the average less divided. Capitula (1)3-10(15), aggregated at stem
apex into rather compact, simple or compound corymb, sometimes
(apparently, under unusual habitat conditions) more lax, and then
peduncles to 3 cm long. Involucres 7-12 mm in dia and 5-7 mm long,
on the average more [densely] tomentose than in previous species;
involucral bracts herbaceous, with wide, dark brown, membranous
border. Corolla of tubular florets 2.8-3.5 mm long. Achenes 2.5—3.2
mm long and about 0.6—0.8 mm wide, lacking corona. Flowering July
to August.

Rocks, stony slopes; above 2,500 m.—Soviet Central Asia: Tien
Shan (central part), Dzhungaria-Tarbagatai (Dzhungarian Alatau),
Pamiro-Alai Region (Alai and Trans-Alai ranges). General distribution:
Dzhungaria-Kashgaria (western part). Described from Trans-Ili Alatau.
Type and isotype in Leningrad.

Note. I do not find any significant difference between H. herderi
and Tanacetum leucophyllum, described from eastern Tien Shan (Kash
River). These species are identical in the nature of their pubescence
and the size of their capitula, and individuals of H. herderi with longer
capitular peduncles (f. racemigera Poljak. in litt.) apparently have no
taxonomic significance.

3. H. darvasica (Winkl.) Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVIII (1957) 289.—Tanacetum darvasicum Winkl.
in Tr. Peterb. Bot. Sada, XI (1890) 317; O. and B. Fedtsch. Perech.
Rast. Turk. IV, 205.—Chrysanthemum darvasicum (Winkl.) B. Fedtsch.
Rast. Turk. (1915) 738.

416

595

Perennial. Plant 10-50 cm high, with thick, many-headed rhizome
producing stems and condensed nonflowering shoots, grayish from
profuse pubescence of appressed bifid hairs. Stems usually few, often
as if axillary in relation to strongly condensed nonflowering shoots,
ascending or erect, sparsely leafy, branched at apex (in inflorescence).
Leaves grayish-green from rather profuse pubescence, with numerous
punctate glands; basal leaves to 8-10 cm long and 4 cm wide, with
long (often 1.5 times as long as lamina), basally strongly thickened
and profusely hairy petioles; obovate or broadly oval, twice pinnately
parted (or cut) up to narrow-winged rachis, with 2-4 pairs of lateral
lobes, terminal lobes (of second order) lanceolate to linear, 0.5—2.0
mm wide, subobtuse or subacute, sometimes in turn 2—4-lobed (or
parted); cauline leaves few, reduced, with short petiole or sessile,
cuneately narrowed toward base, less divided. Capitula (2)5—10(15),
aggregated at stem apex into compact or lax, compound corymb,
peduncles of capitula to 4 cm long. Involucres 6-10 mm in dia and 6—
8 mm long, usually only at base finely tomentose. Involucral bracts
coriaceous-herbaceous; outer lanceolate-ovate, with very narrow
membranous border, inner oblong-ovate, with wide, brownish, less often
light-colored, membranous border, appendiculately broadened at apex.
Corolla of tubular florets 2.5—3.2 mm long. Achenes 2.0—2.8 mm long
and about 0.6 mm wide, lacking corona. Flowering July to August.
(Plate XVIII, Fig. 2).

Rocks, stony slopes, above 2,000 m.—Soviet Central Asia: Pamiro-
Alai Region (Darvaz and Shugnan). General distribution: Afghanistan
(northern part). Described from Darvaz Range. Type and isotype in
Leningrad.

Note. It is distinguished from the similar Himalayan species H.
tomentosa (DC.) Tzvel. comb. nova (=Tanacetum tomentosum DC.
Prodr. V1, 130) by larger capitula and, on the average, wider leaf
lobes. There is an isotype of H. tomentosa in the Herbarium of the
Botanical Institute of the Academy of Sciences of the USSR.

4. H. schugnanica (Winkl.) Poljak. in Bot. Mat. Gerb. Bot. Inst.

‘Akad. Nauk SSSR, XVIII (1957) 289.—Tanacetum schugnanicum

Winkl. in Tr. Peterb. Bot. Sada, XI (1890) 319; O. and B. Fedtsch.
Perech. Rast. Turk. 205.—Chrysanthemum schugnanicum (Winkl.) B.
Fedtsch. Rast. Turk. (1915) 738.

Perennial. Plant 15-40 cm high, with thick, many-headed rhizome
producing strongly condensed nonflowering shoots as if forked in
relation to stems, dull- or grayish-green from rather profuse pubescence
of appressed bifid (in some, transitional to simple) hairs. Stems usually
few or solitary, basally ascending, sparsely leafy, simple or with few

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396

lateral branches in upper part. Leaves usually dull-green from rather
profuse pubescence, with numerous punctate glands, leaves as in
previous species but less divided (lobes of basal leaves often pinnately
or palmately lobed), terminal lobe oblong-lanceolate to broadly linear,
1-3 mm wide, subacute. Capitula solitary or 2-3 at stem apex, on
rather long (to 4—5 cm) peduncle. Involucres 7-11 mm in dia and 6—
8 mm long, usually finely tomentose; involucral bracts coriaceous-
herbaceous, all bracts with rather wide, usually light-colored
membranous border. Corolla of tubular florets 2.5-3.2 mm long.
Achenes 2.0-2.8 mm long and about 0.6 mm wide, lacking corona.
Flowering August to September.

Rocks, stony slopes, at 1,500—2,500 m.—Soviet Central Asia:
Pamiro-Alai Region (between Darvaz and Shugnan ranges). General
distribution: Iran-Afghanistan? (Afghanistan). Described from the above
mentioned area. Type in Leningrad.

Note. In relation to the previous species, it apparently is a lower
altitude ecogeographical race. However, it is not ruled out that typical
specimens of this species are random, broader leaved plants of second-
ary flowering, which belong to H. darvasica. It is necessary to have
more extensive material for a definitive solution of this question.

GENUS 1546. Filifolium Kitamura!”

Kitamura in Acta Phyt. et Geobot. IX (1940) 157.

Capitula (1)2—10(15), on short erect peduncles, aggregated into
simple or compound corymbose inflorescence, heterogamous, but
lacking ligulate florets, with one row of (1-10) fertile pistillate florets
and numerous (about 30-40) bisexual but usually sterile (with more or
less rudimentary pistil) disk florets bonded together into compact mass
by resinous secretions of punctate glands. Involucres goblet-shaped,
4-5 mm in dia and 4-5 mm long; involucral bracts imbricate, in 3
irregular rows, coriaceous-herbaceous, with rather wide, brownish
membranous border; outer bracts ovate-roundish, middle and inner
broadly elliptical, 1.5 times as long as outer. Receptacle strongly
convex, obtusely conical, almost as long as wide, full, glabrous, very
indistinctly alveolate. Corolla of peripheral pistillate florets tubular,
dorsally compressed and distinctly narrowed at apex, with 2—4 very

‘Treatment by N.N. Tzvelev.
7From the Latin words filum—thread, and folium—leaf,; so named for the very
narrow (almost filiform) leaf lobes.

418

ao]

short, obtuse teeth; corolla of bisexual disk florets also tubular, yel-
lowish, apically not broadened, with 5 (less often 4) broadly
deltoid acute teeth. Anthers basally obtuse, with lanceolate-ovate,
subobtuse apical appendate; Styles[stigmas] short, bifid; style branches
linear or oblong. All achenes similar, glabrous, 1.8-2.5 mm long,
globose-obovoid, strongly compressed dorsally, with 2 distinct
(sometimes narrowly winged) lateral ribs, finely and indistinctly
longitudinally sulcate, slightly narrowed toward base, lacking
pappus (corona), but often with acute, more or less truncate upper
border, pericarp very slippery when soaked.

Wholly glabrous perennial plants, base of shoots profusely covered
with fibrous remnants of old petioles, relatively sparsely leafy erect
stems, condensed vegetative shoots and 2- (sometimes 3-) pinnately
cut alternate leaves with almost filiform terminal lobe.

A monotypic East Asiatic genus.

Note. It occupies a rather distinct position in the tribe Anthemideae,
being closest to the genus Artemisia L. It belongs to the group of
wind-pollinated plants and like the genus Artemisia L. it is distinguished
from the genera Tanacetum L. and Ajania Poljak., aside from the
peculiarities of the structure of the capitula and the achenes, also by
the nonspinulose pollen grains.

1. F. sibiricum (L.) Kitamura, op. cit., p. 157.—Tanacetum
sibiricum L. Sp. pl. (1753) 844; DC. Prodr. VI, 129; Ldb. Fl. Ross. II,
603; Turcz. Fl. baic. dahur. II, 75; Kom. and Alis. Opred. Rast.
Dalnevost. Kr. II, 1032.—Artemisia sibirica (L.) Maxim. in Mél. Biol.
VIII (1872) 524.—Chrysanthemum trinioides Hand.-Mazz. in Acta
Horti Goteoburg. XII (1938) 273, nom. nov.—Ic.: Gmel. Fl. Sib. II
(1749) 134, t. 65, fig. 2—Exs.: GRF No. 1118.

Perennial. Plant 20-60 cm high, with thick oblique rhizome, wholly
glabrous. Stems solitary or few, basally covered with thick layer or
fibrous remnants of old petioles, erect, obtusely ribbed, branched usually
only at apex (in inflorescence). Leaves dull green; basal leaves to 20
cm long and 5.0-6.6 mm wide, with long petiole, not longer than
lamina, obovate to linear-oblong, 2-(sometimes thrice-) pinnately cut,
terminal lobe linear-filiform, to 1 mm wide and 4 cm long, more or
less revolute, subacute; cauline leaves 1—5, like basal leaves but smaller
and with shorter petiole (to subsessile and entire uppermost leaves).
Capitula 1-10 (less often to 15), usually aggregated in rather compact,
simple or compound corymbs, peduncles to | cm long. Involucres 4—
5 mm in dia and 4-5 mm long. Corolla of bisexual florets yellowish,
1.8—2.4 mm long, in pistillate florets usually somewhat smaller. Achenes
pale brownish, 1.8—2.5 mm long and 1.5—2.0 mm wide. Flowering July
to September. (Plate XIX).

398

419

Filifolium sibiricum (L.) Kitam.
pistil of peripheral pistillate

Plate XIX.
Habit, disk florets, peripheral pistillate floret,
floret, anther, achene, involucral bracts.

421

399

Stony slopes, steppes, forest glades, up to lower mountain zone.—
Eastern Siberia: Angara-Sayans, Dauria; Far East: Zeya-Bureya, Ussuri.
General distribution: Mongolia, China (northeastern part), Korea.
Described from Eastern Siberia. Gmelin’s figure cited above is type
specimen.

GENUS 1547. Cotula L.!:?

Li Sp. pl. (1753) 892.

Capitula many-flowered, small, yellow; tubular florets hetero-
gamous, peripheral female florets lacking ligule, homogenous, with
compressed tube, often at base with 2 spurs and 4-notches. Receptacle
smooth, lacking pales, after fruit-setting with papillae (stalks), more or
less distinct. Achenes flat-compressed, wide, along edges glabrous,
lacking papillae; outer achenes with stalk, achenes of central tubular
florets often sessile, narrow.

Leaves toothed, pinnate or bipinnate, with narrow lobes; branches
terminating in peduncles with solitary capitula. Annual, small, branched
herbs.

All species of this genus (about 75) are found mainly in the
Southern Hemisphere, most of them originating from South Africa (in
the USSR one introduced species), a smaller number from Australia
and South America.

The achenes of most species of this genus are inadequately and
incompletely described.

1. C. coronopifolia L. Sp. pl. (1753) 892; DC. Prodr. VI, 78;
Roth in Engl. Bot. Jahr. 5, 337; Hegi, Ill. Fl. VI, 2, 622.—Ic.: Hegi,
op. cit. fig. 335; Rchb. Ic. Fl. Germ. XVI, t. 998.

Annual. Stems weak, erect or procumbent, simple or branched
from base. Leaves lanceolate-linear or lanceolate, entire, toothed or
pinnate, with linear, narrow, small toothed segments, obtuse or short-
acuminate, at base amplexicaul; upper leaves narrow, linear, entire.
Capitula solitary at apices of stems and branches; receptacle
hemispherical, covered with papilliform outgrowths (stalks). Involucral
bracts few, elliptical, obtuse, green or reddish with narrow, white,
membranous border, glabrous. Peripheral female florets one-whorled
with long, compressed, slender stalks; ovary large, with wide, white
edges; corolla scarcely visible in depression of ovary; tubular disk

'Treatment by E.G. Pobedimova.
*From the Greek word cotule—palm; so named for the form of the leaves.

422

400

florets with long tube (2 times as long as narrow achene), with 4 short
roundish teeth; anthers and styles inserted in corolla tube. Achenes of
peripheral florets 1.0-1.25 mm long and 0.75-1.0 mm wide, flat,
compressed, with white broad ribs, along border; achenes of disk florets
narrower, oblong, 0.75-1.0 mm long, and 0.25-0.5 mm wide; all
achenes smooth, dorsally with papilliform stalk, lacking pappus.
Flowering July to August. (Plate VI, Fig. 3).

Along seacoasts, mainly in muddy-marshy low-lying areas.—Far
East: Ussuri, Sakhalin. General distribution: Scandinavia, Atlantic
Europe, Western Mediterranean, North and South America, Australia,
Africa. Described from South Africa. Type in London.

GENUS 1548. Centipeda Lour’”

Lour, Fl. Cochinch. II (1790) 492; Benth. and Hook. Gen. pl. I
(1873-1876) 430; O. Hoffm. in Pflzfam. IV, 5 (1897) 280.—Myriogyne
Less. in Linnaea, VI (1831) 219.

Capitula solitary, 2.5—-8.0 mm in dia, many-flowered, hetero-
morphic, globose, subsessile in racemes at apices of branches and in
axils of pseudo-opposite leaves. Involucres hemispherical; involucral
bracts biseriate, imbricate, oblong or obovate, scarious,
along margin hyaline, greenish in middle, upright, concave, ventrally
glabrous, dorsally with sparse glandular pubescence. Receptacle flat
or hemispherical, glabrous. Florets greenish or reddish, very small,
regular, tubular, with very short corolla tube shorter than
ovary; peripheral florets pistillate, fertile, numerous, with very narrow
tube and 2—3-lobed narrow limb, central florets bisexual, fertile, few
(4-8), campanulate, with 4-lobed very short limb; stamens 4,
anthers connate into tube surrounding style, with short, undivided,
subsessile basal appendage; style short, wide; stigma bifid, lobes
short, truncate; ovary oblong. Achenes oblong, angular, with 4, less
often 3-5 obtuse ribs, with crisped glandular pubescence of
hairs, lacking pappus. Leaves alternate, approximate, narrowed toward
base, spatulate, oblong or oblong-obovate, undivided or toothed at
apex. Annual or perennial herbs, glabrous or with whitish-grayish
pubescence.

Of the five species of this genus, three grow in Australia, and one
has been reported from Chile. Only one species is found in the USSR.

‘Treatment by L.A. Smoljaninova.
*Named for the habit of the plant; the numerous branches of the plant are
procumbent on the ground.

423

401

1. C. orbicularis Lour, Fl. Cochinch. II (1790) 493; Maxim. Prim.
Fl. Amur. 163; Kom. Fl. Manchzh. III, 650.—Myriogyna minuta Less.
in Linnaea, VI (1831) 219.

Annual. Stem to 10 cm high, elongate procumbent or assurgent,
strongly branched, glabrous or weakly silky pubescent at apex. Leaves
6-17 mm long and 2-4 mm wide, evenly distributed, alternate, oblong,
entire or with 2—4 teeth in upper part, subobtuse or acuminate, cuneately
narrowed toward base, glabrous. Capitula small, 2 mm long and 2-4
mm in dia, heteromorphic, hemispherical, solitary, axillary, subsessile.
Outer involucral bracts 1.5 mm long and 0.5 mm wide, obovate, white,
membranous, along margin hyaline, light green in middle, ventrally
glabrous, dorsally with sparse, long, appressed, glandular hairs,
unevenly small-toothed; inner bracts 1 mm long and 0.5 mm wide.
Receptacle hemispherical, glabrous. Pistillate florets 0.25 mm long,
much shorter than ovary, with small, greenish, 3-lobed limb; style as
long as corolla, flattened, glabrous; stigma bifid, weakly exerted from
corolla tube, stigma lobes ovoid, short, flattened, erect, approximate,
papillate; ovary 1 mm long and 0.3 mm wide, oblong, slightly curved,
narrowed toward base, with 3-4 ribs, with stalk, at apex with 3 fleshy
thick, short, indistinct lobes, with small glandular hairs on ribs, with
sparse, long, erect, apically curved glandular hairs on surface; bixexual
florets few, their corollas 0.5 mm long, 4-lobed, lobes ovate, erect,
reddish, glandular along margin; style 0.5 mm long and 1.2 mm wide,
subellipsoidal, flattened, glabrous; stigma bifid, with small papillae,
stigma lobes short, flattened, upward directed, weakly divergent,
falcately incurved; ovary 1 mm long and 0.3 mm wide, oblong, narrowed
toward base, with short stalk, at apex with 4 fleshy, short lobes, each
with one short glandular hair at apex, with occasional, erect and
squarrose hairs curved at their ends; stamens 4, anthers 0.3 mm long,
connate into tube surrounding style, roundish, apically truncate, with
very short, undivided basal appendage, subsessile; anther filaments
flattened, free, glabrous. Achenes 1 mm long, oblong, with 3-4 ribs,
with small glandular hairs along ribs, on body with occasional, long
glandular hairs curved at apex. Flowering July to August.

Muddy shoals of banks of rivers, lakes and streams, marshes, as
weed on wet and clayey arable lands, roadside ditches and fields.—
Far East: Uda River Area, Ussuri, Zeya-Bureya. General distribution:
Madagascar Island, Mascarene Islands, peninsular Indochina (Burma,
Malaya), Indonesia, New Caledonia Island, northern Australia, New
Zealand, Philippine Islands (Luzon Island), Tahiti Island, China (Taiwan
Island, provinces: Kwangtung, Fukien, Shansi, Shantung, northeastern
provinces of Yunnan), Japan, Korea, Mongolia. Described from
Indochina. Type in London.

424

402
GENUS 1549. Kaschgaria Poljak.':?

Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVIII
(1957) 282.

Capitula ovoid, diversiflorous; involucres narrow cup-shaped, 2-
4 whorled, imbricate. Receptacle convex, conical, glabrous. Florets
heterogeneous, peripheral florets 3—S, pistillate, their corolla narrowly
tubular, broadened toward base, with 2-3 teeth, lobes of stigma linear,
acute, divergent; disk florets 11-17, bisexual, their corolla tubular, 5-
toothed, sparsely stellate-hairy; anthers lanceolate-linear, apical
appendages of anthers acute, angular, basal appendages short,
subobtuse; stigma lobes short, linear, weakly thickened above, short
ciliate, after anthesis divergent. Achenes ovate, with flattened apex,
with small uneven border. Pappus absent. Low semishrubs with simple,
linear partly ternate, pinnately cut leaves.

The genus includes two species distributed in lower mountains,
partly in western Mongolia, western China (Sinkiang) and eastern
Kazakhstan.

Type of genus: Kaschgaria brachanthemoides (Winkl.) Poljak.

1. Fertile branches usually strongly branched, with more or less long,
divergent shoots; leaves mostly narrow, lanceolate-linear, simple;
capitula \clustered ims fasciele ty. ..vss2:522402. eee. ls atleast
Ss he Aude Many. Sad. date 1. K. brachanthemoides (Wink1.) Poljak.

+ Stems with rather short branches; most leaves ternate or pinnatisect,
with wide lobes; capitula in compact corymbose or partly fascicular
INMOFESCENCEy Reals. b..chessay. eee 2. K. komarovii (Krasch.) Poljak.

1. K. brachanthemoides (Winkl.) Poljak. comb. nov.— Artemisia
brachanthemoides Winkl. in Tr. Peterb. Bot. Sada, IX, 2 (1886) 422;
B. Fedtsch. Perech. Rast. Turk. 1V, 200.—Tanacetum brachanthemoides
(Winkl.) Krasch. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, I (1933)
175; Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, IX, 168.

Cultivated. Smooth semishrub, 40-55 cm high. Tap root woody,
thick, producing perennial, woody, short, strongly branched stem, like
branches covered with grayish, cracking bark; fertile branches numerous,
upright, 20-35 cm long, slender, angular-ribbed, in lower part stiff,
virgate, stramineous, smooth, light green above, repeatedly branched
with long, slender, divergent leafy branches. Leaves simple, sessile,

'Treatment by P.P. Poljakov.
?Named after the old name of the Chinese province—Kashgar, with which the
distribution of the genus is linked.

425

403

longitudinally rugose, glabrous or with sparse stellate hairs, narrow-
lanceolate-linear, linear, or narrow-oblong-linear, short-cuspidate,
usually entire, or sometimes apically trilobate; lower leaves often 1.2-
2.0 cm long and 1.7—5.0 mm wide, upper considerably shorter, narrow-
linear. Capitula on short peduncles, sometimes subsessile, upright, ovate,
4-5 mm long and 1.5-2.7 mm wide, 2-5 at apices of branches,
aggregated in fascicles, partly solitary, remote. Involucral bracts
herbaceous, with scarious border; outer bracts small, almost roundish,
inner considerably large, oval or lanceolate-elliptical, apically roundish.
Receptacle convex, narrow-conical, glabrous. Peripheral florets
pistillate, fewer, their corolla narrow-tubular, broadened toward base,
3-toothed, punctate-glandular, stellate-hairy, stigma lobes linear, acute,
after anthesis scarcely exserted from corolla tube; central florets
bisexual with tubular 5-toothed corolla, filaments quite long, anthers
lanceolate-linear, with acute angular apical appendage, basal appendage
short, subobtuse, stigma lobes short, linear, slightly thickened upward,
divergent, short ciliate. Achenes ovoid, with flattened apex, weakly
bordered. Flowering August. (Plate XX, Fig. 2).

On rocks of low desert hills —Soviet Central Asia: Tien Shan
(eastern part of Trans-Ili Alatau). General distribution: Dzhungaria-
Kashgaria (Chinese part of Tien Shan). Described from Tien Shan.
Type in Leningrad.

2. K. komarovii (Krasch.) Poljak. comb. nova.—Tanacetum
komarovii Krasch. and Rubtz. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, [X (1946) 168; Kryl. Fl. Zap. Sib. XI, 2759.

Cultivated low semishrubs, 25-45 cm high; tap root woody, 2 cm
thick, with short, perennial, woody, ascending vegetative shoots above,
covered with brownish-gray cracking bark. Fertile stems (or branches)
numerous, upright, erect or more often weakly bent, woody below,
stramineous, lustrous, light green above, angular-ribbed, sparsely stellate
hairy. Leaves dark green, 1-3 cm long and 0.7—1.5 cm wide, 3- or 5-
lobed, less often pinnately cut into lanceolate, short, acute, or subobtuse,
0.5-1.5 cm long and 1-4 mm wide lobes, sparsely stellate hairy on
both sides, lowermost leaves sometimes undivided, oblanceolate, upper
bracteal leaves, small. Capitula on short peduncles, ovoid, 3.5-4.0 mm
long and 2-3 mm wide, aggregated at apices of branches and stems
into short compact corymbs or fascicles. Involucres narrow cup-shaped,
imbricate; involucral bracts herbaceous, scaly along margin, outer
almost round, inner larger, oval. Peripheral florets pistillate, with narrow
tubular corolla broadened toward base, 3-toothed; central florets
bisexual, with tubular 5-toothed corolla. Achenes ovoid, with flattened
apex. Flowering August.

404

Stony slopes of low desert hills.—Soviet Central Asia: Lake
Balkhash Region (southern part of Altai in Zaisan District and northern
banks of Lake Balkhash, Dzhungaria-Tarbagatai (Dzhungarian Alatau).
General distribution: Dzhungaria-Kashgaria (western Mongolia and
western China). Described from Kaldzhir River valley. Type in
Leningrad.

GENUS 1550. Artemisia L.!-?

L. Sp. pl. (1753) 845 and Gen. pl. ed. V (1754) 357.

Capitula numerous, small, aggregated in paniculate, racemose, less
often spicate or subcapitate inflorescence, heterogamous, with pistillate,
tubular, peripheral florets in one row and more numerous bisexual
tubular central florets. Involucres hemispherical, ovoid, goblet-shaped
or cup-shaped, up to 8(10) mm wide; involucral bracts herbaceous or
coriaceous-herbaceous, imbricate, in 2-6 irregular rows, outer bracts
gradually reducing and considerably shorter than inner bracts or all
bracts almost equal, all or some with more or less distinct membranous
border. Receptacle usually convex, conical, or hemispherical, less often
almost flat, punctate-alveolar, glabrous or somewhat hairy. Corolla of
pistillate florets very narrow, sometimes almost filiform tubular, 2-,
less often 3-toothed, almost colorless, or corolla tubular, 2-3—4-toothed,
colored; corolla of central florets usually campanulately or cup-like
broadened in upper part, 5-toothed, yellow of various shades or reddish-
violet; anthers lacking distinct basal appendage, linear, with lanceolate
or oblong-lanceolate, acute, less often sub-obtuse apical appendage;
pollen grains globose, with smooth or almost smooth surface, always
nonspinulose. Style[stigma] bifid, lobes of stigma in peripheral pistillate
florets usually narrow-linear, apically more or less narrowed, acute or
subobtuse, lacking hairs or cilia, in bisexual central florets linear,
apically truncate, barbate from upright hairs; style of rudimentary pistil
of staminate florets apically not bifid, infundibuliform, with erect ciliate
hairs or short connate lobes. Achenes small, homogeneous, obovate or
oblong-ovate, almost terete or flattened, glabrous, weakly sulcate or
with indistinct, longitudinal ribs of which only 2 almost winged, distinct,
brownish, apically more or less roundish, lacking pappus but sometimes
with slightly raised upper edge (as if rudiment of corona). Perennial,
biennial and annual, geiconogamous [with fertilization between different
flowers on the same plant] or autogamous [self fertilized] herbs or

'Treatment by P.P. Poljakov.
*The name of the common wormwood, derived from the Greek name “Artemisa”
or “Artemisia” (“Artemisa” was the wife of king Mausolus).

427

405

semi-shrubs with erect or ascending, sometimes procumbent stems and
alternate leaves, with more or less divided, less often undivided or
entire lamina.

A very large, undoubtedly heterogeneous genus comprising more
than four hundred species distributed mainly in the temperate zone of
Europe, Asia, and North America. One hundred and seventy-four species
are found in the Soviet Union.

Type of genus: Artemisia vulgaris L.

KEY TO IDENTIFICATION OF SUBGENERA

Capitula homogamous, with few ( to 8—10) bisexual florets; involucre
ovoid, with 3-6 rows of imbricate, unequal bracts; receptacle very
smatiiconical ~.!0..0.0i..ba: Subgen. 3. Seriphidium (Bess.) Rouy.
Capitula herterogamous, with pistillate peripheral florets and usually
more numerous bisexual or staminate central disk florets; involucre
of 2-3 rows, the bracts differing very little in size; receptacle quite
PUR RRMIOIN TAO BA iE cnecvenevcnsnerssesecensensnes ciel scsbiaeber TORMOROidessoovents 2.
Disk florets staminate, but with abortive pistil.. ................008
SEES, Sue lets. Gili. {33 Subgen. 2. Dracunculus (Bess.) Rydb.
Die MaretS DISeKual .............n.-nnncsanrnnnns Subgen. 1. Artemisia Less

Subgenus 1. Artemisia.—Less. Syn. Comp. (1832) 264.—Capitula
heterogamous; peripheral florets pistillate, with narrow-tubular corolla
having 2 teeth, disk florets bisexual, fertile, with 5(4) teeth. Receptacle
glabrous or hairy.

Type of subgenus: type of genus.

L.
+
2:

2

*

Anmuals, or, biensials, .......D6Gididg- «BX sein dba Gigli Bie etahes owdiens 2.
Pete miniats- ore emits hr se. «2206s 63s. ..0 cen ides Leavy sateen nee 13,
Plants more or less densely white hairy, less often pubescence
partly lost by end of vegetative growth; receptacle hairy, less
Bibeay patra iS i. Shs) Berk aes chan dl tess A Mae E. Geaeee vo odeeses 3.
Plants green, glabrous or weakly hairy; receptacle glabrous ...... 8.
Capitula subglobose or broadly campanulate, 2-4 mm wide
Wee BROGS, LEY TROL EARS. AUNTS. SRS See. DELL. cals eee ncartaces 4.
Capitula hemispherical, usually 5—8(10) mm wide ................05 >.
Plants up to 80-120 cm high, grayish from dense pubescence;
terminal lobes of leaves oblong-linear, 3—8 cm long .............:0006
rrbitssne pMAASINUh Anz HLS «chats Shes oUREIe dock dit og 77. A. kulbadica Boiss.
Plants shorter, glabrescent, green; terminal lobes of leaves shorter,
ree gS SS ca Ne J 81. A. anethifolia Web.

428

406

8(2).

13.5

Capitula numerous, in more or less lax paniculate inflorescence
shih URES ART, op SIRE ED gee ae SORES, og okies Mae, nl ae 6.
Capitula relatively fewer, in simple raceme or partly spicate
inflorescence %:..2L.GNas rater uae ease, mame Aes Me A
Stem 60-100 cm high, ribbed; lower cauline leaves large, to 12
cm long, their terminal lobes oblong-linear ...............c:.csseeeseeeees
axpavadeudesdlodtaustavancarterar seduces ee aa 75. A. sieversiana Willd.
Stem 25—40 cm high, round, sulcate; lower cualine leaves 2.0-4.5
cm long, their terminal lobes narrowly linear ................csseeseeeeeee
wa sleshls i aR cat it ste ae ego 80. A. jacutica Drob.
Lower cauline leaves 2-3 cm long, pinnately cut or thrice pin-
nately cut, their terminal lobes linear or linear-lanceolate, 1.0—1.5
ITUAT WAGES 215. Lik 2 Set See 78. A. samoiedorum Pampan.
Lower cauline leaves mostly up to 1.5 cm long, twice pinnately
cut, their terminal lobes narrower, up to 0.5 mm wide .................
is AGNI SL SANSA GI NOE CORLL ENE Mr HEE 76. A. macrocephala Jacq.
Stem 120-190 cm high, brownish-violet, almost simple terminat-
ing into long, narrow, dense paniculate inflorescence with short
SIGNER DEARCHES occcks exes ce nsccscspaeecnias 52. A. tournefortiana Rchb.
Plants. shorter, characters otherwise (sccck.-t...cesess se eee eee nce .
Capitula hemispherical, 46 mm in dia, with goblet shaped, woolly-
RATE! IN VONICRO i ceisecentecpeniiensemsstsannerenand SARE 76. A. succulenta Ldb.
Capitula smaller, 2-3 mm in dia, subglobose, with cup-shaped,
glabrous; smoothiinvolucre 2.4%. eee 2... ae eee 10.
Stemis 310; "slender, '8=12 «cm phigh! «si0.22.-2isc;.,cceeeesteeaeeeeeeeeaee
Leo. AEE Me i At 81. A. anethifolia var. korotkyi (Krasch.) Poljak.
Stem solttary, taller... .......0ccescmancdtereess<sosvecercapa te Rene. eat. enn fl.
Lower cauline leaves thrice pinnately cut, their terminal lobes
oblong-lanceolate, entire or with 1—2 teeth ....... 53. A. annua L.
Leaves. once. Or, (WICE: pIntate .......8ss isi taysth. ooh eee esa 2.
Stem 10-50 cm high; lower cauline leaves 2—5 cm long, once or
twice pinnately cut; terminal lobes of leaves filiform-linear
wis sGeaeede hartabet acu he Or bed ele gEM OR OHERNEE , SEP ROCIO WES A! 54. A. palustris L.
Stem 15-60(80) cm high; leaves 6-12 cm long, twice pinnately
cut; terminal lobes of leaves narrowly linear-lanceolate...............
Leadereadaententapeaonveeaelbttwts Mende eta.. Wet. MUM UERRE LE sta 55. A. aurata Kom.
Leaves, for the most part, green, subglabrous above, whitish be-
neath from short arachnoid tomentum; corolla of pistillate periph-
eral florets small, almost filiform tubular, with 2 teeth, almost
colorless, much smaller than corolla of disk florets; style lobes of
pistillate florets acute, those of disk florets obtuse ................. 58.

*Whole of couplet 13 as per changed version given in the errata to Vol. XXVI
published on p. 756 of Vol. XXVII.—-General Editor.

429

++

14.

5

16.

17.

19.

20.

407

Leaves usually concolorous; corolla of pistillate peripheral florets
with 2-3 teeth, somewhat smaller than corolla of disk florets;
style lobes of pistillate florets subobtuse or subacute, those of
Gabe Trt DUES in sninnisncnknes Sage easdte oleae L Ae dh gual nee oes 14.
Leaves usually concolorous; corolla of pistillate peripheral florets
with (2)3—5 teeth, scarcely smaller than or equal to corolla of disk
florets, both colored; style lobes of all florets obtuse, truncate
hee abies bea twekeon avandan Sot ldlaba: 78.
Capitula large, 5-8 mm long and 4-6 mm wide, in spicate
inflorescence; achenes 3-4 mm long and 0.3-0.4 mm wide, linear-
oblong, smooth, loosely enclosed in membranous sheath ...............
fete WR ldare ty a Fass Vado cavtdpstsasonsidavsdepadthaonsd 21. A. stelleriana Bess.
Capitula smaller; achenes different, in tightly adhering sheath

Capitula in dense fasciculate corymbs at apices of more or less
inclined branches, usually forming corymbose-paniculate
inflorescence; corolla with appressed hairs at apex ...........scseeseeeeeess
sammuantaies eis ie tie a Liat Ca NE sce eats. clive 82. A. incana (L.) Druce.
Capitula differently borne; corolla glabrous at apex or with straight
PB te ciceetes flap. alain pete tices ees 16.
Plants more or less densely covered with white appressed or erect
hairs; receptacle hairy, less often glabrous; corolla of bisexual
flonets:partly hairy. in upper partes. isnsic8. cect thas. Ae
Plants with green or glaucescent-green, usually sparsely hairy or
almost glabrous leaves, or leaves green above and whitish-
tomentose beneath; receptacle glabrous, less often hairy; corolla
small yc laters yoctlesl cae esos Sains Desh eka cats a aea gE CEES 30.
Stem sulcate, 60-100 cm high; lower cauline leaves 6-9 cm long
and 3-7 cm wide, lamina broadly oval, almost thrice pinnately cut
aia iitinaritadbeinhinneinsinhapninionscn 74. A. absinthium L.
Stem not sulcate, less tall; leaves smaller ..............cceecsseesseees 18.
Lower cauline leaves thrice pinnately cut; semishrub, glaucescent-
grayish from dense crisped hairs (mountains of Soviet Central

PB) sevesteoncs serve tivectuts tcl vasa leltece testedadlnc tin 2S 67. A. persica Boiss.
Leaves less complexly divided; plants whitish or gray, appressed
MAN eal waht sa bias cca chy «nc cae dS RNID 19.
Lower cauline leaves ternately dissected or ternately toothed or
Gven! cutire <2, xiiscoupiees: aii, shel x sei ane, tas. 5 2 20.
Lower cauline leaves, all or some, bipinnatifid or bi- or tripinnate,
an liianstininnoatifid) miciet.ci:sibs i150. 0. OA RN 22.

Semishrub, 30-60(70) cm high; capitula hemispherical 4-7 mm
in dia, with goblet-shaped involucre; leaves more weakly hairy
above than beneath ..................... 65. A. lagocephala (Boiss.) DC.
Perennials, 3-15 cm high; capitula subglobose, 2~4(5) mm in dia,

430

408 -

25:

26.

20:

30(16).

with cup-shaped involucre; leaves whitish-hairy on both sides
widcvhe tke aditel Pedetch «Manad. RUBREAUEUEL. LEU Wier, OEE erate, MRE et atta pA
Capitula 2-3 mm in dia; receptacle glabrous; stems numerous or
few, .(3) 10-15. cra iS nscsvusicctuemcee ee 60. A. caespitosa Ldb.
Capitula 4-5 mm in dia, receptacle hairy; stems 1-2, to 6-10 cm
High's, £55. Sese0ed, Mek USS). eee tte oe a 61. A. cuspidata Krasch.
All or most capitula on long (1-3 cm) peduncles .................. 23.
Most capitula on shorter peduncles or subsessile ................... 24.
Capitula in simple racemose inflorescence; peduncles not leafy
AEA TELETE, Sestdvad btascbdne oceebaeh leetan, deekos ceanebetes 57. A. splendens Willd.
Capitula in paniculate inflorescence; peduncles leafy ....................
PI WENT ot oe cos swoscncvasaerauecanmeureane 62. A. schmidtiana Maxim.
Capitula hemispherical, usually wider than long; involucre goblet-
shaped; terminal lobes of leaves mostly oblong-oval, lanceolate-
oblong, of lanceolate-lmear sito. ek. S20 ee 2D.
Capitula subglobose or broadly ovate, less or as wide as long;
involucre cup-shaped; terminal lobes of leaves narrowly linear or
Laneel ate WMCAL... ..050i5:cisoree.«dearscouacacnapanineawenrnccterstrteeReren aca eae Dis
Semishrubs, with tap root; terminal lobes of leaves oblong-oval or
linear-oblong, round or scarcely pointed .............cseeseeseeeseeeneeeeeeee
Lyd ACECUR, RRO, BRELYZ, PEAVEN S8 4, A ba irceiaoek eaten 66. A. rutifolia Steph.
Perennials with creeping rhizomes; terminal lobes of leaves lan-
ceolate-linear,. acuminate as, tose. Ake, Rete eee 26.

Capitula in racemose-paniculate inflorescence; lower leaves 3-5
cm long; stem 40-70 cm high ........ eee 63. A. sericea Web.
Capitula in simple racemose inflorescence; lower leaves 1.5—2.5
em. _long;, stem usually..15—35 cm Wigh..:.cu Mec. seeteeres onmcea ares
eectds GERAIS, BORIS ED IE RES, 8 64. A. aschurbajevii Winkl.
Receptacular hairs absent (receptacle glabrous) .................s:seseee
LTA 2: a AO ROOD NO EE eV RO eS hit Aa 59. A. austriaca Jacq.
Receptacular hairs always present (receptacle hairy) ............... 28.
Terminal lobes of leaves narrowly linear or narrow lanceolate-
linear, 5-10 mm long; plant of Caucasus ..............cceeceeeeeeseees 29.
Terminal lobes of leaves oblong-lanceolate or lanceolate-linear,
smaller, 1.5—5.0 mm long; plants of Siberia and Kazakhstan ........
ssa snnchsweseceatutplentudooOadee ie ict ein FERRE Ne doakete ne 58. A. frigida Willd.
Capitula 4.0-6.5 mm in dia, in simple racemose inflorescence;
leaf lobes narrowly lanceolate-linear usually 5-8 mm long .............
seh oWhdes Sih abet eae MeL IEG, OTIIR, Manga ecteeas 56. A. caucasica Willd.
Capitula up to 3 mm in dia, in racemose-paniculate inflorescence,
often in simple racemose; leaf lobes narrowly linear, usually 5-10
mm long ....... 56. A. caucasica var. grossheimii Krasch. ex Poljak.
Receptacle baa. 2k. .i2 csecessesevvsiss ovens MURROUM AMO Mec enn 3i.

431

39:

34.

35(30).

38.

Receptacle glabrous; hairs lacking or caducous ...............:00 a5:
Perennial herb, root in upper part having perennial, decumbent,
vegetative shoots and ascending, simple, herbaceous, brownish-
WAI RE As Brae ses sai sins a nedes bth Vea Leite RM Ra aR dee ces 32:
Semishrub; root thick, sometimes many-headed, bearing peren-
nial, woody, ascending, short, vegetative shoots and straight brown-
ish, branched or almost simple, stiff, virgate stems ................. 33%
Stem 20-50 cm high; leaves up to 1.5—5.0 mm long, glabrous or
weakly hairy only beneath; plants of plains and mountains ...............
sheaves <7 bette alls dlc uiatak 69. A. rupestris L.
Stem 10—25(35) cm high; leaves up to 1.5—2.0 cm long, glabrous
or weakly hairy above, usually hairy beneath; plants of mountains
etadetie treet ale cassasteszanns 69. A. rupestris var. viridis (Willd.) DC.
Capitula small, mostly 2—3 mm in dia, in rather narrowly panicu-
late inflorescence; terminal lobes of leaves oblong-lanceolate or
Gs es ee a a or 70. A. obtusiloba Ldb.
Capitula 3-5(6) mm in dia, in simple raceme or very narrow
racemose-paniculate inflorescence; terminal lobes of leaves lan-
ceolate or linear, short-acuMinate ..............:ccscessesssecessessseeeenes 34.
Fertile stems 5—20 cm high; terminal lobe of leaves lanceolate,
1.5—3.0 mm long; plants hairy in beginning of vegetative growth,
fate? SUDBIAGAINNO AS, oi. LU MERTSS 73. A. altaiensis Krasch.
Fertile stems 25—35(45) cm high; terminal lobe of leaves linear,
up to 3-6 mm long; plant generally densely appressed-hairy to
end of vegetative growth ............seeeeeee 71. A. subviscosa Turcz.
Semishrubs or perennials with woody tap root, in upper part pro-
ducing short, perennial, woody shoots and more or less numerous

Min Bate Testwhe SMG 26821 OL A ZA ce 36.
Herbaceous, rhizomatous perennial, mostly with solitary stem (very
PAN OR yc erste tseseetite cs sacocscccssaccscestsstaaccanseseter AR oeaaminens 46.
Whole plant or leaves, involucre, and stems in part whitish, densely
abachnei@hainy Mi A TS a RA A A, RA antes a7;
Plants green, weakly hairy or glabrous, very rarely leaves more or
less.oubescent bétieath......23.4 102242 ER ee etl. 38.

Semishrubs, 35-60 cm high; rachis of lower and middle cauline
leaves with lanceolate, acute, usually entire lobes between pri-
mary lobes; capitula-plobdse 3. .2.10:8.52.2.225. SOR ARO
i ae 27. A. gmelini Web. var. messerschmidtiana (Bess.) Poljak.
Perennial, 15—35 cm high; rachis of leaves lacking lobes; capitula
Cain tila Wa 2a ak MOOS 68. A. hololeuca MB.
Rachis of leaf flattened, partly with lanceolate, acute, entire lobes
between primary lobes; terminal lobes of leaves flat, narrowly
lanceolate or oblong-linear, entire, sometimes toothed ........... 39.

432

410

41(38).

+

42.

+

43.

44.

45.

46(35).

Rachis of leaf terete, narrowly linear, usually lacking lobes; terminal
lobes of leaves filiform, filiform-linear or linear-lanceolate ......... 41.
Leaves ternate; capitula comparatively large, 3.5—4.0(5.0) mm wide
Sabie bill beck Mh. tat Neate castes ssecntaa eas 28. A. santolinifolia Turcz.
Leaves twice pinnate; capitula smaller, 2.0-3.5 mm wide ...... 40.
Semishrub, 50-100 cm high; leaves rather large, 3—7(15) cm long,
their lobes and lobules remote, lobes subulate-toothed or entire
iodscawmeaale guaguan Bt Mec Balt senescent sual eit eile abihes 27. A. gmelini Web.
Semishrub, less tall, leaves smaller, with approximate acute lobes
and lobules; lobules often, entire ...........:...0¢¢.-.28<< tees kee
ann. ttesa fda eer en coins 29. A. freyniana (Pamp.) Krasch.
Lower leaves ternate or almost ternate; their terminal lobes filiform

Of filaformy-limeany des ssescd cave sind! Waris baat dbiscens Gersucns dace eee ee 42.
Lower leaves twice ternate; their terminal lobes linear-lanceolate,
subobtuse rs nssauga Salta. deste: bs Loeactnatt nsseimrtyepeantce Oaeeh- edet cas eee 45.
Semishrub, 80-150 cm high; terminal lobes of leaves 6-15 mm
long bs. coseaaenpbetele wae als. abt EN 22. A. abrotanum L.
Semishrubs or perennials, 15—70(80) cm high; terminal lobes of
leaves. short, :2=5(6)ammplonigiceysvscr dessert Perel ea eee de oe 43.

Semishrub, 40-80 cm high; capitula quite numerous, hemispherical,
small: 15=2.0 mmamna tn Gia ¢ cacainccd. csgeedab es. comptcp iiebdeth Ck Gee aes, ee
‘ Secepobenth 22. A. abrotanum L. var. proceriformis (Krasch.) Poljak.
Plant 15-40 cm high; capitula subglobose, large, 2-4 mm in dia

Leaves short arachnoid-hairy or subglabrous, their terminal lobes
with very short, cartilaginous tips; capitula 2.5-4.0 mm in dia.
(Gauicasus) vs dscudieks ii eertok 23. A. chamaemelifolia Vill.
Leaves arachnoid-hairy and punctate-glandular; their terminal lobes
lacking cartilaginous tips; capitula 2.0—-2.5(3.5) mm in dia. (Baikal
jer 0) 1) OP oe eR Ae eee SOREN RSEEEPE MIE om RHA 26. A. adamsii Bess.
Leaves green, punctate, rugose, appressed-hairy; capitula 4-5 mm
in dia, with short peduncles, inclined or drooping .............:sseseeeeees
sunthasdossdalapageme dearest eee 72. A. martjanovii Krasch. ex Poljak.
Leaves dark glaucescent green, glabrous; capitula 2.0—-3.5 mm in
dia; erect OF ime bimed axis avtesiy texas. Wiss ehets- gxdste dA neeeciet nha. ace.
set cease 70. A. obtusiloba var. glabella (Kar. and Kir.) Poljak.
Lower cauline leaves simple, twice or almost thrice pinnately cut,
very rarely undivided, usually punctate-alveolate, concolorous,
green, glabrous or relatively less densely hairy, very rarely when
grayish beneath from dense pubescence, twice or almost thrice
pinnately cut; involucre goblet-shaped or cup-shaped, wider than
long or as long as wide; corolla of peripheral pistillate florets
somewhat shorter and narrower than that of bisexual disk florets

433

47.

48.

49.

50.

54.

411

Leaves simple, entire, shallow-lobed, toothed or deeply pinnatifid
or pinnately cut, lamina green, glabrous or weakly pubescent above,
whitish- or grayish-tomentose beneath, very rarely weakly hairy;
involucre cup-shaped or campanulate, mostly not as wide as long;
corolla of peripheral pistillate florets more than a half as long as
or a fifth to a third as narrow as corolla of bisexual disk florets

TORN LENS IO Shed oT OL LTA A, cson cbt 58.
Leaves weakly hairy above, whitish or grayish beneath from dense
pres cance aie 2),. KAN A AD ARR, fc. ok. 48.
Leaves on both sides glabrous or not densely hairy ............... 50.

Lower leaves twice pinnately cut, terminal lobes lanceolate, usually
1—2 mm wide; uppermost floral bracts simple, lanceolate; involucre
verily. hai: 054.2800. 286.0021... LAS 34. A. armeniaca Lam.
Lower leaves twice or thrice pinnately cut, their terminal lobes
lanceolate-linear or oblanceolate, 0.5—1.5 mm wide, floral bracts
often pinnately cut; involucre whitish-tomentose .................... 49.
Lower leaves 4-9 cm long and 2-7 cm wide, floral bracts usually
exceeding branches of inflorescence and projecting beyond;
capitula large, 4-7 mm in dia ............... 25. A. macrantha Ldb.
Lower leaves 2-5 cm long and 1-3 cm wide, floral bracts smaller,
not exceeding inflorescence; capitula smaller, 2.5-4.0 mm in dia
PUTER, SEDEMTR: MEIGS AMR AeA Rl 24. A. pontica L.
Capitula in simple raceme; involucral bracts blackish- or dark
brown along margin, scarious; florets numerous (45-156)
ee | ee ee 38. A. laciniatiformis Kom.
Capitula in racemose-paniculate inflorescence; involucral bracts
whitish- or brownish-scarious along margin; florets fewer .......

PRL. IRI QARRAL AR ARAL BUA ba. 1 A Ln Al 51.
Capitula less numerous, partly in racemose and partly in racemose-
paniculate inflorescence ihn. oii ean ath 52.
Capitula rather numerous, in paniculate inflorescence ............. 52.

Leaves sessile, usually simple, oblong-lanceolate ...........0....eeee
TREN. titi cs ivcivsi inset sstienathends 39. A. macrorhiza Turcz.
Lower leaves petiolate, twice or almost thrice pinnately cut ..
SO PR NIRS TS 08 0 ND). GREED IA ATOR, SIO SR So.
Lower cauline leaves usually hairy beneath, 2—6 cm long, oblong-
oval, twice or almost thrice pinnately cut, with approximate lobes
manned Meinl is hada a, MILE 37. A. phaeolepis Krasch.
Lower cauline leaves glabrous, 10-18 cm long, oval, bipinnatisect,
with) remote dobes. “and: lobules! ‘i121 ) annie cst Riss cathe.
Gy, . dee, ..ainaduantids, 33. A. remotiloba Krasch. ex Poljak.
Leaves pinnatifid, i.e., their lobes deeply divided, but not up to
midrib, with toothed, acute, usually entire lobules ................. 85.

434

412

56.

Sik

58(46).

+

59:

+

60.

62.

Leaves twice or almost thrice pinnatifid ............ ce ceeeeeeeeeeeees 56.
Stem 90-125 cm high and to 4-5 mm thick; basal leaves long-
petiolate, 25-35 cm long including petiole and 10-15 cm wide
shsdtas Galle Maat reap 32. A. maximovicziana Krasch. ex Poljak.
Stem 15-90 cm high; leaves smaller, usually 4—12(15) cm long
and 2 7.em Wide tack wale ae ee ea 35. A. latifolia Ldb.
Terminal lobe of leaves narrow, 0.5 or about 1 mm wide; capitula
small, usually 2-3 mm in dia .................. 36. A. laciniata Willd.
Terminal lobe of leaves wide, usually 1.5—2.0 mm wide; capitula
4-6 mit diastase. ds seeks cabin eee. ceed Sie 57;
Rhizome producing few, straight, long, vegetative leafy shoots
and usually one fertile stem 70-120 cm high, leaves twice pinnately
cut, their terminal lobes linear-lanceolate, often 4-10 mm long,
entire, scarcely acuminate; achenes almost winged, scaly-ribbed
2G REDE Ail whale vas 30. A. medioxima Krasch. ex Poljak.
Rhizome bearing rosette of basal leaves and single, 45—90(110)
cm-high stem; leaves twice or almost thrice pinnately cut, their
terminal lobes shorter, acute, often with short toothed; achenes
ribeye CRS Pe) A oe eter Ee 31. A. tanacetifolia L.
All leaves or some simple, entire, toothed or shallow-lobed
sated Ch eOhd eee, BRA os colt ne mngnh canter eae ae 59.
Lower and middle cauline leaves once or twice pinnately cut or
deeplyspinnatifid ee iieid.. cei. See ee eee 63.
All leaves narrow-linear, 2—5 mm wide, entire .................csseeeeee
ere E nate ANE I EES OSE 14. A. subulata Nakai.
Lower and middle cauline leaves much wider, usually toothed or

shallowlylobed sc: css. auwannecend eh eee 60.
Middle and lower cauline leaves linear-lanceolate or lanceolate,
acuterfrt uc Fins Ancweicleea eee eee eal ee. LAR 61
Leaves obovate, oblong-cuneate Or Oval ...........ecseesseeseeeeeeeeeee 62;

Middle cauline leaves 1-2 cm wide, more or less deeply divided-
toothed, usually lacking auricles at base ................ceeeceeeeeees
eladeeseabasdeees di QE cansiisctasadsctieweiskeeasd emeeteesem 13. A. integrifolia L.
Middle cauline leaves usually 2.0-2.5 cm wide, small-toothed or
with short, somewhat obtuse teeth, in part entire with auricles at
DASE! «tsucs scree Mee ee eR a 12. A. komarovii Poljak.
Stem 60—100(140) cm high; middle cauline leaves 6-10 cm long
and 3—S cm wide, narrowly ovate, sharp-toothed; capitula narrowly
campanulate pies Shah ea 9. A. stolonifera Maxim.
Stem 20—25(65) cm high; middle cauline leaves 4—6(8) cm long
and 2-4 cm wide, oval or oval-cuneate, short and wide lobed-
toothed; capitula broadly campanulate or subglabrous ...............
os vbiletusupabiapcturk weap ai el faa ae ee ed 11. A. saitoana Kitamura

435

413

63(46). Lobes of middle cauline leaves usually narrowly linear and entire

+

64.

a a
65.

66.

67.

68.

69.

70.

onjmith-emdll shan tecthyssxixl...tlaneee Wave. nis... 64.
Lobes of leaves wider, usually pinnately cut or deeply lobed-
toothed, less often with small teeth or entire ............c cece 67.

All or some lobes of leaves small and sharp-toothed ....................
tinegant, beteneteennunla...yotdl ean 20. A. selengensis Turcz.
Lobes of leaves lacking teeth, usually entire ...............cceeeeee 65.
Capitula campanulate, 3.5-4.0 mm long and 2.0-2.5 mm in dia;
leaves not densely appressed hairy above, lobes 4-6 mm wide
BE Spledioain; ose eda ial cient 19. A. umbrosa Turcz. ex DC.
Capitula narrowly campanulate or almost narrowly cylindrical,
2.5-3.0 mm long; leaves glabrous above, lobes 2-4(5) mm wide
acbadey deireceneny. uhssrs a (idini..d ea See I. AS... 66.
Middle cauline leaves usually 5-8 cm long, their lobes 2—3(4.5)
cm long, narrowly linear, short-acuminate; capitula up to 10—1.2
mm in dia, few-flowered (10—14 florets) ..........ccccccccsssceseesseesseeeees
Laas Sew ;ouids..ures..cl daaus.cns 18. A. lavandulifolia DC.
Middle cauline leaves often 8-12 cm long, their lobes somewhat
longer (4—6 cm long), attenuate-acuminate; capitula 1.2-2.0 mm
in dia, many-flowered, (20-26 florets) ...........cccccccsccessseceeeseeeeeees
ab Niecy ALG QE ARS. LO estd, Reel, wees 15. A. rubripes Nakai
Capitula subglobose or campanulate-globose, usually 4—5(6) mm

long Aidvaleidst.48 much in. dia..cciwa Aciiéialn cath Lameedel veces 68.
Capitula campanulate or ovoid, smaller, 2.0—-3.5(4.0) mm long
cGgembdne Balsewrtal eae cna res Ld RRR PY:

Lower and middle cauline leaves 3—7 cm long and 2-5 cm wide,
deeply pinnatifid into lanceolate, entire or lobes with very few
Rink stivdonbelercy st obese, cis tutewliles esis Stet aR. @ 69.
Leaves large, 8-13 cm long and 7-19 cm wide, pinnately cut,
twice pinnate-pinnatifid; primary lobes incised, with sharp teeth;
teeth simple, entire or with short, acute, small teeth ................ 70.
Lobes of leaves linear-lanceolate, acute, entire or with few, acute,
linear teeth or even (mainly in lower leaves) in turn divided into
2-3 lobes; lamina green above, subglabrous or moderately
ginehunid-haiti: duk:o5..dode shied 4. A. tilesii Ldb.
Lobes of leaves lanceolate, entire or with few small teeth, lamina
greenish-gray above, rather densely arachnoid-hairy ...............
bate, arnallios, Bilin’. seabumeens.: 10. A. koidzumi Nakai
Middle cauline leaves woolly, (8)10-13 cm long and 7-10 cm
wide, pinnately cut, their lobes linear-lanceolate, with fewer acute
teeth, lobes of upper cauline leaves simple ..................008
et EES No Se ann REET ren RaC Dee ere reer 2. A. opulenta Pamp.
All or almost all leaves 6—9.5 cm long and as wide, twice pinnate

436

414

JAR

74.

OP

76.

Uh:

to pinnatifid, lobes lanceolate, deeply pinnatifid; lobes of upper
cauline leaves narrowly lanceolate, with sharp teeth ...............
LRAT datce Eee a ee 3. A. unalaskensis Rydb.
Capitula small, 2.0-2.5 mm long, always remote on long, very

slender, glabrous branches ...............:ccceee 7. A. silvatica Maxim.
Capitula 2.5-4.0 mm long, always aggregated densely on short
branches: .i:.. 020 Cy ak LE td SRT RA he.
Middle and some upper leaves divided in narrowly linear or linear
lobes; capitula’6=4 nim long eRns. Le a 7.
Leaves deeply divided into lanceolate lobes; capitula 2.5—-3.0 mm
Leora 1S. ER CY 75.

Leaf lobes with small and sharp teeth, lower-most pair of lobes
always with short stalk; lamina bright green and glabrous above
Sack RS Pe a aes 8. A. ussuriensis Poljak.
Leaf lobes sessile, entire or with remote short teeth; lamina dull
green above and appressed-hairy or glabrous above ............... 74.
Leaves glabrous or weakly hairy above; lower leaves 12—17(20)
cm long and 6—10(12) cm wide, deeply pinnatifid, their lobes
entire, sometimes with short teeth ............... 17. A. montana Pamp.
Leaves punctate-glandular and more or less densely appressed-
hairy above; lower leaves 6—8(9) cm long and 5-7 cm wide,
pinnately cut, their lobes with short somewhat sharp teeth,
sometimes almost entire ................... 16. A. argui Lev. and Van.
Stem 20—65 cm high, simple, branched only above with short
adpressed branches; lower cauline leaves 5-8 cm long ............ 76.
Stem 65-160 cm high, branched in upper third, branches more or
less long, upward spreading or inclined ...........cccccccccseesseeeeeeees fee
Leaves green above, glabrous or weakly pubescent; lobes of middle
and some upper leaves narrowly linear, mostly 1.5—2.0 mm wide,
ACUIET REL A 6. A. mongolica Fisch. ex Nakai
Leaves dull green or grayish-green above, arachnoid hairy; lobes
of middle and some upper leaves wider, short-pointed apex ........
(YER EY, BRABANT AOSD SE TRIED Mees 5. A. leucophylla Turcz.
Whole plant dull green or grayish, arachnoid-hairy; branches long,
lower 30-40 cm long; leaves on branches with paired small
lanceolate auricles at base, lower leaves with 1-2 pairs of lobe-
like, somewhat obtuse teeth, others mostly narrowly linear or linear-
lanceolate,‘entire; capitula to’S mn long... cae ee
RR A, Bed eA 1. A. vulgaria L. var. jaxartica Poljak.
Plant green, branches usually shorter, upward-spreading; leaves
on branches pinnate or linear, usually lacking auricle; capitula 3—
Amimelong in. ithe, aR, Sh Se 1. A. vulgaris L.

78(13).

83.

84(79).

+

85.

86.

415

Capitula small, 2.5—-3.0 mm in dia, subglobose, in narrowly race-
mose-paniculate inflorescence; stigma lobes apically truncate, with
narrowly linear-cuneate appendage —............cceeeeeeeeeeeee
Lh eehet MARINI IRANS wad shit coe enesonoftbdiVi¥Old 51. A. keiskeana Miquel.
Capitula usually 3.5—8(10—-12) mm in dia, in simple raceme,
corymb, or capitate-corymbose inflorescence; stigma lobes obtuse,
truncate; lacking appendage ..2.........02..0i.cccccscsesesssesssedegseedeosareces 79.
Capitula in simple racemose or spicate-racemose inflorescence,
only some of them apically loosely clustered .............c::csee 84.
Capitula in more or less dense capitate or oblong-ovoid head,
veny rarely in sumpleraceme:., 2.2.0. ici: ok ocdh ss idan RNIN ps seasons 80.
Florets and achenes quite densely covered with long, straight,
erect, white hairs ............... 48. A. leontopodioides Fisch. ex Bess.
Plorets glabrousor weakly hairy «...2.4...:4..85.,.04. ae 81.
All florets with wide, cup-shaped-conical, dark-purple and punctate
glandular corolla, with 5 rather large deltoid recurved teeth ........
eels Bedi dees. .. MENIAG. Aca. 43. A. globularia Bess.
Florets yellow, characters Otherwise ............:::cssceesseesseseeeeees 82.
Capitula fewer, in capitate-corymbose inflorescence; stem usually
10-15(25) cm high; lower leaves 1.5—2.0 cm long ...............
ent Aes URIS, DENG I Ris MEL aden dede. 44. A. glomerata Ldb.
Capitula numerous, in dense oblong-ovoid or globose inflorescence;
latwiris camiaablen ser .tirat ilies Bie E hi Sidadede debe cg Nie 83.
Plant with short, erect, perennial, sterile, woody branches densely
covered with small, overlapping leaves making them appear like
hemispherical or oblong “heads;” leaves 4-7 mm long, ternately
pi viebees }. Gem hs 6... GES BR 47. A. senjavinensis Bess.
Perennial, branches prostrate, not resembling “heads,” lower
cauline leaves 1.0—-1.8 cm long, twice ternately dissected or twice
ternate-palmate or pinnately cut ............... 50. A. triniana Bess.
Plants whitish from dense silky hairs; leaves twice ternately
dissectedto Pirthatipartite 420.4%. .0.4.. KARR a RO ge 85.
Plants glabrous or green, comparatively less densely hairy; leaves
twice pinnately cut or simply ternate .............:cssceeseseeeseeereeeeees 86.
Lower cauline leaves 2—5 cm long, twice ternately dissected
terminal lobes 5—15 mm long; capitula large, 5-8 mm wide .......
SUDA M ARR eRe Qyi28is Mag AA 46. A. insulana Krasch.
Leaves smaller, 0.7—1.2 cm long, twice pinnately dissected, terminal
lobes 1.5—3.0 mm long; capitula smaller ..............ccccscesseseeeseeseeeeeeees
OS FMRI MAN EANIAL, aiid. cs Madre 49. A. lagopus Fisch. ex Bess.
Leaves simple, ternate-palmate; capitula subglobose, 46 mm wide
SOIT. 1 BIA Ad ved PA ol 45. A. furcata MB.

416

+ Leaves twice pinnately dissected; capitula hemispherical, 5—10(12)

mm Widesw14.2.02. amine aaomeceie Real pees ae 87.

87. Corolla in all florets glabrous, punctate-alveolate leaves on both
sides punctate-alveolate ......... 41. A. punctigera Krasch. ex Poljak.

+ Corolla hairy, if glabrous then capitula large, 10-12 mm wide
si biodofaS2da deny MRONE MM ae ad oo Gene ar ce name eis el tennalaanes cae ale 88.

88. Capitula 10-12 mm in dia, flowers numerous, to 150; primary
lobes of lower cauline leaves usually 2-3 pairs ...........eeceseceeeeeees
andanndonche none OOOO. tl eobeetel nel Lamhe eee 42. A. norvegica Fries.

+ Capitula 5-8 mm in dia; flowers fewer (50-70); primary lobes of
lower cauline leaves usually 5-7 pairs ........ 40. A. arctica Less.

Section 1. Artemisia.—Artemisia sect. Abrotanum subsect.
Polycarpicae DC. Prodr. VI (1837) 105.—Herbaceous perennials with
short rhizomes; leaves simple, entire or serrate, with lobe-like teeth or
even pinnately lobed, green and glabrous or weakly hairy above,

438 whitish- and grayish tomentose beneath, less often glabrous on both
sides. Receptacle glabrous; florets glabrous.

Series 1. Vulgares Rydb. in North. Amer. Fl. 34, 3 (1916) 265.—
Lower cauline leaves pinnately cut or deeply pinnatifid into linear-
lanceolate, lanceolate, oblong or broadly-rhombic, more or less toothed
lobes; capitula mostly 3-4(5) mm long, in more or less dense, narrow,
paniculate inflorescence.

1. A. vulgaris L. Sp. pl. (1753) 848; Ldb. Fl. Alt. IV, 52; DC.
Prodr. VI, 112; Ldb. Fl. Ross. li, 585 p. p.; Maxim. Prim. Fl. Amur.
160; Boiss. Fl. or. III, 371; Kryl. Fl. Alt. II, 650; Kom. Fl. Manchzh.
III, 671; Fedtsch. Perech. Rast. Turk. IV, 199; Hegi, Ill. Fl. VI, 2, 637;
Grossh. Fl. Kavk. IV, 141; Krascheninn. in Kryl. Fl. Zap. Sib. XI,
2812.—A. samamisica Bess. Tent. de Abrotan. (1834) 50, pl. Gmel.
Jun.—A. superba Pampan. in Nuov. Giorn. Bot. Ital. n. s. XXXVI
(1930) 473.—Exs.: GRF No. 1618.

Perennial. Rhizome strong, in upper part thickened; stem
herbaceous, erect, (45)65—160 cm high, angular-ribbed, more or less
branched, usually brownish-violet, leafy, weakly hairy. Leaves green
above, glabrous or not densely arachnoid hairy, grayish, arachnoid-
hairy beneath, lower-most leaves petiolate, rest sessile, 3-15 cm long
and 1.5-11 cm wide, deeply pinnately dissected or pinnately cut into
elliptical-lanceolate or linear-lanceolate, usually deeply incised or
toothed, lobes 1-10 cm long and 2.5 cm wide; upper leaves small,
floral bracts entire, linear. Capitula oblong or narrowly campanulate,
3—4 mm long and 1.5-3.0 mm in dia, slightly inclined, less often

439

417

almost drooping, in dense racemes on secondary branches forming in
general, more or less narrow or rather wide panicle. Involucre arachnoid-
hairy, involucral bracts scarious along margin, outer bracts ovate, acuminate,
inner bracts longer, elliptical, obtuse. Receptacle convex. Peripheral pistillate
florets 7-10; their corollas narrowly tubular, stigma lobes narrowly linear,
straight, erect; disk florets bisexual, (5)8—20, their corollas narrow, cup-
shaped-conical, smooth, reddish-brownish; anthers linear, on rather long
filaments, apical appendages of anthers acute, basal appendages
subacute; stigma lobes at maturity slightly exserted from tube, arcuate,
thickened upward, truncate, with long dense cilia.

Forest, partly forest-steppe and steppe zones, in coniferous-de-
ciduous open deciduous forests, around forest glades, edges of forests,
river valleys, ravines, gullies, mountain slopes, meadows, scrubs, fal-
low lands, near fields and dwellings.—European part: Karelia-Lapland,
Dvina-Pechora, Ladoga-IImen, Baltic Region, Upper Volga, Volga-
Kama, Upper Dnieper, Middle Dnieper, Volga-Don, Trans-Volga,
Upper Dniester, Bessarabia, Black Sea Region, Crimea, Lower Don,
Lower Volga; Caucasus: Ciscaucasia, Dagestan, Eastern and Western
Transcaucasia; Western Siberia: Ob River Area, Upper Tobol, Irtysh,
Altai; Eastern Siberia: Yenisei, Lena-Kolyma, Angara-Sayans, Dauria,
Zeya-Bureya; Soviet Central Asia: Lake Balkhash Region, Dzhungaria,
Tarbagatai, Tien Shan, Syr-Darya, Pamiro-Alai Region. General dis-
tribution: Scandinavia, Central and Atlantic Europe, Mediterranean
Region, Balkans-Asia Minor, Mongolia, Dzhungaria-Kashgaria, North
America. Described from Western Europe. Type in London.

Economic Importance. The plant contains essential oil; the re-
covery of the oil from the dry mass from the Saratov Region is 0.3—
0.61% (Kazakevich, 1923). The recovery from Central Asian plants is
lower—not more than 0.03-0.09% (Kudryashev, 1936; Voroshilov,
1941). The oil content of the roots reaches 0.1%. The oil is light
yellow with the mild characteristic aroma of the plant. According to
S.M. Strepkov (1937), the recovery of the oil in plants from Uzbekistan
is 0.061-0.077%, on absolutely dry matter—0.156-0.17%; D.—0.9634;
acid number 7.79; ester number 144.94.

The European oil has the following constants: D?° = 0.907-0.927;
L. from 8 to 19; n>?° = 1.4776-1.485; acid number 1.2; ester number
16-103; ester number of acetylation 50-206. Solubility in 90%
alcohol—1:1. The oil contains: cineole, a-thujone, paraffin and borneol
(cited from Goryaev). Traces of alkaloids have been found in the leaves
and stems (Bankovskii, Zarubina and Sergeeva, 1947). A plant collected
in the flowering stage from the floodplains of the Yenisei River
(Vershinin) contained: 21.3% protein, 24.5% cellulose, 1.9% oil, and
43.2% nitrogen-free extractable substances. Reports about the edibility
of this species for animals are contradictory (Larin, 1956).

440

418

2. A. opulenta Pamp. in Nuov. Giorn. Bot. Ital. n. s. XXXVI
(1930) 464.—A. vulgaris var. kamtschatica Bess. in Nouv. Mém. Soc.
Nat. Mosc. III (1834) 54; DC. Prodr. VI, 113; Kom. Fl. Kamch. III,
156.—A. vulgaris var. communis Ldb. Fl. Ross. II (1844-1846) 586 p.
p.—A. verlotorum auct. non Lamotte: Hult. Fl. Kamtch. IV (1930)
189.—A. koidzumii Nakai in Bot. Mag. Tokyo, XXV (1911) 56 p. p.;
Sugawara, Pl. of Saghal. (1937) 303 and Illustr. Fl. Saghal. ITV (1940)
1823.—Ic.: Sugawara, ibid. tab. 835.

Perennial. Rhizome creeping, woody; stem erect, 80-125 cm high,
angular-sulcate, sparsely hairy or subglabrous, leafy, simple or with
short adpressed branches in upper part. Leaves green and glabrous
above, sometimes weakly pubescent, whitish arachnoid-tomentose
beneath, lower leaves withering early: middle cauline leaves short-
petiolate with short, linear-lanceolate auricles at base, (8)10-13 cm
long and 7-10 cm wide, pinnately cut, their lobes linear-lanceolate,
acute, most often 4—6 cm long, with few, narrow, sharp teeth; upper
cauline leaves mostly with entire lobes, sometimes with occasional
short teeth; floral bracts undivided, lanceolate or linear-lanceolate,
upper-most small, not exceeding inflorescence. Capitula on short
branches, usually densely clustered, sessile, broadly campanulate, 4—5
mm long, in narrow, dense, racemose-paniculate inflorescence, less
often branches elongated. Involucre weakly hairy; involucral bracts
lanceolate, subacute, with wide scarious border; inner bracts somewhat
larger than outer. Receptacle convex. Peripheral florets 7-8, pistillate,
fertile; their corollas filiform-tubular; central disk florets bisexual,
fertile, 14, their corollas narrow, cup-shaped-conical, with recurved
teeth, glabrous; anthers exserted from tube, apical appendages of anthers
cuneate, basal appendages round; stigma lobes arcuate, short-ciliate.
Flowering August.

Slopes of ridges in birch forests, alder forests, meadows, coastal
areas, often near dwellings as a weed.—Far East: Kamchatka, Sakhalin.
General distribution: Northern Japan. Described from Kamchatka. Type
in Florence?

Note. Judging from the diagnosis, A. verlotorum Lamotte, according
to Hultén (1930), is not identical to the plant from Kamchatka. But from
the data presented by Pampanini (1930), Lamotte’s species is found in
China and partly in Japan. As regards Kamchatka, A. opulenta Pampan.,
corresponding to A. verlotorum Hult., is found within its boundaries.

3. A. unalaskensis Rydb. in North. Am. Fl. 34, part 3 (1916) 266;
Hulten, Fl. Aleut. 326 and Fl. Al. X, 1577.— A. wrighti Gray, Proc.
Am. Acad. 19 (1883) 48.—A. vulgaris wrighti (Gray) Hall and
Clements, The North Am. Sp. of Artemisia (1923) 80-87.—A. ursorum

44] Plate XX.
1 — Artemisia stelleriana Bess. Habit, peripheral pistillate floret, style, bisexual
disk florets, stamen, achene. 2 — Kaschgaria brachanthemoides (Winkl.) Poljak.
Upper part of plant, peripheral pistillate floret, bisexual disk floret, stamen,
style, schematic diagram of involucre with general receptacle, achene.

443

420

Hult. Fl. Kamtch. IV (1930) 180.—A. opulenta Pampan. f. laciniata
Pampan. in Nuov. Giorn. Bot. Ital. n. s. XXXVI (1930) 465.

Perennial. Stem up to 1 m high, erect, to 5 mm in dia in middle,
ribbed, glabrous or weakly arachnoid-hairy, simple, leafy. Leaves green
and weakly hairy above, whitish arachnoid-tomentose beneath; all leaves
except uppermost petiolate, middle and some upper leaves 6-8 cm
long, oval or almost round, deeply pinnately divided, their lobes oblong
or lanceolate, more or less deeply incised, lobules narrowly lanceolate,
acute, toothed-incised; upper cauline leaves subsessile, divided usually
into narrowly lanceolate, entire lobes or with few short teeth; uppermost
lobes sessile, entire, linear-lanceolate. Capitula broadly campanulate
or subglobose, to 5 mm long, clustered on very short branches, in
short, narrow and dense paniculate inflorescence. Involucre arachnoid-
hairy; involucral bracts scarious along margin, outer bracts oval,
subacute, inner oblong. Peripheral pistillate florets 5—8; disk florets
bisexual, 20-25. Achenes small, brown, to 1 mm long. Flowering
August.

Meadows, seacoasts.—Far East: Commander Islands, Kamchatka.
General distribution: North America (Aleutian Islands, Alaska).
Described from Unalaska Island. Type in New York.

4. A. tilesii Ldb. in Mém. Acad. Sc. Pétersb. V (1813) 568; Bess.
in Mém. Soc. Nat. Mosc. VIII, 70; DC. Prodr. VI, 113; Rydb. in North
Am. Fl. 34, part 3, 267, excl. syn. A. tilesii arctica and A. tilesii
unalaskensis; Kom. Fl. Kamch. III, 155; Hult. Fl. Kamtch. IV, 186;
Krascheninn. in Spisok Rast. Gerb. Fl. SSSR, X, 84 and in Kryl. FI.
Zap. Sib. XI, 2815.—A. vulgaris var. tilesii Ldb. Fl. Ross. II, 2 (1844—
1846) 586.—A. vulgaris ssp. tilesii Hall. and Clem. in The North
Amer. Spec. of Artem. (1923) 72.—Exs.: GRF No. 3166.

Perennial. Stem few or solitary, arising from more or less thick
rhizome, erect or ascending only at base, simple, (12)15-—35(45) cm
high, more or less densely hairy, mostly above, with fine long entangled
appressed hairs, sometimes completely glabrous below. Leaves
subglabrous and green or even weakly arachnoid-hairy above, grayish,
whitish-tomentose beneath; lower and middle cauline leaves 3-7 cm
long and 2-5 cm wide, pinnately cut into lanceolate or linear-lanceolate,
acuminate lobes, their lobes entire or with very few (1-3) teeth, or
divided (mainly lower) into 2-3 lobules; uppermost leaves simple,
entire, linear. Capitula broadly campanulate or subglobose, 4-6 mm in
dia on long or short peduncles, in dense or lax racemose-paniculate
inflorescence 3—12 cm long. Involucral bracts dark violet-brown along
margin with wide scarious border; outer bracts oval, arachnoid-hairy,
inner narrower, elliptical or lanceolate. Receptacle convex,

44

421

hemispherical, glabrous. Peripheral florets 9, pistillate, corolla nar-
rowly tubular, with 2 teeth, stigma lobes exserted from tube after
anthesis, linear, straight; central disk florets numerous (to 60), bi-
sexual, their corollas cup-shaped-conical, colored, anthers lanceolate,
apical and basal appendages of anthers acute, stigma lobes not ex-
serted from tube, short, linear, erect, short-ciliate. Achenes (immature)
oblong-linear, angular; pappus absent. Flowering August.

Circumpolar zone on coastal, mostly sandy slopes, less often in
dry tundra.—European part: Arctic Europe, Novaya Zemlya, Arctic
Siberia, Chukotka, Anadyr, Kamchatka. General distribution: North
America. Described from Kamchatka. Type in Leningrad.

5. A. leucophylla Turcz. ex C.B. Clarke Comp. Ind. (1876) 162;
Kom. Fl. Manchzh. III, 674; Krascheninn. in Kryl. Fl. Zap. Sib. XI,
2814.—A. leucophylla var. typica f. genuina Pamp. in Nuov. Giorn.
Bot. Ifal. n. s. XXXVI, No. 4 (1930) 414.—A. vulgaris var. leucophylla
Turcz. ex Bess. in Nouv. Mém. Soc. Nat. Mosc. III (1834) 54.—A.
vulgaris &. minor Ldb. Fl. Alt. IV (1833) 82, excl. syn. Tilesii.—A.
vulgaris €. leucophylla Turcz. ex Ldb. Fl. Ross. II, 2 (1844-1846)
585.

Perennial. Rhizome creeping in many-stemmed specimens,
nodulose, with numerous fibrous roots; stem 35-65 cm high, solitary
or few, ribbed, sometimes violet, arachnoid-hairy, with short branches
above. Leaves dull green or greenish-gray and arachnoid-hairy above,
whitish-tomentose beneath, indistinctly punctate-glandular; lowermost
leaves petiolate, 5-8 cm long and 4—7 cm wide, pinnately cut, with
broadly rhombic, coarse-toothed, 1—2 pairs of lobes 0.5—1.0 cm wide;
middle and upper cauline leaves sessile, smaller, less deeply dissected
and with narrower lobes; lower floral bracts also pinnatisect, with
lanceolate or linear lobes, uppermost undivided, linear. Capitula broadly
campanulate, 3-4 mm long and 2-4 mm in dia, aggregated or solitary
on branches, usually forming narrow panicle. Involucre arachnoid-hairy,
greenish or violet; involucral bracts with wide scarious border; outer
bracts oval, inner considerably larger, obovate, subobtuse. Peripheral
florets pistillate with filiform-tubular corolla; disk florets bisexual,
with conical corolla. Flowering July to August.

Steppe slopes, river valleys, in alkaline meadows.—Western
Siberia: Altai (Chui steppe). General distribution: Mongolia, Tibet.
Described from northern Mongolia (Lake Khubsugul). Type in
Leningrad.

6. A. mongolica Fisch. ex Nakai in Bot. Mag. Tokyo, XXXI (1917)
112 and Fl. Sylv. Kor. XIV (1923) 101; Pampan. in Nuov. Giorn. Bot.

445

422

Ital. n. s. XXXVI, 409; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II,
1041; Krasch. in Spisok. Rast. Gerb. Fl. SSSR, X, 83.—A. vulgaris
mongolica Fisch. and y. coarctata Forsell ex Bess. in Nouv. Mém.
Soc. Nat. Mosc. III (1834) 53; DC. Prodr. VI, 113.—A. vulgaris var.
tenuifolia Turcz. ex DC. Prodr. VI (1837) 1113; Ldb. Fl. Ross. II,
585.—A. mongolica var. tenuifolia f. genuina Pampan. op. cit. 409.—
Exs.: GRF No. 3164a, b, c.

Perennial. Rhizome woody; stem erect, (20)40—75 cm high, angular-
sulcate, more or less pubescent or glabrous, simple, with short branches
only in upper part. Leaves green, usually glabrous above, whitish-
arachnoid-pubescence beneath; lower cauline leaves withering early,
long-petiolate; middle leaves 3-9 cm long, deeply pinnately divided,
lobes oblong-lanceolate or lanceolate, more or less deeply divided into
teeth, teeth acute, upper sessile, stipulate, pinnate, lobes mostly simple,
linear-lanceolate, acute, less often with short teeth; uppermost floral
bracts small, simple, linear-lanceolate. Capitula campanulate, about 3
mm long and 1.5 mm in dia, clustered on short branches in narrow
elongated panicle. Involucre arachnoid-hairy-tomentose. Peripheral
florets pistillate, their corollas narrowly tubular; disk florets bisexual
with conical corolla. Flowering August.

Steppes, alkaline meadows, river valleys, southern steppe slopes
of [mud] volcanic cones.—Eastern Siberia: Angara-Sayans, Dauria.
General distribution: Mongolia. Described from Mongolia and Dauria.
Type in Leningrad.

Series 2. Silvaticae Poljak.—Leaves thin; lower cauline leaves
large, deeply pinnately divided into lanceolate, lobes with small sharp
teeth. Capitula small, 2.0—-2.5 mm long, on very long, slender, strongly
inclined branches.

7. A. silvatica Maxim. Prim. Fl. Amur. (1859) 161; Kom.
Fl. Manchzh. III, 670; Pampan. in Nuov. Giorn. Bot. Ital. n. s.
XXXVI, No. 4, 443; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II,
1043.

Perennial. Rhizome short, thickened in upper part; stem erect, 80-
130 cm high, angular, sulcate,:weakly hairy almost simple, leafy, in
upper part with relatively short branches. Leaves green above, glabrous
or weakly pubescent, whitish- and grayish-arachnoid-hairy-tomentose
beneath, lower leaves withering early; middle cauline leaves petiolate,
10-15(19) cm long and 7-11 cm wide, deeply pinnately divided or
lobes lanceolate, small-toothed, upper smaller, with 3-5 narrowly
lanceolate, acute, mostly entire lobes; uppermost floral bracts small,
linear-lanceolate. Capitula pedunculate, broadly campanulate, 2.0—2.5

446

423

mm long and 1.5—2.0 mm in dia, distant, inclined, in lax, pyramidal,
racemose-paniculate inflorescence. Involucral bracts not densely
arachnoid-hairy, smooth, with more or less wide, scarious border; outer
bracts oval, inner ones wide, oblanceolate, larger. Peripheral florets
pistillate, 7, filiform, tubular, very small, with 2 teeth, stigma lobes
strongly exserted from tube, filiform-linear, long, erect, somewhat
inclined; disk florets 10, bisexual, their corollas conical, punctate-
glandular, with convergent or divergent teeth, anthers on short filaments,
linear; apical appendages of anthers acute angled, basal appendages
not distinct; stigma lobes short, arcuate, after anthesis. Achenes to 1
mm long, flattened, narrowly ovoid, dark brown. Flowering August.
(Plate XXI, Fig. 1).

Forest zone, in deciduous , coniferous-deciduous forests, in river val-
leys in bottom-land deciduous forest, sometimes along river banks.—Far
East: Zeya-Bureya, Uda River Area, Ussuri. General distribution: North-
ern China. Described from mouth of Ussuri River. Type in Leningrad.

Series 3. Nutantiflorae Poljak.—Leaves glandular-hairy above
along veins, lower and middle cauline leaves pinnately parted into
elliptical or broadly lanceolate, small-toothed, divided, petiolate lobes;
capitula on short branches, drooping.

8. A. ussuriensis Poljak. nom. nov.—A. nutantiflora Kom. in Izv.
Bot. Sada Akad. Nauk SSSR, XXX, 1-2 (1932) 219 non Nakai.

Perennial. Rhizome short, woody; stems straight, up to 50 cm
high, ribbed, quadrangular in upper part, not densely hairy, with short,
inconspicuous hairs, leafy, simple, with short branches. Leaves green
above, glandular-hairy on veins, yellowish-white and tomentose beneath;
lower, middle and partly upper, cauline leaves 6-10 cm long and 4.0-
7.5 cm wide, with flattened petioles, basally with pinnately parted
auricles; leaves short petiole, broadly lanceolate or elliptical, acute-
toothed; upper floral bracts small, sessile, not exceeding inflorescence,
simple, linear-lanceolate or lanceolate. Capitula sessile, strongly
approximate, narrowly campanulate, 2.5-3.5 mm long, with dense
racemes on branches, forming relatively narrow racemose-paniculate
inflorescence. Involucral bracts arachnoid-hairy, with wide scarious
margin; outer bracts broadly elliptical or oval, inner large, lanceolate.
Peripheral florets pistillate, 8, with narrowly obconical corolla; disk
florets bisexual, fertile, 10; corolla narrowly conical with narrow and
long, deltoid, divergent teeth, weakly punctate-glandular; stigma lobes
not exserted from tube, initially approximate, later strongly recurved,
apically thickened, short ciliate. Flowering August.

447

424

Inundated meadows in forest zone.—Far East: Ussuri. Endemic.
Described from Far East. Type in Leningrad.

Series 4. Simplicifoliae Poljak.—Leaves lanceolate or ovate, en-
tire or with short teeth, or even toothed-lobate, usually with auricles.

9. A. stolonifera (Maxim.) Kom. Fl. Manchzh. III (1907) 676;
Nakai, Fl. Korean. II, 29; Kom. and Alis. Opred. Rast. Dalnevost. Kr.
II, 1043; Hara, Enum. Spermatoph. Japon. II, 126 p. p.—A. vulgaris
var. stolonifera Maxim. Prim. Fl. Amur. (1859) 161; Franch. and Sav.
Enum. I, 239.—A. vulgaris var. stolonifera lusus incana Regel Tent.
Fl. Ussur. (1862) 95.—A. integrifolia var. stolonifera (Maxim.) Pampan.
in Nuov. Giorn. Bot. Ital. n. s. XXXVI (1930) 481.—A. stolonifera
var. laciniata Nakai in Bot. Mag. Tokyo, XXXIV (1920) 53.—Ic.:
Kom. and Alis. Opred. Rast. II, Plate 310.

Perennial. Rhizome creeping, with stolons; vegetative aerial shoots
erect, 30-50 cm high, leafy. Stem 60-100(140) cm high, glabrous or
with appressed hairs, with short adpressed branches in upper part.
Leaves sessile or subsessile, often with short auricles at base, dark
green and glabrous or arachnoid-hairy above, whitish-arachnoid-
tomentose beneath; lower cauline leaves and ones on non-flowering
shoots caducous, middle cauline leaves often 6—9(10) cm long and 3—
4(5) cm wide, broadly lanceolate or obovate, cuspidate, with cuneate
base and small subacute teeth; upper cauline leaves smaller, lanceolate,
entire or with short acute teeth; uppermost leaves simple, narrowly
lanceolate or almost linear. Capitula on narrowly winged peduncles,
4-5 mm long, approximate, in narrow, dense, racemose-paniculate
inflorescence. Involucres arachnoid-hairy; outer involucral bracts ovate;
inner bracts larger, obovate, glabrous, with wide scarious margin.
Receptacle convex. All florets fertile; peripheral pistillate florets 11 or
12, their corollas narrowly tubular; stigma lobes conspicuous, narrowly
linear, straight or weakly inclined; disk florets bisexual, 12-14, their
corollas narrowly conical, glabrous, with recurved teeth; anthers linear,
apical appendages acute, subulate, exserted from corolla, basal
appendages acute; style somewhat shorter than stamen, filiform-linear,
stigma lobes after flowering exserted from corolla, arcuate, thickened,
with divergent cilia. Achenes 1.5 mm long, narrowly ovate, somewhat
flat, sulcate, dark brown. Flowering July to August. (Plate XXII, Fig.
1).

Forest zone on ridges, along slopes, slopes of [mud] volcanic cones
in deciduous forests (often in oak groves), in scrubs, sometimes in
river valleys, meadows, old fields—Far East: Zeya-Bureya (south),
Uda River Area, Ussuri, Sakhalin. General distribution: Japan, China.
Described from Far East. Type in Leningrad.

448

425

10. A. koidzumii Nakai in Bot. Mag. Tokyo, XXV (1911) 56;
Pampan. in Nuov. Giorn. Bot. Ital. n. s. XXXVI, No. 4, 482, excl. var.
mandshurica Pamp.—A. samamisica auct., non Bess. (1834); Hulten,
Fl. Kamtch. IV, 185.—A. vulgaris Kom. in Tr. Spb. Bot. Sada, XXV,
2 (1907) 671, pro parte.—A. vulgaris var. samamisica Maxim. in
Melanges biol. VIII (1872) 535, pro parte.

Perennial. Rhizome woody, slightly thickened in upper part,
producing straight, up to 15 cm high, vegetative leafy shoots. Stem
erect, sometimes arcuate, (40)50—70 cm high, strong, sulcate, arachnoid-
hairy, leafy, almost simple, in upper part with short or somewhat long
branches. Leaves greenish-gray above, rather densely covered with
appressed arachnoid hairs, whitish-tomentose beneath; lower cauline
leaves short-petiolate, usually with pinnately cut auricles, withering
early; middle leaves subsessile, 5-8 cm long, broadly ovate or obovate,
cuneately narrowed toward base, deeply pinnately divided, their lobes
lanceolate, acute, with acute, small teeth; upper leaves sessile, often
with auricles, divided into 3-5, usually entire, lanceolate lobes; floral
leaves in lower part of inflorescence simple, lanceolate, 2-3 cm long,
uppermost smaller, narrowly lanceolate. Capitula campanulate-
hemispherical, 3-5 cm long and 4-7 mm in dia, usually approximate
in more or less dense, less often lax panicle. Involucre whitish,
arachnoid-hairy; involucral bracts oblanceolate, subacute, scarious along
margin; outer bracts somewhat shorter than inner. Receptacle plano-
convex. Peripheral pistillate florets 12-14, narrowly tubular, with 2
teeth; stigma lobes filiform-linear; disk florets bisexual, 42-44, narrowly
cup-shaped-conical, glabrous; anthers linear, on short filaments, their
apical appendages sharply angular, basal appendages almost
undeveloped; stigma lobes after anthesis strongly recurved, densely
ciliate. Flowering August.

Coastal areas, stony slopes on seacoasts, in forests—Far East:
Kamchatka, Okhotsk, Uda River Area, Ussuri, Sakhalin. General dis-
tribution: Japan. Described from Japan. Type in Tokyo.

Plants from Gmelin’s expedition (1773-1774) originating from
Gilyan, and possibly collected by G. von Hoblitz, are preserved in
Leningrad.

11. A. saitoana, Kitamura in Acta Phyt. et Geobot. VII (1938)
63.—A. brachyloba auct. non Franchet; Kom. in Izv. Bot. Sada Akad.
Nauk SSSR, XXX, 1-2 (1932) 218.—Kom. and Alis. Opred. Rast.
Dalnevost. Kr. II, 1043.

Perennial. Plant more or less whitish-tomentose; rhizome creeping,
cordlike, bearing rather long, straight, leafy, nonflowering shoots. Stem
erect, (25)30—65 cm high, simple or with short branches in upper part.

449

426

Leaves sessile or short-petiolate, whitish-tomentose beneath, green
above, not densely appressed arachnoid-hairy; leaves of nonflowering
shoots as also lower and middle cauline leaves 4—5 cm long and 3.5-
4.0 cm wide, broadly ovate, with 5- or 7-lobed, scarcely acute teeth;
upper cauline leaves smaller, with 3—5-lobed teeth; uppermost leaves
simple, lanceolate. Capitula pedunculate, campanulate, 4-5 mm long,
approximate in almost simple raceme or in narrowly racemose-
paniculate inflorescence. Involucral bracts more or less densely covered
with long, arachnoid-hairs, scarious along margin; outer bracts ovate,
inner bracts oblate, along margin violet. Receptacle convex. Peripheral
florets pistillate, 4—5, filiform-tubular, weakly hairy; stigma lobes long,
linear, straight, weakly inclined, exserted from corolla tube; disk florets
bisexual, fertile, 25-26, narrowly conical, with 5 teeth; anthers linear,
apical appendages long subulate after anthesis, basal appendage sharply
angular, anther filaments short; style narrowly linear, long, stigma lobes
after flowering linear, arcuate, at apex funnel-like expanded, with
profuse squarrose cilia. Achenes 0.8—1.1 mm long, ovoid, terete, dark
brown, with longitudinal filiform stripes, apically with roundish smooth
areola. Flowering August.

Forest zone, on dry slopes of [mud] volcanic cones, rock outcrops,
oak groves.—Far East: Ussuri. General distribution: Korea. Described
from Korea. Type in Tokyo.

12. A. komarovii Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVII (1955) 402.

Perennial. Rhizome short, cord-like. Stem erect, 50-80 cm high,
angular-ribbed, appressed-hairy or sub-glabrous, leafy, with short
appressed branches in upper part, sometimes with long branches, stem
almost simple. Leaves glabrous above, almost coriaceous, whitish- or
grayish-arachnoid-tomentose beneath; basal leaves and lower and middle
cauline leaves 4—7 cm long and 2.0—2.5 cm wide, lanceolate, subacute,
entire or with remote short teeth; floral bracts smaller, narrowly
lanceolate, entire. Capitula on short peduncles, upright, campanulate,
3.5—4.0 mm long and 2-3 mm in dia, strongly approximate in dense
panicle. Involucral bracts scarious along margin, colored, outer ovate,
arachnoid-hairy; inner larger, oblanceolate, glabrous. Peripheral
pistillate florets 9, their corollas narrowly tubular; disk florets bisexual,
22, their corollas narrowly cup-shaped, conical, glabrous; anthers not
exserted from tube; stigma lobes arcuate after flowering. Achenes to
1 mm long, narrowly oblong, somewhat flat, brown, finely sulcate.
Flowering August.

Meadows, on dry slopes in forest zone.—Far East: Ussuri.
Endemic. Described from Pacific Coastal area. Type in Leningrad.

450

427

13. A. integrifolia L. Sp. pl. (1753) 848; Bess. Tent. Abrot. 58;
DC. Prodr. VI, 114; Turcz. Fl. baic.-dahur. II, 2, 63; Kom. Fl. Manchzh.
III, 675 p. p.; Kom. and Alis. Opered. Rast. Dalnevost. Kr. II, 1042;
Krascheninn. in Spisok. Rast. Gerb. Fl. SSSR, X, 82; Krascheninn. in
Kryl. Fl. Zap. Sib. XI, 2815.—A. integrifolia var. typica f. genuina
Pamp. in Nuov. Giorn. Bot. Ital. n.s. XXXVI (1930) 478.—A. vulgaris
var. integrifolia Ldb. Fl. Ross. (1844-1846) 585.—Exs.: GRF No.
3163.

Perennial. Rhizome not thick, creeping. Stem simple, single, 50—
100 cm high, weakly ribbed, usually reddish-violet. Leaves simple,
oblong, lanceolate or lanceolate-linear, acuminate, usually deeply lobed-
toothed to different degrees, with 1-3 pairs of erect teeth on each side,
narrowed toward base, glabrous above or indistinctly pubescent, dark
green, finely whitish-tomentose beneath, 5-10 cm long and 1-2 cm
wide; floral bracts smaller, usually entire or weakly toothed. Capitula
in narrowly paniculate or spicate inflorescence, on lateral, more or less
long branches and on main axis of inflorescence on short peduncles,
drooping, oblong-campanulate or campanulate, 3-4 mm in dia.
Involucral bracts green in middle, brownish-reddish-brown on sides;
outer bracts ovate; inner elliptical, along with wide scarious margin.
Peripheral florets 14-15, pistillate; corolla narrowly tubular, with 2
teeth; stigma lobes exserted from tube after flowering, narrowly linear,
straight or curved; disk florets 27-30, bisexual; corolla conical, with
recurved teeth, glabrous; anthers not exserted, lanceolate-linear; apical
and basal appendage of anthers narrowly acute; stigma lobes arcuate
after anthesis, apically thickened, flat, with long, straight cilia.
Flowering August.

Forest zone along banks of rivers, coastal thickets, forest meadows
and along edges of grassy marshes.—Western Siberia: Ob River Area
(Tomsk, Mariinsk); Eastern Siberia: Yenisei, Angara-Sayans, Lena-
Kolyma (southern part), Dauria; Far East: Zeya-Bureya, Uda River
Area. General distribution: Mongolia, Northern China. Described from
Siberia. Type in London.

14. A. subulata Nakai in Bot. Mag. Tokyo, XXIX (1915) 8, non
Willd. ex Steud. Nom. ed. II, 140, nom. nud.; Kitam. Com. Japon. II,
417.—A. lavandulaefolia Nakai Fl. Koreana, II (1911) 29, non DC.—
A. integrifolia var. subulata Pampan. Nuov. Giorn. Bot. Ital. n. s.
XXXVI, No. 4 (1930) 480.—A. stenophylla Kitam. in Acta Phyt. et
Geobot. V. No. 2 (1936) 97, nom. nov. —

Perennial. Rhizome slender, creeping, producing straight, rather
long, leafy, vegetative shoots. Stem straight, 45-60 cm high, glabrous
or weakly pubescent, leafy, simple or with short branches above. Leaves

45

—

428

linear, linear-lanceolate, acute, entire, involute, glabrous above, whit-

ish-arachnoid-tomentose beneath, sessile, or lowermost leaves short-
petiolate, at base with short linear auricles; lower and partly middle
cauline leaves withering early, 5-8 cm long and (1.5) 2-5 mm wide,
adpressed or somewhat inclined; upper cauline leaves smaller;
uppermost floral bracts small, not exceeding inflorescence. Capitula
pedunculate or subsessile, campanulate, 4 mm long, uppermost
somewhat remote, lower in short branches, in narrowly linear, racemose-
paniculate inflorescence. Involucre arachnoid-hairy; involucral bracts
scarious On margin; outer bracts ovate, subacute; inner longer,
oblanceolate. Peripheral florets pistillate, 10-11; disk florets bisexual,
to 11. Flowering August.

Slopes of [mud] volcanic cones, among shrubs in forest zone.—
Far East: Ussuri. General distribution: Northern China, Korea.
Described from Korea. Type in Tokyo.

Series 5. Bipinnatilobae Poljak.—Lower cauline leaves twice
pinnately cut into broadly lanceolate pinnately divided lobes.

15. A. rubripes Nakai in Bot. Mag. Tokyo, XXXI (1917) 112;
Kitam. Compos. Japon. II, 426.—A. venusta Pampan. in Nuov. Giorn.
Bot. Ital. n. s. XXXVI, No. 4 (1930) 470; Kom. and Alis. Opred. Rast.
Dalnevost. Kr. 104; Pampan. in Lavor. Inst. bot. Univ. Cagliar. XL
(1940) 570. —A. vulgaris var. parviflora Maxim. Pr. Fl. Amur. (1859)
160.—A. vulgaris y. vulgatissima Kom. Fl. Manchzh. III, 673; Nakai,
Fl. Koreana, II, 31; Ling Jong in Contrib. Inst. of Bot. nat. Acad.
Peiping.., (323.

Perennial. Rhizome slender, creeping. Stem erect 75-120 cm high,
weakly pubescent or subglabrous, angular-sulcate, leafy, with short
branches in upper part. Leaves green and glabrous above, whitish-
tomentose beneath; lower cauline leaves petiolate, 7-13 cm long, deeply
twice pinnately divided, their lobes broadly lanceolate, acute, lobate-
pinnately divided, lobules linear or lanceolate, acute; middle and upper
cauline leaves smaller, sessile or short-petiolate, pinnately cut, their
lobes narrowly linear, long-acuminate, entire, involute or with
occasional linear acute teeth; floral bracts small, not exceeding
inflorescence, almost linear, acute. Capitula numerous, strongly
appressed, narrowly campanulate, 3.0-3.5 mm long, in narrow, mostly
elongated and dense paniculate inflorescence. Involucres arachnoid-
hairy; involucral bracts lanceolate, subacute, with wide, scarious margin;
inner bracts longer than outer. Peripheral florets pistillate, usually 9-
10, their corollas filiform-tubular; disk florets bisexual, 12-14, their
corollas narrowly cup-shaped-conical, in upper part purple, with

452

429

divergent teeth; anthers exserted from tube, apical appendages of anthers
subulate, basal appendages undeveloped; stigma lobes short, slightly
divergent, almost straight. Achenes small, 0.6 mm long, narrowly ovate,
somewhat flattened, brown, smooth. Flowering August.

Forest zone, in river valleys, among shrubs, meadows, edges of
deciduous forests, as weed in old fields.—Far East: Zeya-Bureya, Uda
River Area, Ussuri. General distribution: Northern China, Korea.
Described from Korea. Type in Tokyo.

Series 6. Angustilobae Poljak.—Leaves pinnately cut or deeply
divided into narrowly linear or lanceolate-linear, entire lobes, or lobes
with short acute teeth, or leaves even entire, narrowly linear.

16. A. argyi Lev. and Van. in Fedde Repert. VIII (1910) 138;
Pamph. in Nuov. Giorn. Bot. Ital. n. s. XXXVI, No. 4, 450-452; Kom.
and Alis. Opred. Rast. Dalnevost. Kr. II, 1036; Kitam. Comp. Japon.
II (1940) 412.—A. vulgaris var. incana Maxim. Prim. Fl. Amur. (1859)
160.—A. vulgaris var. incanescens Franch. in Nuov. Arch. Mus. V, 2
ser. (1884) 169.—A. nutans Nakai in Journ. Coll. Sc. Univ. Tokyo,
XXXI (1911) 33.—A. leucophylla auct. non Turcz.: Kom. Fl. Manchzh.
Ill, 674.

Perennial. Rhizome creeping. Stems erect, 85-120 cm high,
angular-sulcate, grayish-arachnoid-hairy, leafy, with short branches in
upper part. Leaves, except the uppermost, petiolate, punctate-glandular,
dull green above, more or less densely appressed hairy, white-tomentose
beneath; lower and middle cauline leaves 6—8(9) cm long and 5-7 cm
wide, pinnately cut, their lobes lanceolate, with short, subacute teeth
or entire; upper cauline leaves somewhat smaller, pinnate or ternate,
their lobes narrowly lanceolate, entire or with occasional small teeth,
or uppermost floral bracts small, not exceeding inflorescence, linear-
lanceolate. Capitula sessile or on short peduncles, upright, campanulate
or almost oblong, 2.5—3.0 mm long, in narrowly pyramidal panicle.
Involucres arachnoid-hairy; involucral bracts scarious along margin,
colored; outer bracts ovate, somewhat smaller than inner narrowly
obovate. Peripheral florets pistillate, 10, their corollas filiform-tubular,
stigma lobes filiform, linear; disk florets bisexual, 10, their corollas
narrowly campanulate, conical, punctate-glandular; stigma lobes short,
truncate, strongly recurved after anthesis.

Forest zone, in meadows, scrubs, deciduous forests, as well as old
fields—Far East: Zeya-Bureya, Ussuri. General distribution: Northern
China, Korea. Described from northern China. Type in Paris.

453

430

17. A. montana Pamp. in Nuov. Giorn. Bot. Ital. n. s. XXXVI
(1930) 461; Kitam. Comp. Japon. II, 427; Hara, Enum. Spermatoph.
Japon. II, 122.—A. vulgaris var. indica f. montana Nakai in Bot. Mag.
Tokyo, XXXVI (1912) 104.—A. vulgaris var. kamtschatica Nakai op.
cit. (1912) 104, non Bess.—A. gigantea Kitamura in Acta Phyt. et
Geobot. II (1933) 172; Hara Bot. Mag. Tokyo, LII, 4, includ. f. montana
(Nakai), f. electa (Pamp.) and f. shikotanensis (Kitam.) Hara.—A.
shikotanensis Kitamura in Acta Phyt. et Geobot. III (1934) 128.—A.
gigantea var. shikotanensis (Kitam.) Tatewaki, Rep. Veg. Shikotan
(1940) 21.

Perennial. Stem 80—95 cm high, cylindrical, erect, sulcate, leafy,
in basal part glabrous, above weakly hairy. Leaves dark green and
glabrous or weakly pubescent above, whitish-arachnoid-tomentose
beneath; lower and middle cauline leaves petiolate, 12—17(20) cm long
and 6—10(12) cm wide, deeply (but not up to midrib) pinnately divided,
lobes 5, lanceolate, acute, entire or sometimes with short acute teeth;
upper leaves shorter, 3-8 cm long, short-petiolate, simple, lanceolate
or narrowly lanceolate, acute, longer than floriferous branches or
somewhat shorter; uppermost floral bracts small, narrowly linear or
lanceolate-linear, usually exceeding capitula. Capitula subsessile, ovoid,
2.6—3.0 mm long and 1.5—1.8 mm in dia, dense, spicate on branches,
usually forming long paniculate inflorescence. Involucres cup-shaped,
arachnoid-hairy; involucral bracts with wide scarious border; outer
bracts ovate, acuminate, dorsally green; inner large, oblong-ovate.
Receptacle convex, short-clavate, glabrous. All florets fertile; peripheral
florets 8, pistillate, their corollas narrowly tubular; stigma lobes long,
narrowly linear; disk florets bisexual, 5, their corolla tubular-conical,
glabrous, anthers on short filaments, linear, their apical appendages
acute, stigma lobes arcuate after flowering, short, densely ciliate at
apex. Flowering August.

Forest meadows.—Far East: Ussuri, Sakhalin (and Kuril Islands).
General distribution: Japan. Described from Honshu Island. Type in
Tokyo.

18. A. lavandulifolia DC. Prodr. VI (1837) 110; Kom. FI.
Manchzh. III, 678; Pampan. in Nuov. Giorn. Bot. Ital. n. s. XXXVI,
No. 4 (1930) 466 p. p.; Kom. and Alis. Opred. Rast. Dalnevost. Kr.
II, 1041; Kitam. Comp. Japon. II (1940) 411.—A. vulgaris Maxim. in
Bull. Acad. Sc. Pétersb. VIII (1872) 536, pro parte.

Perennial. Rhizome slender. Stem erect, 50-80 cm high, angular-
sulcate, short-pubescent, sometimes subglabrous, leafy, usually with
short adpressed branches in upper part, less often with long, weakly
spreading branches. Leaves glabrous above, grayish-tomentose beneath;

454

431

lower cauline leaves withering early; middle cauline short-petiolate,
5-8 cm long, pinnately cut, their lobes 2.0-3.0 (4.5) cm long and 2-
3(4) mm wide, linear, simple, acuminate, gradually narrowed toward
base; upper leaves sessile, smaller, with fewer lobes (usually 3-5);
uppermost floral bracts small, not exceeding inflorescence, narrowly
lanceolate-linear. Capitula numerous, subsessile, upright, narrowly
campanulate or almost narrowly cylindrical, 2.5—-3.0 mm long, in dense,
narrowly pyramidal or linear-paniculate inflorescence. Involucres short
arachnoid-hairy; involucral bracts lanceolate-elliptical, scarious along
margin. All florets fertile; peripheral florets pistillate, 5—6, their corollas
almost filiform-tubular; disk florets bisexual, 5-6, their corollas
narrowly cup-shaped-conical, glabrous, usually with recurved teeth,
stigma lobes strongly recurved after flowering, not exserted from tube.
Flowering August.

Forest zone, in meadows, scrubs, often in old fields and weedy
places.—Far East: Ussuri. General distribution: Northeastern China,
Korea. Described from China. Type in Geneva.

19. A. umbrosa Turcz. ex DC. Prodr. VI (1837) 111.—A. umbrosa
var. typica Pampan. in Nuov. Giorn. Bot. Ital. n. s. XXXVI, No. 4
(1930) 448; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1041.—A.
vulgaris var. umbrosa Bess. in Nouv. Mém. Soc. Nat. Mosc. III (1834)
52; Turcz. Fl. baic.-dahur. II, 2, 63.—A. selengensis B. umbrosa Ldb.
Fl. Ross. II, 2 (1844-1846) 584.—A. selengensis lusus umbrosa Rgl.
Fl. Ussur. (1862) 96.—A. vulgaris Kom. Fl. Manchzh. III (1907) 673,
pro parte.

Perennial. Rhizome short, slender, woody. Stem 65-95 cm high,
erect, ribbed-sulcate, sparsely covered with inconspicuous appressed
hairs, branched above, sometimes almost simple, leafy. Leaves green
and mildly appressed-hairy above, whitish-arachnoid-tomentose beneath;
basal and, in part, middle cauline leaves 8-12 cm long, short-petiolate;
petioles at base with lanceolate auricles, their lamina broadly ovate or
wide, deltoid-ovate, pinnately cut; lobes linear, 5-7 cm long and 3-6
mm wide, acute, entire or with 1-2 short, acute, linear teeth; upper
cauline leaves divided into 3 lobes; uppermost floral bracts linear,
more or less exceeding inflorescence. Capitula pedunculate or sessile,
campanulate, 3.5-4.0 mm long and 2.0—2.5 mm in dia, densely crowded
on branches, usually forming dense pyramidal inflorescence. Involu-
cres whitish, arachnoid-hairy; involucral bracts elliptical, scarious along
margin; outer bracts smaller than inner. Peripheral florets pistillate, 5—
8, their corollas filiform-tubular; stigma lobes narrowly linear, exserted
from corolla; disk florets 13-18, campanulate, in upper part purple-

455

432

pink, scatteredly glandular, their teeth broadly deltoid, slightly recurved;
anthers somewhat exserted from tube, linear, apical appendages acute,
basal appendages sharply angular; stigma lobes short, oblong-linear,
recurved apically, densely ciliate. Flowering August.

Forest zone in river valleys, wet meadows, scrubs, sometimes along
sandy banks, and in old fields.—Eastern Siberia: Dauria; Far East:
Zeya-Bureya, Uda River Area, Ussuri. General distribution: Mongolia,
northern China. Described from Selenga River. Type in Leningrad.

20. A. selengensis Turcz. ex Besser in Nouv. Mém. Soc. Nat.
Mosc. III (1834) 50; DC. Prodr. VI, 112; Ldb. Fl. Ross. II, 2, 584;
Turcz. Fl. baic.-dahur. II, 2, 62; Maxim. Pr. Fl. Amur. 160; Nakai, FI.
Korean. II, 32; Pampan. in Nuov. Giorn. Bot. Ital. n. s. XXXVI, No.
4, 473; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1041.—A.
vulgaris var. selengensis lusus serratifolia and lusus typica Rgl. FI.
Ussur. (1862) 96.—A. vulgaris var. selengensis Maxim. in Bull. Acad.
Sc. Pétersb. VIII (1872) 536.—A. vulgaris a. selengensis f. serratifolia
Kom. Fl. Manchzh. III (1907) 673.—Exs.: GRF No. 3165.

Perennial. Rhizome 0.4—0.6 cm thick, woody. Stem straight, 75-—
120(130) cm high, light green or reddish brown, usually glabrous,
sulcate, simple or almost simple, leafy. Leaves green and glabrous
above, whitish-arachnoid-tomentose beneath. Lower and in part, middle
cauline leaves withering at anthesis, petiolate, 8-12 cm long, lamina
broadly ovate or almost ovate, pinnately cut, with 5 lobes, less often
7, linear or linear-narrowly lanceolate, acute, mostly 5—7(8) cm long
and 3-5 mm wide, with short acute teeth, in part, entire; upper cauline
leaves sessile, divided into 3-5 lobes or simple, linear narrowly
lanceolate; uppermost floral leaves, short, not exceeding inflorescence,
linear. Capitula pedunculate or sessile, campanulate, 3-4 mm long and
2.0—2.5 mm in dia, strongly crowded, on short, adpressed or weakly
spreading branches, usually forming long, narrow, paniculate
inflorescence. Involucres weakly arachnoid-hairy or subglabrous;
involucral bracts with wide, scarious border, outer bracts roundish or
oval, acuminate, inner larger, broadly oblanceolate, somewhat colored.
Peripheral pistillate florets 8-12, their corollas filiform-tubular; disk
florets bisexual, 10-12; their corollas conical, punctate-glandular;
anthers linear, apical appendage acute, basal appendages roundish-
cuneate; stigma lobes arcuate after flowering. Achenes 1.2 mm long,
ovate, somewhat flattened, brown, smooth, apically with uneven, weakly
stretched margin.

Forest zone in river valleys, in floodplain meadows, along sandy-
gravelly banks, forest glades, burned-over forest areas.—Eastern
Siberia: Dauria; Far East: Zeya-Bureya, Uda River Area, Ussuri.

456

433

General distribution: Mongolia, northeastern China. Described from
vicinity of Selenginsk. Type in Leningrad.

Note. A. selengensis Turcz. is reported for the flora of North
America by Rydberg (Rydberg, North Am. FI., 34, part 3 (1918) 48).
Judging from the diagnosis, the American plant differs from the eastern
Siberian plant by fewer peripheral florets, glandular pubescence and
shape of the corolla, much larger capitula, and the shape and size of
the leaf lobes. On this basis, it can be inferred that this is some other
species of wormwood growing in North America, which is mistakenly
called A. selengensis Turcz. ex Bess.

Section 2. Stellerianum (Rydb.) Poljak. comb. n.; Rydb. in North
Amer. Fl. XXXIV, 3 (1918) 277 pro serie.—Capitula relatively large,
to 8-10 mm wide; anthers with filaments sparsely and not densely
covered with very slender hairs; achenes narrowly oblong-linear,
smooth, in hollow, membranous, semitransparent, saccate pericarp.

21. A. stelleriana Bess. in Nouv. Mém. Soc. Nat. Mosc. III (1834)
79; DC. Prodr. VI, 119; Ldb. Fl. Ross. II, 592; Maxim. in Bull. Acad.
Sc. Pétersb. VIII, 534; A. Gray, Synopt. Fl. North Am. I, 2, 371; Kom.
Fl. Manchzh. III, 668; Rydberg, North Am. Fl. 34, part 3, 277; Hall
and Clements. The Phylogenet. method in Taxon. 67; Pampan. in Nuov.
Giorn. Bot. Ital. n. s. XXXIV, 704; Kom. Fl. Kamch. III, 154; Hult.
Fl. Kamtsch. IV, 185; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II,
1036; Sugaw. Ill. Fl. Kamtsch. IV, 1814; Hara Enum. Spermatoph.
Japonic. II, 125.—A. stelleriana var. sahalinensis Nakai in Bot. Mag.
Tokyo. XXVI (1912) 102.—Ic.: Bess. loc. cit. tab. V; Hall. and
Clements. loc. cit. tab. 5, fig. 1-7; Sugaw. op. cit. 831.

Perennial. Whole plant whitish, hairy-tomentose. Rhizome slender,
creeping or obliquely upright, producing leafy nonflowering shoots.
Stem single, less often 2—3, rather thick (3-5 mm), (15)20-55 cm
high, erect or ascending, leafy. Leaves weakly pubescent above, less
often subglabrous; leaves of nonflowering shoots and lower cauline
leaves petiolate, 3-7 cm long and 1.5—3.0 cm wide, oval or lobate,
lobate-toothed, divided into 3—5(7) oblong-roundish lobes, or scarcely
acuminate; middle cauline leaves short-petiolate, pinnately divided or
pinnately cut, more often with entire lobes; upper floral bracts pinnate,
ternate or undivided, lanceolate-linear. Capitula cup-shaped, 5-8 mm
long and 4-6 mm wide, erect or inclined, pedunculate, all or only
upper capitula crowded in usually dense, spicate inflorescence, but
middle and lower capitula 2—8 on long appressed branches. Involucral
bracts whitish-arachnoid-hairy, linear or narrowly lanceolate, brownish-
scarious, imbricate; receptacle flat. Peripheral florets pistillate, usually

457

434

16, their corollas almost filiform, conical-tubular; stigma lobes dis-
tinct, filiform-linear, erect, subobtuse; disk florets bisexual, 27-30,
their corollas tubular-conical, glabrous or weakly hairy, sometimes at
base hairy; anthers with filaments sparsely covered outside with very
fine hairs visible only at high magnification; style long, thickened in
upper part, with stigma lobes erect or recurved after anthesis, apically
ciliate. Mature achenes narrowly oblong-linear, 3—4 mm long and 0.3-
0.4 mm wide, smooth, dark brown, in membranous, semitransparent,
saccate pericarp to 1.0-1.3 mm wide. Flowering August. (Plate XX,
Fig. 1).

In river mouths and usually along sandy-gravelly seacoasts, some-
times on rubbly slopes near the sea.—Far East: Ussuri, Uda River
Area, Okhotsk, Sakhalin, Kamchatka. General distribution: Scandinavia
(Norway), Japan, North America (Alaska). Described from Kamchatka.
Type in Kiev.

Section 3. Abrotanum Bess. in Bull. Soc. Nat. Mosc. I (1829)
222.—Abrotanum (Bess.) Rydb. North Am. FI. 34, part 3 (1916) 247,
pro subgen.—Semishrubs, less often perennials, with woody tap root,
short, perennial, woody shoots above and more or less numerous, usually
twiglike floriferous stems; leaves twice- or thrice pinnately dissected
into filiform-linear, linear, or linear-lanceolate pinnae, green, glabrous
or hairy beneath, less often whitish from dense pubescence; capitula
globose or hemispherical; receptacle glabrous.

Series 1. Ponticae Rydb. ibid.—Leaf lobes filiform-linear, linear,
or linear-lanceolate.

22. A. abrotanum L. Sp. pl. (1753) 845; Rydb. North Am. FI. 34,
part 3, 280; Hall and Clements. Artemisi: 49; Hegi, III, Fl. VI, 2,
634.—A. paniculata Lam. Encycl. meth. I (1783) 265, non Bess.;
Poljak. in Majevski, Fl. 587. — A. procera Willd. Sp. pl. (1800) 1818;
Bess. in Nouv. Mém. Soc. Nat. Mosc. III, 29; Ldb. FI. alt. 1V, 77; DC.
Prodr. VI, 108; Ldb. Fl. Ross. II, 579; Boiss. Fl. or. III, 370; Kryl. Fl.
Alt. III, 643; Fedtsch. Perech. Rast. Turk. IV, 197; Grossh. Fl. Kavk.
IV, 140; Krascheninn. in Fl. Yugo-Vost. Evrop. Ch. SSSR, VI, 359;
Kryl. Fl. Zap. Sib. XI, 2803.—A. herbacea Willd. Sp. pl. III, 3 (1800)
1828; Bess. op. cit. p. 33; Ldb. Fl. Ross. II, 579.—A. proceraeformis
Krasch. in Otchet o Rabote Pochv.-bot. Otryada Kazakhstan. Eksp.
Akad. Nauk SSSR (1930) 274; Spisok. Rast. Gerb. Fl. SSSR, X, 82;
Fl. Yugo-Vost. Evrop. Ch. SSSR, VI, 360; Kryl. Fl. Zap. Sib. XI,
2802.—Exs.: GRF No. 1024a, 3162 (sub A. procera), Herb. Norm.
No. 4893.

458

435

Perennial. Semishrub, 70-150 cm high, with rather thick, woody
root. Stems erect, woody at base, simple (not branched). Leaves short
appressed hairy beneath, later green, glabrous, petiolate, ovate, usually
4-8 cm long and 3-6 cm wide, twice or thrice pinnately dissected into
filiform-linear, 5-15 mm long and 0.2—1.0 mm wide lobes; floral bracts
less divided to simple pinnate, but uppermost leaves undivided,
generally exceeding inflorescence. Capitula ovoid-globose or
subglobose, 2.0—2.5 mm wide, drooping, crowded in corymbs on lateral
branches of long, narrow, dense paniculate inflorescence. Involucre
weakly short-hairy; outer involucral bracts herbaceous, oblong-elliptical
or oblong-ovate; inner elliptical, with wide, incised, scarious border.
Peripheral florets pistillate, 8, their corollas narrowly tubular, broadened
toward base, bidentate, glabrous, punctate glandular hairy; stigma lobes
exserted, linear, truncate, usually recurved; disk florets 14-16, their
corollas tubular, punctate-glandular hairy, glabrous or with sparse hairs
outside of teeth; anthers slightly exserted from tube after anthesis,
linear, apical appendages sharply angular, basal appendages short,
roundish; stigma lobes shorter, linear, apically thickened, ciliate, after
anthesis spreading. Achenes to 1.2 mm long, ovoid-oblong, somewhat
flattened, sulcate, with a roundish areola at apex, and scarcely toothed
border. Flowering July to August. (Plate XXIII, Fig. 1).

Forest and often forest-steppe zones, along banks of rivers, strips
of riverine deciduous forest, flood-plains, less often wet meadows,
edges of deciduous forests, sometimes as weed in vicinity of
dwellings.—European part: Dvina-Pechora, Ladoga-Ilmen, Volga-
Kama, Upper Volga, Middle Dnieper, Volga-Don, Lower Don, Trans-
Volga, Lower Volga, Black Sea Region; Caucasus: Ciscaucasia south-
ern Transcaucasia; Western Siberia: Upper Tobol, Ob River Area,
Irtysh, Altai; Soviet Central Asia: Aralo-Caspian Region, Lake Balkhash
Region. General distribution: Western Mediterranean Region, Central
Europe, Balkans-Asia Minor, Armenia and Kurdistan. Described from
Dalmatia. Type in London.

Note 1. The confusion relating to the name of this species was due
partly to the appearance after Linneaus of the new names, A. paniculata
Lam. and A. procera Willd. Of these, the second one was accepted by
many authors of the Russian flora despite the absence in fact of a basis
for distinguishing it from A. abrotanum L. This is clearly confirmed
by the collections in the Botanical Institute of the Academy of Sciences
of the USSR. Although Krascheninnikov (Mat. po Istor. Fl. i Rast.
SSSR, No. III, 1958, p. 73) mentions the existence of “intermediate”
forms between A. abrotanum and A. procera, they are so insignificant
(not beyond the limits of an ecological form) that they cannot serve as
the basis for segregating the separate species, A. procera Willd.

46

_—

436

Note 2. A. proceraeformis Krasch. differs from A. abrotanum L.
by the somewhat smaller leaves (2-5 and not 4—8 cm long), the size
of the leaf lobes (not 5—15, but 2-4(7) mm long), a somewhat shorter
stem (not 70-150 but 40-85 cm), and also by the size of the capitula,
i.e., by purely quantitative characters.

Economic Importance. According to L. Kazakevich and O.
Sobolevskaya (1928), the essential oil has the following constants:
D,°-0.9214; D,°-0.9166; a D-9.3°; n,”°-1.4692. Dissolves in a 0.5
volume of 80% alcohol, acid number—0.74—1.32; ester number—
20.46—29.81; ester number after acetylation—55.72-72.58.

According to M.I. Goryaev (1951), the yield of essential oil from
the plants collected near the village of Irgiz was 0.32%. The oil is
orange-yellow with a sharp, pleasant smell. The constants of the oil
are: D,,”°-0.948; n,*°-1.4670; it dissolves in 70% alcohol in the ratio
1:3, in 80% alcohol—1:1 and in 90% alcohol—entirely. Reaction to
aldehyde—positive, reaction to alcoholic solution of FeCl,—negative.

According to I.V. Larin (1956), cattle do not relish this wormwood.

23. A. chamaemelifolia Vill. Pros. 32 and Fl. Delph. III (1785)
250; MB. FI. taur.-cauc. II, 296; DC. Prodr. VI, 108; Ldb. Fl. Ross.
II, 380; Boiss. Fl. or. III, 370; Grossh. Fl. Kavk. IV, 140; Grossh. Opr.
Rast. Kavk. 464.—A. stechmanniana $. armeniaca Bess. in Nouv.
Mém. Soc. Nat. Mosc. III (1834) 35; DC. Prodr. VI, 109; Ldb. FI.
Ross. II, 580.—A. lobelii All. ex Bess. op. cit. 32.—A. chamaemelifolia
Y. armeniaca Hohenack. Enum. Taliisch. (1838) 57.—Ic.: Vill. Fl.
Delph. tab. 35; Rchb. Ic. Fl. Germ. tab. 1034.

Perennial. Root relatively thicker, woody. Stem basally woody,
branched, covered with dark brown bark, producing divergent,
vegetative, leafy shoots, stems few or numerous, erect, brown, short-
pubescent, fertile, with short branches in upper part. Leaves short-
arachnoid-pubescent, hairy or subglabrous, green; basal leaves petiolate,
withering before anthesis, 2.0—-3.5 cm long, thrice pinnately cut, their
terminal lobes linear-filiform, 2-4 mm long, sub-obtuse, with very
short, acuminate, cartilaginous tip; middle cauline leaves sessile, twice
pinnately cut, upper leaves less completely divided; bracteal leaves
undivided. Capitula globose, 2.5—4.0 mm in dia, on peduncles, inclined
or drooping, crowded or not lax, on short branches adpressed to stem,
in narrowly corymbose-paniculate inflorescence. Outer involucral bracts
oblong-ovate, somewhat hairy, with narrow scarious border; inner bracts
somewhat larger, ovate, glabrous, with wide scarious border. Receptacle
glabrous. Peripheral florets 10-12, pistillate, their corollas tubular with
3 obtuse teeth, glabrous, stigma lobes linear, divergent with border,
obtuse; disk florets bisexual, 30-35, their corollas obpyriform with

437

Plate XXI.
Upper part of the plant and lower leaf; bisexual

459

1 — Artemisia silvatica Maxim.,

achene, peripheral pistillate floret, pistil,

’

disk floret, inner involucral bract

stamen. 2 — A. persica Boiss., disk floret, achene.

462

438

convergent teeth; anthers linear, apical appendages of anthers subu-
late-linear, basal appendages less sharply angular; stigma lobes not
exserted from tube, short, linear, with border, obtuse, ciliate. Achenes
prismatic, 5-angled, apically with a flat areola. Flowering August.
Mountain steppes and meadow slopes.—Caucasus: Ciscaucasia,
Eastern, Western, and Southern Transcaucasia, Dagestan. General
distribution: Armenia and Kurdistan, Iran. Type in Paris.

24. A. pontica L. Sp. pl. (1753) 847; Bess. in Nouv. Mém. Soc.
Nat. Mosc. III, 37; Ldb. Fl. Alt. IV, 77; DC. Prodr. VI, 109; Ldb. FI.
Ross. II, 580; Boiss. Fl. or. III, 369; Kryl. Fl. Alt. III, 644; Fedtsch.
Perech. Rast. Turk. 4, 197 p. p.; Rydb. North Am. FI. 34, part 3, 280;
Hall and Clements. Artemisi (1923) 52; Hegi, Ill. Fl. VI, 2, 644; Grossh.
Fl. Kavk. IV, 141; Krascheninn. Fl. Yugo-Vost. Evrop. Ch. SSSR, VI,
359; Kryl. Fl. Zap. Sib. XI, 2803.—A. balsamita Willd. Enum. h.
berol. Suppl. (1822) 57.—A. altaica Desf. Hort. Paris (1829) 160.—
Exs.: GRF No. 3161, 3280.

Perennial. Rhizome creeping, 1.3—3.0 mm thick. Stem 40-100 cm
high, herbaceous, erect, except in lower part densely leafy, with short
hairs. Lower cauline leaves petiolate, others sessile, 2-5 cm long and 1-
3 cm wide, grayish-green above, with scattered appressed hairs, whitish-
or grayish-tomentose beneath, twice or almost thrice pinnately cut, lobes
apically somewhat broadened and then acuminate, 2-6 mm long and 0.5—
1.0 mm wide, leaf rachis with few small, undivided lobes in upper part
between lobes, often lacking them; lower bracteal leaves pinnately cut,
upper undivided, linear, not exceeding inflorescence. Capitula
subglobose, 2.5-4.0 mm in dia, drooping, crowded in narrowly
paniculate inflorescence. Involucres whitish, finely tomentose; outer
involucral bracts herbaceous, oblong-ovate; inner elliptical, with wide
scarious margin. Peripheral florets pistillate, 12, their corollas narrowly
tubular, broadened toward base, punctate-glandular; stigma lobes exserted
from tube, linear, truncate, recurved; disk florets numerous (40-45), bisexual,
in part, undeveloped, their corollas conical, glabrous, less often hairy,
punctate-glandular; anthers on short filaments, linear, apical appendages
of anthers sharply angular, basal appendages almost indistinct; stigma
lobes short, not exserted from tube, short-ciliate, after anthesis recurved.
Flowering August.

Forest-steppe and steppe zones, in thickets of shrubs, birch groves
(small islands in forest-steppe), ravines, river valleys on steppes, and
meadow-saline depressions, steppe slopes.—European part: Volga-
Kama, Volga-Don, Trans-Volga, Middle Dnieper, Lower Don, Black
Sea Region, Bessarabia, Crimea, Caucasus: Ciscaucasia; Western
Siberia: Upper Tobol, Irtysh, Ob River Area, Altai (foothills). General

463

439

distribution: Central Europe, Balkans-Asia Minor; in cultivation in
southern Canada and the USA. Described from Hungary. Type in
London.

25. A. macrantha Ldb. in Mém. Acad. Pétersb. V. (1815) 573;
Bess. in Nouv. Mém. Soc. Nat. Mosc. III, 34; Ldb. Fl. Alt. IV, 76;
DC. Prodr. VI, 109; Ldb. Fl. Ross. II, 581; Fedtsch. Perech. Rast.
Turk. IV, 193; Krascheninn. in Fl. Yugo-Vost. XI, 2805.—Ic.: Ldb.
Ic. Fl. Ross. V, f. 452; Fl. Yugo-Vost. Evrop. Ch. SSSR, VI, fig.
686.—Exs.: GRF Nos. 1226, 3160a, b, c, 3279.

Perennial. Rhizome creeping. Stem woody, solitary or few, erect,
20-100 cm high, except lower part densely leafy, with fine appressed
hairs. Leaves, except basal, sessile, 4-8 cm long and 2-7 cm wide,
almost ovate, densely covered with whitish, fine hairs beneath,
scatteredly hairy and grayish-green above or subglabrous, green, twice,
less often almost thrice pinnately cut, with common pectinate rachis,
i.e., with small undivided lanceolate-linear lobules between lobes; lobes
divergent at acute angle; terminal lobe lanceolate-linear, scarcely
acuminate, sometimes weakly falcate, 2-10 mm long and 0.5-1.5 mm
wide; bracteal leaves pinnately cut, usually exceeding reduced branches
of inflorescence projecting from it, uppermost bracteal leaves undivided,
lanceolate-linear. Capitula subglobose or hemispherical, 4-7 mm wide,
drooping, clustered in corymbs on short branches forming narrowly
paniculate inflorescence. Involucres whitish-tomentose; outer involucral
bracts lanceolate-oblong, herbaceous; inner bracts larger, roundish-
ovate, obtuse, with wide brownish scarious margin. Receptacle plano
convex, smooth. Peripheral florets pistillate, 23, their corollas narrowly
tubular, punctate-glandular, stigma lobes exserted from tube, linear,
truncate, after anthesis arcuate; disk florets bisexual, 34, their corollas
narrowly cup-shaped-tubular, punctate-glandular, hairy outside of teeth,
glabrous below, anthers linear, apical appendage of anthers acute, basal
appendage subacute, stigma lobes after anthesis arcuate, short-ciliate.
Achenes to 2 mm long, oblong-ovoid, sulcate, brown. Flowering August.

Steppe-meadows, thickets of shrubs, near steppe birch groves (small
islands in forest-steppe), edges of pine forests, stony steppe slopes.—
European part: Volga-Kama, Trans-Volga; Western Siberia: Upper
Tobol, Ob River Area, Irtysh, Altai; Eastern Siberia: Yenisei, Angara-
Sayans, Dauria, Lena-Kolyma. General distribution: Northern Mongolia
(Lake Khubsugul area). Described from Siberia. Type in Leningrad.

26. A. adamsii Bess. in Nouv. Mém. Soc. Nat. Mosc. III (1834)
27; DC. Prodr. VI, 107; Ldb. Fl. Ross. II, 577; Maxim. in Bull. Acad,
Sc. Petersb. VIII, 531.

464

440

Perennial. Root relatively slender, woody. Stem woody at base,
usually branched, with dark brown bark, producing short, divergent
vegetative leafy shoots, few, erect, rather slender, usually arachnoid-
hairy, branched, fertile, 15—30(35) cm high shoots. Leaves green,
punctate-glandular, sparsely tomentose; lower and middle cauline leaves
petiolate, 2-4 cm long, their laminas oblong or elliptical, twice or
thrice pinnately cut, terminal lobe filiform, subobtuse, 2—4(6) mm long;
upper leaves sessile, less complexly divided; bracteal leaves linear-
filiform. Capitula globose, 2-4 mm in dia, pedunculate, drooping,
usually crowded on branches, in narrowly paniculate inflorescence.
Involucral bracts with wide scarious border; outer bracts ovate-oblong,
with short hairs, inner roundish or broadly ovate, glabrous. Receptacle
convex, hemispherical, smooth. Peripheral florets pistillate, usually 11-
12, narrowly tubular, somewhat broadened toward base, punctate-
glandular, their stigma lobes slightly exserted from tube, linear,
divergent; disk florets bisexual, numerous (40-44), their corollas
narrowly cup-shaped-conical, glabrous; anthers lanceolate-linear, apical
appendages sharply angular, basal appendages much smaller, acute;
style almost a half as long as stamens, stigma lobes linear, straight,
short-cilliate, divergent. Achenes to 1.2 mm long, oblong-conical,
somewhat flattened, apically with round convex areola, with border on
one side along margin. Flowering August. (Plate XXIV, Fig. 1).

River valleys, lakes, on alkaline and saline-meadow areas, rubbly
and clayey steppe slopes of [mud] volcanic cones and mountains.—
Eastern Siberia: Angara-Sayans, Dauria. General distribution:
Mongolia. Described from Selenga and Baikal. Type in Kiev.

Note. Erroneously reported by Fedtschenko (Perech. Rast. Turk.,
V (1911) 196) for Akmolinsk (now Tselinograd) Region.

Series 2. Vestitae Poljak.—Semishrub; terminal lobes of leaves
linear-lanceolate or lanceolate, margins serrate or entire.

27. A. gmelinii Web. ex Stechm. Dissert. de Artem. (1775) 17;
DC. Prodr. VI, 106; Krascheninn. in Spisok Rast. Gerb. Fl. SSSR. XI,
41; Krascheninn. in Kryl. Fl. Zap. Sib. XI, 2790.—A. gmelinii var.
legitima Bess. in Nouv. Mém. Soc. Nat. Mosc. III (1834) 26.—A.
gmelini Web. var. intermedia (Ldb.) Krasch. op. cit. 2790.—A.
messersch-midtiana Bess. cum var. viridis Bess. op. cit. 27, 28; DC.
op. cit. 107; Nakai, Fl. Korean. II, 31.—A. sacrorum Ldb. in Mém.
Acad. SC. Pétersb. V (1815) 571; Kryl. Fl. Alt. III, 642 p. p.—A.
sacrorum B. intermedia Ldb. FI. Ross. II (1844-1846) 578; Kom. and
Alis. Opred. Rast. Dalnevost. Kr. II (1932) 1042.—A. sacrorum var.
intermedia f. incana Ldb. op. cit. 578.—A. sacrorum var. intermedia
Ldb. f. nivea and var. minor Ldb. f. vestita Kom. Fl. Manchzh. III

441

(1907) 664.—A. sacrorum var. latiloba Ldb. f. incana and f. intermedia
Pampan. in Nuov. Giorn. Bot. Ital. n. s. XXXIV (1927) 686; 688;
Sugawara, Fl. Saghal. IV, 1809.—A. iwayomogi Kitamura in Acta Phyt.
et Geobot. VII, 2 (1938) 64.—Ic.: Gmel. Fl. Sib. II, pl. 56, f. I-V;
Ldb. Ic. pl. Fl. Ross. V (1834) f. 470; Kom. and Alis. op. cit. plate
309.—Exs.: GRF Nos. 3276, 1024.

Perennial. Semishrub, 50-100 cm high. Root woody, thick, to 3
cm, at collar bearing perennial, woody, more or less erect or ascending,
branched shoots, with brownish gray peeling bark. Stems (branches)
herbaceous, rather numerous, erect, dull-violet or brownish, glabrous
or in upper part weakly pubescent. Leaves glandular-hairy, green and
glabrous or weakly pubescent above, grayish- or whitish-tomentose
beneath from appressed dense hairs or on both sides whitish tomentose
(var. messerschmidtiana (Bess.) Poljak.), except upper leaves petiolate,
lower leaves with auricles at base, 3-15 cm long and 1-8 cm wide,
lamina ovate or oblong-ovate, twice pinnately cut, lobes lanceolate,
pinnately cut into lanceolate or linear-lanceolate, serrate- or entire
lobes; part of petioles also with similar lobes; upper cauline leaves
much smaller, petiolate or sessile, uppermost bracteal leaves simple
pinnate or entire, linear. Capitula subglobose, 2.0-3.5 mm wide, droop- *
ing, in short corymbs, aggregated into more or less dense narrow
panicle. Outer involucral bracts densely hairy or subglabrous, lanceolate;
inner bracts elliptical, with wide, scarious border. Peripheral florets
pistillate, fewer (10-12), their corollas narrowly tubular, glabrous,
punctate-glandular, stigma lobes exserted from tube, linear, truncate;
disk florets bisexual, numerous, their corollas conical, punctate-glan-
dular; anthers oblong-lanceolate, apical appendages of anthers obtuse,
basal appendages short, acute, stigma lobes broadly linear, short, at
apex long-ciliate. Achenes 1.5 mm long, narrowly oblong-ovoid or
narrowly conical, longitudinally sulcate, apically with flat, roundish
areola with goffered edge. Flowering August. (Plate XXIII, Fig. 2).

Open coniferous and deciduous forests, forest meadows and
glades.—Western Siberia: Ob River Area, Upper Tobol, Irtysh, Altai;
Eastern Siberia: Yenisei, Angara-Sayans, Dauria, Lena-Kolyma; Far
East: Zeya-Bureya, Uda River Area, Ussuri, Sakhalin; Soviet Central
Asia: Dzhungaria-Tarbagatai, Tien Shan. General distribution: North-
ern China, Mongolia. Described from Lena and Angara rivers. Type in
Berlin, co-type in Leningrad.

Note 1. As regards the specimens issued under No. 3276 (GRF),
Krascheninnikov notes that the original description and the drawing of
A. gmelini Web. ex Stechm. are given in Gmelin’s flora, but the bino-
mial name of this species was used for the first time by Weber, whose
manuscript was used by Stechmann for his monograph on wormwoods.
However, Ledebour, in 1812, while describing a number of Siberian

442

wormwoods, established a new species, A. sacrorum Ldb., which he
later accepted in the Flora of Altai, although this species should have
been identified with A. gmelini Web. ex Stech. The authentic specimen
that evidently has served as the type for Gmelin’s drawing in the Flora
of Siberia convinces me of this. Judging from the label affixed to this
very specimen written in Besser’s own hand, Besser considered this
form to be his own A. gmelini var. legitima Bess. The name proposed
by Ledebour is rather firmly established in the Russian literature. All
the same, according to the rules of nomenclature, the much older name
proposed by Weber must be given preference.

Note 2. Hooker (FI. Br. Ind., 1882) reports this species for west-
ern Tibet; however, on the basis of the description, the Tibetan plant
most probably belongs to A. vestita Wall. ex DC.

28. A. santolinifolia Turcz. ex Bess. in Nouv. Mém. Soc. Nat.
Mosc. III (1834) 87; Krascheninn. in Spisok Rast. Gerb. Fl. SSSR, XI,
43; Krascheninn. in Kryl. Fl. Zap. Sib. XI, 2791.—A. sacrorum var.
minor Ldb. FI. Alt. IV, (1833) 72; Ldb. Fl. Ross. II, 578; Fedtsch.
Perech. Rast. Turk. IV, 196; Krascheninn. in Fl. Yugo-Vost. Evrop.
Ch. SSSR, VI, 358.—A. sacrorum var. santolinifolia Pampan. in Nouv.
Giorn. Bot. Ital. n. s. XXXIV (1927) 693.—A. gmelini var.
biebersteiniana Bess. and var. legitima Bess. op. cit. 26.—A. gmelini
var. turtschaninovianum Bess. op. cit. 87; DC. Prodr. VI, 107.—A.
turschaninoviana Krasn. Spisok Rast. Vost. Tien Shan. (1887) 68.—
A. gmelini Turcz. Fl. baic.-dahur. II (1845) 57.—Ic.: Ldb. Ic. pl. FI.
Ross. V (1834) tab. 471; Gmel. Fl. sib. op. cit. tab. 56 f. 1.— Exs.:
GRF Nos. 3278a, b, c.

Perennial. Semishrub, 12—45(80) cm high, very rarely. 7-10 cm
high (f. pygmaea (Kryl.) Krasch.), with thick woody root bearing
perennial, woody, usually more or less short, branched, spreading,
vegetative shoots covered with dark gray bark; fertile shoots numerous,
erect, straight, ribbed, woody in lower part. Leaves punctate-alveolar
or alveolate-rugose, dark green, glabrous or weakly pubescent above,
grayish- or yellowish arachnoid-woolly beneath; lower leaves with
pinnatisect auricles at base, usually 2—4 cm long and 1-2 cm wide;
lamina ovate, twice or thrice pinnately cut, lobes approximate, terminal
lobes numerous, small, linear or lanceolate, undivided or serrate-
toothed, sub-obtuse; part of rachis between lobes also with similar
lobes; middle and upper leaves smaller, sessile or short-petiolate, twice
pinnately cut; upper bracteal leaves simple pinnate or entire, linear.
Capitula subglobose, 3-5(6) mm in dia, sessile or short-pedunculate,
mature capitula inclined or drooping, crowded on short, upright branches
Or On main axis of stem in narrowly racemose or paniculate
inflorescence. Outer involucral bracts glabrous or pubescent, oblong-

467

443

lanceolate, herbaceous with narrow, uneven, scarious margin; inner
bracts ovate, glabrous, with wide scarious margin. Receptacle plano
convex, smooth. Peripheral florets pistillate, few (10-12), their corollas
narrowly tubular, punctate-glandular, broadened toward base, stigma
lobes linear, truncate, divergent; disk florets bisexual, numerous, usually
52-58, of which some undeveloped, corolla tubular-conical, glabrous,
punctate-glandular, anthers linear, apical and basal appendages of
anthers acute, stigma lobes linear, straight, divergent or slightly inclined.
Flowering August.

Stony, clayey-rubbly slopes of mountains, ascending to timber
line.—European part: Volga-Kama (southern Urals); Western Siberia:
Altai; Eastern Siberia: Angara-Sayans, Dauria; Soviet Central Asia: Pamiro-
Alai Region, Tien Shan, Dzhungaria-Tarbagatai. General distribution:
Mongolia. Described from the Ona River. Type in Leningrad.

Economic Importance. According to the data of O.1. Prokhorova
and I.M. Lebedev (1928), the yield of essential oil was about 1% in
plants collected in the Altai; according to A.F. Gammerman and
Shchupinskaya (1937)—up to 0.56%; and according to S.N. Kudryashev
(1936)—0.1%. According to the studies of G.Z. Chistova (1935), the
oil contains 16% phenols, 14% aldehydes, 19-26% cineole and 5.9%
camphors. Besides, hydrocarbons and alcohols are also found. According
to the data of M.M. Goryaev, G.K. Kruglykhina, M.G. Pugachev, and
I.M. Shabanov (1953), the essential oil in plants collected from the
Dzhungarian Alatau contains 30% carbonyl compounds, mainly as thujone,
13.2% free alcohols, and 19.24% bound esters. In the oil, thujyl alcohol
is the main alcohol; also present are about 2.3% borneol and, possibly,
phenyl alcohol. Besides, the oil contains about 1% cineole, 8% organic
acids (among them the presence of the formic and isovaleric acids has
been confirmed), 9% phenols (mainly t-cresol), terpenic hydrocarbons
and azulene-forming sesquiterpenes (their type not identified), aldehydes
giving only a qualitative reaction (these could not be isolated); azulene
forming sesquiterpenes on dehydrogenation produce azulenes of blue
color. The oil might be a source of thujone and thujyl alcohol.

According to the data of Bankovsky (1941) and P.S. Massagetov
(1947), small quantities of alkaloids are contained in the capitula and
leaves; according to Pekhachek (1948), 100 g of the green herb contains
4-5 mg of vitamin C. Plants collected at anthesis contain 19.0% protein
and very little cellulose (17.7%). Despite some beneficial fodder quali-
ties mentioned in the literature, according to I.V. Larin (1956), this
plant should be considered undesirable in pastures.

29. A. freyniana (Pamp.) Krasch. in Spisok Rast. Gerb. Fl. SSSR,
XI (1949) 42.—A. sacrorum Ldb. var. latiloba Ldb. f. e. freyniana
Pamp. in Nuov. Giorn. Bot. Ital. n. s. XXXIV (1927) 688.—A. sacrorum

468

et

var. viridis f. minor Freyn. in Oest. Bot. Zeitschr. XLV (1895) 345.—
Exs.: GRF No. 3277.

Perennial. Root woody, slender, bearing strongly reduced perennial,
vegetative, woody, branched shoots covered with brownish-gray, peeling
bark; fertile shoots 18—35(45) cm high, slender, virgate, erect, brownish-
violet, glabrous below, weakly hairy above, usually densely leafy, simple
or with short branches. Leaves dark green, punctate-alveolar, glabrous
above, weakly hairy beneath; lower leaves petiolate, withering early;
middle leaves short-petiolate, in part sessile, with pinnately cut auricles
at base, 2-3 cm long and 0.7—1.5 cm wide, oblong-ovate or oblong-
lanceolate, twice pinnately cut, their lobes oblong, approximate,
divergent, pinnately divided, lobes numerous, short, narrowly lanceolate,
acute, entire or with short acute teeth; upper leaves bracteate, smaller,
sessile, simple pinnate, some simple, linear-lanceolate. Capitula globose,
2-3 mm in dia, on short peduncles, inclined or drooping, crowded on

-Short, slender, slightly inclined branches, in narrowly paniculate or

racemose-paniculate inflorescence. Outer involucral bracts glabrous or
subglabrous, small, oblong, herbaceous, with narrowly scarious margin;
inner bracts larger, oval, with narrow (linear) green midrib, remaining
part wide, white-scarious. Peripheral florets pistillate, 6, their corollas
narrowly tubular, gradually expanding toward base, glabrous, punctate
glandular, anthers somewhat exserted from tube, linear; disk florets
bisexual, usually 18, their corollas conical, punctate-glandular; anthers
on short filaments, lanceolate, apical appendages of anthers acute, basal
appendages shorter; stigma lobes linear, straight, divergent, short-ciliate.

Dry southern slopes of [mud] volcanic cones and hills, along edges
of deciduous forests as well as in dry meadows, river valleys, in forest
zone.—Far East: Zeya-Bureya, Uda River Area, Ussuri. General dis-
tribution: Mongolia, northern China. Described from Nercha River.
Type in Florence.

Series 3. Latifoliae Poljak.—Rhizomatous herbaceous perennials;
leaves pinnately cut, twice or almost thrice pinnately cut or divided,
primary lobes lacking petiolule, less often with it, upward spreading.

30. A. medioxima Krasch. ex Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVII (1955) 405.—A. latifolia Maxim. Prim. FI.
Amur. (1859) 159 p. p.

Perennial. Rhizome long, creeping, bearing few vegetative,
divergent, leafy shoots. Stem usually single, 70-120 cm high, erect,
herbaceous, glabrous in basal part, dull violet, weakly hairy above,
leafy. Leaves punctate-alveolar, green above, glabrous or subglabrous,
somewhat pale beneath, scatteredly covered with forked hairs; lower

469

445

and middle cauline leaves petiolate, 6-8(12) cm long and 2-4 cm
wide, their lamina elliptical or oblong-ovate, twice pinnately cut; lobes
lanceolate, petiolate, divergent, deeply pinnately divided into lanceolate
or linear-lanceolate, obtuse, entire lobules; often 4-10 mm long; upper
leaves smaller, sessile, or short-petiolate, pinnate or bipinnate;
uppermost bracteal leaves simple, narrowly lanceolate, inclined, slightly
exceeding inflorescence. Capitula hemispherical, 4-6 mm in dia, on
long peduncles, drooping, more or less remote, lower capitula on long
branches, upper on short, upward-spreading branches and in part on
stem axis, into narrowly lax racemose-paniculate inflorescence. Outer
involucral bracts narrowly elliptical or oblong-lanceolate, hairy; inner
bracts larger, oval, glabrous, or subglabrous, with wide scarious border,
general receptacle smooth. Peripheral florets pistillate, 9, their corollas
narrowly tubular, with 2, teeth, stigma lobes shorter than tube; disk
florets bisexual, numerous (to 35), their corollas tubular-conical, with
5 teeth, glabrous or in upper part scatteredly hairy; anthers lanceolate,
their apical and basal appendages acute, stigma lobes erect, linear,
after anthesis inclined, short-ciliate. Achenes somewhat flattened,
narrow, oblong, brown, to 1.2 mm long, membranous-ribbed. Flowering
August. (Plate XXV, Fig. 2).

Forest zone, deciduous forests and in meadows.—Far East: Zeya-
Bureya, Uda River Area, Ussuri, Sakhalin. General distribution: North-
ern China. Described from Amur. Type in Leningrad.

31. A. tanacetifolia L. Sp. pl. (1753) 843; Willd. Sp. pl. III, 3,
1825; Krascheninn. In Spisok Rast Gerb. Fl. SSSR, X, 77; Krascheninn.
in Kryl. Fl. Zap. Sib. XI, 2806.—A. laciniata Willd. Sp. pl. Ill, 3 (1800)
1843.—A. laciniata &. glabriuscula Ldb. Fl. Alt. TV (1833) 75.—A.
krascheninnikoviana Bess. in Nouv. Mém. Soc. Nat. Mosc. III (1834)
41, pro parte.—A. punctata Bess. ibid., p. 43, pro parte.—A.
macrobotrys Ldb. Fl. Alt. TV (1833) 73; DC. Prodr. VI, 111; Ldb. FI.
Ross. II, 582, pro parte: Kryl. Fl. Alt. III, 647; Rydb. North Am. FI. 34,
part 3 (1916) 264.—Ic.: Gmel. Fl. sib. II, tab. 58.—Exs.: GRF No.
3156a, b, c.

Perennial. Rhizome long, creeping, nodulose, besides shoots bearing
rosettes of basal leaves. Stem usually single, herbaceous, 45—90(110)
cm high, glabrous in lower part, sparsely arachnoid-hairy above, weakly
leafy. Leaves punctate-alveolar, weakly hairy above, with bifid forked
hairs, more densely hairy beneath, petiolate, their lamina oblong-ovate,
4-8 cm long and 2-4 cm wide, twice-, in part almost thrice pinnately
divided, lobes oblong-elliptical, upward-spreading or almost
perpendicular to rachis, sometimes weakly arcuately recurved, more or
less deeply pinnately divided, lobes linear-lanceolate, acute, undivided

470

446

or more or less deeply incised with sharp teeth; basal leaves larger,
long-petiolate, upper cauline leaves sessile, less complexly incised,
uppermost bracteal leaves undivided, lanceolate-linear, short, not
exceeding inflorescence. Capitula globose, 4—5(6) mm in dia, drooping,
in long racemes, aggregated into lax panicle, 20-40 cm long, with
distant and strongly elongated lower branches. Outer involucral bracts
elliptical, hairy, white-scarious along margin; inner bracts larger, oval,
glabrous; with wide brownish-scarious margin. Peripheral florets
pistillate, 11-12, their corollas narrowly tubular, with 2 teeth, punctate-
glandular, stigma lobes exserted from tube, narrowly linear, straight,
slightly recurved; disk florets bisexual, 16-24, their corollas conical,
glabrous, anthers linear, apical and basal appendages of anthers acute,
stigma lobes exserted from tube after anthesis, arcuately recurved, short-
ciliate. Achenes to 1.5 mm long, ovoid-oblong, somewhat flattened,
dark brown, filiformly ribbed, with round convex areola at apex,
undulately bordered. Flowering August.

Open coniferous and deciduous forests, forest meadows, glades,
sometimes on stony slopes.—European part: Dvina-Pechora (low-ly-
ing areas of northern Dvina), Volga-Kama (middle and southern Urals);
Eastern Siberia: Yenisei, Angara-Sayans, Lena-Kolyma, Dauria; Far
East: Zeya-Bureya, Uda River Area. General distribution: North
America (Alaska). Described from Siberia. Type in London.

32. A. maximovicziana Krasch. ex Poljak. in Bot. Mat. Gerb.
Bot. Inst. Akad. Nauk SSSR, XVII (1953) 403; Krascheninn. in Mat.
po Istor. Fl. i Rast. SSSR, II (1946) 137.—A. laciniata var. latifolia
f. maximoviczii Pampan. in Nuov. Giorn. Bot. Ital. n. s. XXXVI (1938)
470.—Ic.: Krascheninn. op. cit. 120.

Perennial. Plant wholly glabrous or weakly pubescent with bifid
forked hairs. Rhizome long, rather thick, stems single or 2-3,
herbaceous, straight, 90-125 cm high, up to 4.5 mm thick, weakly
leafy. Basal leaves long-petiolate, including petiole 25-35 cm long,
and 10-15 cm wide, their lamina ovate or roundish-ovate, pinnately
divided to dissected, lobes oblong-elliptical or lanceolate, short-
decurrent on stalks, more or less deeply divided, lobes lanceolate or
deltoid, sharply serrate-toothed; lower and middle cauline leaves with
shorter petiole, upper sessile, less compound, uppermost bracteal leaves
undivided, linear-lanceolate. Capitula hemispherical, 4-6 mm in dia,
divergent, approximate or somewhat remote on short or elongate
branches adpressed to stem, in narrow, long, paniculate inflorescence.
Involucres glabrous or subglabrous; outer involucral bracts oblong,
herbaceous, narrowly scarious along margin; inner bracts larger, oblong,
obtuse, with wide scarious border. Receptacle glabrous. Peripheral

47

—"

447

florets pistillate, 12-24, narrowly tubular, with 2 teeth; stigma lobes
exserted from tube, linear, truncate, somewhat recurved; disk florets
bisexual, numerous (34-36), their corollas conical, glabrous; anthers
lanceolate-linear, apical appendages of anthers acute, basal appendages
short, roundish; stigma lobes linear, short, divergent after anthesis.
Flowering August. (Plate XXV, Fig. 1).

Ravines, river valleys, and north slopes of [mud] volcanic cones,
in open deciduous forests and meadows. —Far East: Zeya-Bureya, Uda
River Area, Ussuri, Sakhalin. Described from Amur River. Type in
Leningrad.

Note. Outside the USSR, it probably is found in northern China
(Amur River basin).

33. A. remotiloba Krasch. ex Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVII (1955) 404; Krascheninn. in Mat. Ist. FI. i
Rast. SSSR, II, 125.—Ic.: Krascheninn. ibid., Plate 9.

Perennial. Plant wholly glabrous or with occasional appressed hairs.
Rhizome slender, ascending. Stems single or 2-3, straight or weakly
flexuous, 50-65 cm high, slender, ribbed, dull stramineous, almost
simple, weakly leafy. Leaves alveolate-punctate-glandular, except
upper—petiolate; basal and lower cauline leaves 10-18 cm long and
4—9 cm wide, their lamina ovate, twice pinnately cut, lobes 3-5 pairs,
strongly remote, perpendicular to rachis or somewhat obliquely upright,
pinnate, lobules remote, lanceolate-linear, short-acuminate, undivided,
less often with occasional, short, lobate teeth, S—15(20) mm long; middle
cauline leaves short-petiolate, smaller; upper bracteal leaves undivided,
linear. Capitula hemispherical, 5-7 mm in dia, on long peduncles,
drooping, remote, racemose inflorescence. Involucral bracts ovate-
oblong, herbaceous, with brown scarious margin, inner bracts oval or
almost round, with wide scarious margin. Receptacle convex, glabrous.
Peripheral florets pistillate, 10; their corollas narrowly tubular, apically
narrowed, punctate-glandular; stigma lobes narrowly linear, exserted
from tube, recurved; disk florets bisexual, numerous (40-45), their
corollas narrowly cup-shaped-conical, punctate-glandular, anthers lin-
ear-lanceolate, apical appendages of anthers acute, basal appendages
short, roundish, stigma lobes linear, short-ciliate, recurved after anthesis.
Achenes ribbed, brown, apically flat, lacking border. Flowering August.

Open deciduous forests, meadow-steppes.—Eastern Siberia: Lena-
Kolyma, upper part of Aldan River basin. Endemic. Described from
Yakutia, Alga River. Type in Leningrad.

34. A. armeniaca Lam. Encycl. meth. I (1783) 293; DC. Prodr.
VI, 111; Ldb. Fl. Ross. II, 583; Boiss. Fl. or. III, 371; Fedtsch. Perech.

472

448

Rast. Turk. IV, 198; Grossh. Fl. Kavk. IV, 140; Krascheninn. in Spisok
Rast. Gerb. Fl. SSSR, X, 80; Krascheninn. in Fl. Yugo-Vost. Evrop.
SSSR, VI, 358; Krascheninn. in Kryl. Fl. Zap. Sib. XI, 2808; Poljak.
in Majevski, Fl. 587.—A. canescens Willd. Sp. pl. III (1800) 1843;
Bess. in Nouv. Mém. Soc. Nat. Mosc. III, 44.—A. macrobotrys Ldb.
Fl. Alt. (1833) 73.—A. potentillaefolia Fisch. ex Spreng. Syst. veg. III
(1826) 494.—Ic.: Rchb. Ic. ex. Cent. 1 (1823) 6.—Exs.: GRF Nos.
978,,, 3159:

Perennial. Rhizome long, creeping. Stem single, less often few,
herbaceous, ribbed, 40-100 cm high, sparsely leafy, grayish above
from fine appressed hairs. Leaves scatteredly hairy and grayish-green
or green above, densely beneath, whitish from long, bifid, appressed,
silky hairs; basal leaves usually withering before anthesis; lower cauline
leaves long-petiolate, their lamina oblong-ovate, 5-10 cm long and
2.5-5.0 cm wide, twice pinnately cut; lobes oblong-elliptical, upward-
spreading, deeply pinnately incised or divided, terminal lobes 3-10
mm long, 1-2 mm wide, lanceolate, with long almost awn-like acuminate
apex, undivided or with few teeth; middle and upper leaves smaller,
sessile, upper bracteal leaves undivided, lanceolate-linear, not exceeding
inflorescence. Capitula globose, 3-4 mm in dia, on short peduncles,
drooping or divergent, crowded on more or less short, weakly divergent
branches, in narrow panicle. Involucral bracts weakly hairy; outer bracts
almost oval, herbaceous, with narrowly scarious margin; inner roundish
oval, obtuse, with wide scarious, slightly incised brownish margin.
Peripheral florets pistillate, 10; their corollas narrowly tubular, with 2
teeth, punctate-glandular; stigma lobes exserted from tube, short, linear,
truncate; disk florets bisexual, 22; their corollas conical, glabrous,
anthers included, on short filaments, linear, apical appendages of anthers
acute, basal appendages short, obtusely acuminate, stigma lobes short,
linear, straight, divergent, short-ciliate. Achenes to 3 mm long, oblong,
somewhat flattened, apically convex, scarcely broadened. Flowering
August.

Plains in meadow and feathergrass-herb steppes, occasionally in
meadow depressions, less often on mountain slopes.—European part:
Volga-Don, Lower Don, Middle Dnieper (Samara River), Trans- Volga,
Volga-Kama; Western Siberia: Upper Tobol, Irtysh, Altai; Caucasus:
Eastern and Southern Transcaucasia, Dagestan. General distribution:
Central Europe (Hungary, Transylvania), Armenia and Kurdistan.
Described from Armenia. Type in Paris.

35. A. latifolia Lam. in Mém. Acad. Petersb. V (1815) 569; Ldb.
Fl. Alt. IV, 70; DC. Prodr. VI, 110, pro parte; Ldb. Fl. Ross. II, 582;
Maxim. Prim. Fl. Amur. (1859) 159; Kom. Fl. Manchzh. III, 667;
Fedtsch. Perech. Rast. Turk. IV, 198; Krascheninn. in Fl. Yugo-Vost.

473

449

Evrop. Ch. SSSR, VI, 357; Kryl. Fl. Zap. Sib. XI, 2810; Poljak. in
Majevski, Fl. 587.—A. punctata B. stricta Bess. in Nouv. Mém. Soc.
Nat. Mosc. III (1834) 43.—A. krascheninni-koviana Bess. Ibid. p. 41,
pro parte.—Ic.: Ldb. Ic. pl. Ross. V, tab. 469.—Exs.: GRF No. 3282a, b.

Perennial. Plant entirely glabrous, less often weakly hairy (f.
pilosiuscula Krasch.). Rhizome long, creeping, woody, apically as-
cending. Stem usually single, herbaceous, straight, ribbed, 15-80 cm
high. Leaves alveolate-punctate-glandular; basal and lower cauline
leaves long-petiolate, their lamina oblong-elliptical, 4-12 cm long, 2-
7 cm wide, pinnately divided, lobes somewhat divergent above, at
base decurrent on petiole, lanceolate or lanceolate-ovate, deeply in-
cised but not up to midrib into teeth-like, 2-15 mm long and 1-3 mm
wide lobules, with almost spine-like acuminate apex, entire, less often
with 1-2 teeth; upper leaves simple pinnate, with serrate-toothed or
entire secondary lobes; upper bracteal leaves linear-lanceolate, not
exceeding inflorescence. Capitula pedunculate, subglobose, 3—5 mm in
dia, approximate, drooping, on more or less short, erect branches, in
narrowly paniculate inflorescence. Involucral bracts usually glabrous;
outer bracts herbaceous, oval, with scarious incised margin; inner
somewhat wider, membranous. Receptacle, hemispherical, glabrous.
Peripheral florets pistillate, 9; their corollas narrowly tubular, with 2
teeth, punctate-glandular, stigma lobes exserted from tube, broadly
linear, truncate, divergent or recurved; disk florets bisexual, rather
numerous (up to 26), their corollas tubular, glabrous; anthers on short
filaments, linear, apical appendages of anthers acute, basal appendages
short, round; stigma lobes short, linear, apically short-ciliate, somewhat
divergent after anthesis. Achenes 1.4 mm long, oblong-prismatic,
somewhat flattened, with edges, brown, at apex with round convex
areola, weakly bordered. Flowering July.

Steppes, less often alkaline meadows, in feather-grass-herb steppes,
open birch forests, along edges of steppe pine woods, in forest zone
in larch-pine forests, in south-facing meadows, sometimes rubbly and
calcareous slopes — European part: Dvina-Pechora (Pinega River), Volga-
Kama, Volga-Don, Trans-Volga; Western Siberia: Upper Tobol, Irtysh,
Ob River Area (southern edge), Altai; Eastern Siberia: Angara-Sayans,
Yenisei, Dauria; Soviet Central Asia: Aralo-Caspian Region. General
distribution: Mongolia. Described from Siberia. Type in Leningrad.

Series 4. Laciniatae Kitamura in Acta Phyt. et Geobot. VIII (1939)
65.—Leaves twice or almost thrice pinnately cut, lobes petiolate,
divergent almost at right angle to petiole.

36. A. laciniata Willd. Sp. pl. II, 3 (1800) 1843, excl. var.; Bess.
in Nouv. Mém. Soc. Nat. Mosc. III, 40; DC. Prodr. VI, 110; Ldb. FI.

474

450

Ross. II, 581; Turcz. Fl. baic.-dahur. II, 2; Maxim. in Bull. Acad. St.
Pétersb. VIII, 530 p. p.; Kom. Fl. Manchzh. III, 665; Fedtsch. Perech.
Rast. Turk. IV, 198; Pampan. in Nuov. Gior. Bot. Ital. n. s. XXXV,
671 p. p.; Hegi, Ill. Fl. VI, 2, 641; Kom. and Alis. Opred. Rast.
Dalnevost. Kr. II, 1042; Kryl. Fl. Zap. Sib. XI, 2806.—A. laciniata
glabriuscula Ldb. Fl. Alt. IV (1833) 75.—A. laciniata f. paniculata
Kryl. Fl. alt. III (1904) 646.—Ic.: Gmel. Fl. sib. II, tab. 57.—Exs.:
GRF Nos. 1127, 3157, 3158, 3283.

Perennial. Rhizome more or less thick, creeping or obliquely as-
cending. Stems few, herbaceous, erect, ribbed, 50-70(90) cm high,
branched above, sparsely leafy, usually glabrous below, appressed-
hairy above. Leaves green, alveolate-glandular, glabrous or scatteredly
hairy above, densely hairy beneath, with distinct longitudinal veins
and thick border along margin; lower and middle cauline leaves long-
petiolate, their laminas almost as long as petiole, 3-12 cm long and
1.5-5.0(8.0) cm wide, oblong-elliptical, twice- or almost thrice- pin-
nately cut, lobes oblong or oblong-elliptical, sometimes slightly curved,
divergent almost at right angle to rachis but lower lobes often even
recurved; petioles at base somewhat thickened and decurrent, lobes
also strongly inclined, oblong-lanceolate or lanceolate-linear, entire or
with aristate-acuminate secondary lobes or teeth; upper leaves sessile,
twice pinnately cut; bracteal leaves undivided except lower, linear,
short. Capitula subglobose or hemispherical, 2-3 mm in dia, drooping
approximate or somewhat remote on branches adpressed to stem, in
narrowly paniculate inflorescence. Involucres glabrous or weakly hairy;
outer bracts oblong-ovate, greenish, with narrow scarious border, in-
ner bracts oval, obtuse, with wide scarious border; common receptacle
hemispherical, glabrous. Peripheral florets pistillate, 12, their corollas
narrowly tubular, with 2 teeth, punctate-glandular, hairy; stigma lobes
exserted from tube, linear, erect, truncate, deflected; disk florets bi-
sexual, numerous (38-40); their corollas narrowly conical, weakly hairy
below teeth; anthers lanceolate, apical and basal appendages of an-
thers short, acute; stigma lobes short, linear, divergent after anthesis.
Achenes oblong-ovoid, somewhat flattened, finely sulcate, dark brown,
apically with round areola, and edges. Flowering July to August.

Forest-steppe with bordering parts of forest and steppe zones,
solonetzic meadows, meadow slopes, among scrub and open decidu-
ous forests.—European part: Volga-Don (near Bobrov); Western
Siberia: Upper Tobol, Irtysh, Altai, Ob River Area (southern part);
Eastern Siberia: Yenisei, Angara-Sayans, Dauria, Lena-Kolyma; Far
East: Zeya-Bureya, Ussuri. Kamchatka. General distribution: Central
Europe, Mongolia, Northern China. Described from Siberia. Type in
Berlin.

477

451

Note. Hooker (Fl. Br. Ind., 1882) reports this species for western
Tibet. Judging from his description, the Tibetan plant cannot be re-
lated to our species.

37. A. phaeolepis Krasch. in Sistemat. Zam. Gerb. Tomsk. Univ.
1-2 (1949); Krascheninn. in Kryl. Fl. Zap. Sib. XI, 2808.—A. laciniata
Q. turczaninowiana Bess. in Bull. Soc. Nat. Mosc. [IX (1836) 48.—
A. laciniata B. turczaninowii DC. Prodr. VI (1837) 111.—A. laciniata
var. glabriuscula f. marginata Pamp. in Nouv. Giron. Bot. Ital. n. s.
XXXIV (1927) 673, pro parte.—A. laciniata f. racemosa Kryl. and f.
tomentosa Kryl. Fl. Alt. III (1904) 645, ex parte.

Perennial. Rhizome slender, creeping or obliquely ascending. Stem
often single, less often 2-3, herbaceous, erect, ribbed, 5—40 cm high,
scatteredly hairy in lower part or glabrous, densely hairy above with
adpressed hairs, sparsely leafy, less often stems and leaves whitish-
tomentose (var. tomentosa (Kryl.) Krasch.). Leaves alveolate-glandular,
usually glabrous above, long-hairy beneath, with distinct longitudinal
veins and thick border along margin; basal and lower cauline leaves
petiolate, their lamina oblong-ovate or elliptical, shorter than petiole,
2-6 cm long and 1.5-3.0 cm wide, twice or almost thrice pinnately
divided, lobes oblong-elliptical, divergent almost at right angle, rachis
somewhat thickened, at base fused with equally thickened petiole,
secondary lobes obliquely divergent, lanceolate, often undivided, less
often divided into small toothed lobes and together apically
cartilaginous, acuminate; upper leaves sessile, once or twice pinnately
cut, uppermost leaves bracteal, short, linear, undivided with few teeth.
Capitula few, hemispherical, 4-6 mm in dia, drooping, in simple long
raceme or less often on short branches in narrowly racemose paniculate
inflorescence. Involucres glabrous or subglabrous; outer involucral
bracts oblong ovate, with narrow brown scarious border; inner bracts
oval, obtuse with wide scarious border. Receptacle convex,
hemispherical, glabrous. Peripheral florets pistillate, 14, narrowly
tubular, with 2 teeth, punctate-glandular, stigma lobes exserted from
tube, linear, truncate, divergent or recurved; disk florets bisexual,
numerous (to 60), some sterile, corolla cup-shaped-conical, hairy and
punctate-glandular; anthers lanceolate-linear, apical appendages of
anthers acute, basal appendages shorter, subacute; style short; stigma
lobes considerably shorter than tube, short-linear, divergent or slightly
deflected, apically short-ciliate. Flowering July to August.

Mountain river valleys, on slopes, often stony places high in
mountains, up to treeline or slightly above, on open southern slopes.—
Western Siberia: Altai, Angara-Sayans. General distribution: Mongolia,
Tibet.— Described from Altai. Type in Leningrad.

452

Plate XXII.

475
pper part of plant, lower cauline leaf,

middle cauline leaf, peripheral pistillate floret, bisexual disk floret, style with
stigma lobes, stamen, outer involucral bract, inner involucral bract, receptacle,
achene; 2—A. rutifolia Steph., habit, achene, bisexual disk floret.

1 — Artemisia stolonifera Maxim.,

478

453

Note. A doubtful species, which should be considered as a separate
mountain race of A. laciniata Willd., distinguished by the less compound
inflorescence (racemose-panicle) and more numerous bisexual florets.

38. A. laciniatiformis Kom. Fl. Kamch. III (1930) 153;
Krascheninn. in Mat. po.Istor. Fl. i Rast. SSSR, II (1946) 136, 128.

Perennial. Root vertical, woody, with more or less elongate rhizome.
Stems few, distant or approximate, 25-40 cm high or 7-10 cm (ssp.
taimyrensis Krasch.), erect, sometimes arcuately bent downward, simple,
leafy, more or less pubescent. Leaves on both sides with long occasional
hairs, denser and more appressed beneath, punctate-glandular; lower
cauline and basal leaves long-petiolate, including petioles to 7-8 cm
long, and 2.0-3.5 cm wide, their lamina ovate, twice pinnately cut,
rachis absolutely entire, pinnae oblong or obovate, pinnately cut, lobes
lanceolate-linear, short-acuminate; upper cauline leaves smaller,
subsessile, pinnately cut; uppermost leaves bracteal, undivided, linear-
lanceolate. Capitula in simple raceme, hemispherical, 4-7 mm in dia,
drooping, lower capitula remote, on long peduncles, upper crowded on
short peduncles. Involucral bracts oval, with wide, brown, scarious,
unevenly incised margin; outer involucral bracts green along midrib,
weakly hairy; inner bracts glabrous. Receptacle convex, hemispherical,
glabrous. Disk florets bisexual, narrowly conical, with hairy or glabrous
teeth. Achenes up to 1.2 mm long, oblong, indistinctly ribbed, apically
with round areola bordered by upper margin of pericarp. Flowering
July to August.

Floodplains of rivers and above, on high terraces of tundra, at
southern border of arctic and forest-tundra zones.—Arctic: Arctic
Siberia (Khatanga River, Taimyr Peninsula); Eastern Siberia: Lena-
Kolyma (Kuma-Sur); Far East: Kamchatka. General distribution: North
America (Yukon). Described from Kamchatka. Type in Leningrad.

Note. The plants of this species growing in Arctic Siberia are
always distinguished from the Kamchatka plants by the hairy corolla
and lower height; among them ssp. taimyrensis Krasch. deviates the
most from A. laciniatiformis Kom.

Series 5. lanceolatae Poljak.—Leaves undivided, oblong-
lanceolate.

39. A. macrorhiza Turcz. in Bull. Soc. Nat. Mosc. XIII (1840)
43;-Ldb. Fi. Ross: II; 590.

Perennial. Plant green, weakly hairy. Rhizome creeping. Stem 25—
35 cm high, herbaceous, curved, weakly leafy, branched, with slender
elongated upright branches. Leaves undivided, sessile, 1.5-2.0 cm long,

479

454

oblong-lanceolate, glabrous; upper leaves smaller, lanceolate. Capitula
on long, in part on short peduncles, in simple racemose inflorescence,
subglobose, 4-6 mm wide, divergent or drooping. Involucral bracts
round-elliptical, herbaceous, glabrous, with wide scarious border.
Receptacle convex, hemispherical, glabrous. Florets glabrous, punctate-
glandular; peripheral florets pistillate, 12, their corollas narrowly
tubular, at base slightly broadened, membranous, with 2 teeth, stigma
lobes exserted from tube, narrowly linear, almost straight, divergent;
disk florets bisexual 35, fertile, in part sterile, their corollas tubular-
conical, with 5 teeth, anthers on long filaments, linear, apical appendage
of anthers acute, stigma lobes almost arcuate after flowering, short
ciliate.

Gardens.—Eastern Siberia: Angara-Sayans (vicinity of Krasno-
yarsk). Endemic? Described by N. Turczaninow from his own
collections from Siberia. No more has been found anywhere so far.
Type in Leningrad.

Note. It is well separated by the simple, oblong-lanceolate leaves.

Series 6. Norvegicae Rydb. North Am. FI. 34, part 3 (1916) 261.—
Capitula many-flowered, in simple racemose inflorescence.

40. A. arctica Less. in Linnaea, VI (1831) 213; DC. Prodr. VI,
119; Ldb. Fl. Ross. 591; Rydb. North Amer. Fl. 34, part 3, 263; Hult.
Fl. Kamtsch. IV, 176, exl. syn.; Kom. Fl. Kamch. III, 150; Krascheninn.
in Sov. Bot. No. 5, 7.—A. arctica ssp. subarctica (Krasch.) Hult. in
Nytt. Mag. Bot. 3 (1954) 67.—A. arctica var. ochotensis (Bess.) Kom.
op. cit. 151.—A. arctica var. beringensis Hult. op. cit. 67.—A.
longepedunculata Bess. and A. chamissoniana Bess. in Nouv. Mém.
Soc. Nat. Mosc. III (1834) 177.—A. montana Schlecht. in herb. reg.
berol. ex Ldb. Fl. Ross. II (1845) 591.—A. norvegica Spreng. Syst.
veg. III (1826) 490, non Fries; Maxim. in Bull. Acad. Sc. Pétersb.
VIII, 533 p. p.; Kom. Fl. Manchzh. III, 668; Sugaw. Ill. Fl. Sachal. IV,
1813.—A. norvegica pacifica A. Gray Synops. Fl. North. Am. I (1884)
371.—A. norvegica Hall and Clements. The Phylogen. meth. (1923)
57, non Fries.—A. subarctica Krasch. in Bot. Mat. Akad. Nauk SSSR,
IX (1946) 176.

Perennial. Root woody, thick, with very short, perennial, sterile
shoots in upper part bearing leaf rosettes. Stems few or single, 25—40
cm high, erect, sometimes arcuately bent, glabrous or hairy, leafy.
Leaves glabrous or hairy; basal leaves and leaves of sterile shoots
long-petiolate, most often 5-8 cm long and 2—3(4) cm wide, twice
pinnately cut lobes 5-7, remote, terminal lobe 3—7 mm long and 1.5-
3.0 mm wide, linear-lanceolate more or less acute, simple, or with

480

455

small acute teeth; middle cauline leaves sessile or subsessile, pinnately
cut, uppermost leaves bracteal, simple, narrowly linear. Capitula in
simple long (up to 10-17 cm long) raceme, hemispherical, S—8 mm in
dia, divergent or drooping, remote, lower capitula on long (to 3-5 cm)
peduncles, upper capitula on shorter peduncles. Involucral bracts
lanceolate-ovate or elliptical, with dark brown scarious margin.
Receptacle convex, hemispherical, glabrous. Peripheral florets pistillate,
12, fertile, their corollas tubular, membranous, hairy, stigma lobes
linear, obtuse, divergent after anthesis; disk florets bisexual, numerous
(50-70), fertile, in part immature, their corollas narrowly conical,
yellow, with dense, long, straight white hairs; anthers on short filaments,
broadly linear, acute, stigma lobes slightly curved after flowering,
divergent, at apex ciliate. Achenes to 2.5 mm long, ovoid-oblong,
angular; pappus lacking.

Mountain, subarctic, partly arctic, and maritime meadows.—Arctic:
Arctic Siberia; Eastern Siberia: Lena-Kolyma; Far East: Kamchatka,
Okhotsk, Sakhalin (Kuril Islands). General distribution: North America.
Described from Aleutian Islands.

Note. A. arctica ssp. subarctica (Krasch.) Hult. is distinguished
by the fewer lobes of the lower leaves (usually 5 and not 7); var.
ochotensis Bess. is distinguished by the wider pinnules of the leaves
and the large sessile capitula.

41. A. punctigera Krasch. ex Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVII (1955) 408.—A. norvegica auct. non Fries:
Sugaw, Ill. Fl. Saghal. (1940) 1813.

Perennial. Root rather thick, woody, with perennial woody, short,
sterile shoots covered with remains of petioles, bearing rosettes of
leaves; fertile shoots usually 2—3, erect, sometimes arcuately bent, 20—
40 cm high angular-sulcate, smooth, sparsely leafy, more or less simple.
Leaves glabrous or usually hairy, on both sides punctate-alveolar; basal
leaves and leaves of sterile shoots long-petiolate, usually 7-9 cm long
and 2-3 cm wide, twice pinnately cut, terminal lobes 4-8 mm long and
1.0-2.5 mm wide, lanceolate-linear, short-acuminate; middle cauline
leaves sessile or short-petiolate, pinnately cut; upper leaves bracteal,
linear-lanceolate, undivided or with 2—4 teeth at base. Capitula in long
racemose inflorescence, hemispherical, 5—7 mm in dia; lower capitula
on long peduncles, remote, upper on short peduncles, approximate.
Involucral bracts oval, smooth, punctate-glandular, light brown, with
wide scarious border, dark brown. Receptacle convex, glabrous.
Peripheral pistillate florets 12, their corollas narrowly tubular, punc-
tate-glandular, stigma lobes narrowly linear, subobtuse, somewhat di-
vergent after anthesis; disk florets bisexual, numerous (to 75), their

48

—

456

corolla conical, narrowly cup-shaped in upper part, with recurved teeth,
yellow, punctate-glandular, anthers on long filaments, lanceolate, acute,
stigma lobes linear, straight, after anthesis divergent, apically densely
ciliate. Achenes small, oblong-ovoid, angular-sulcate, brown, apically with
scarcely attenuate upper margin; pappus lacking. Flowering August.
Sand dunes, banks, rubbly slopes, in dry-valley larch and open
spruce forests.—Far East: Sakhalin Island. Endemic. Type in Leningrad.

42. A. norvegica Fries. Novit. Fl. Suec. ed. I (1817) 56, tab. 3;
Bess. in Nouv. Mém. Soc. Nat. Mosc. III, 76; DC. Prodr. VI, 119;
Krascheninn. in Sov. Bot. No. 5, 11; Kryl. Fl. Zap. Sib. XI, 2819;
Hult. in Nytt. Mag. Bot. 3, 65.—A. norvegica var. walensis Rupr. in
Beitr. z. Pflanzenk. Russ. Reich. 8 (1850) 64 and FI. bor.-ural. (1856)
37; Hult. op. cit. p. 64.—A. norvegica var. villosula Trautv. ex Korsh.
in Mém. Acad. Sc. Pétersb. VIII, VII, 1 (1898) 221; Hult. op. cit. p.
65.—A. rupestris O.F. Muell. in Fl. Danica fasc. decimus. quart. tab.
DCCCI, ed. 4, (1780) non L.—A. ruprechtii Poljak. in Bot. Mat. Gerb.
Bot. Inst. Akad. Nauk SSSR, XVII (1955) 409.—Ic.: Rchb. PI. crit. I
(1823) tab. 89.

Perennial. Rhizome thick, creeping or ascending, often branched
in upper part, bearing rosettes of leaves and one or many, erect or
ascending (only at base) stems, 13-30 cm high, more or less densely
covered with long entangled hairs, sometimes subglabrous. Leaves hairy;
basal leaves with long flattened petioles, almost 2 times as long as
lamina or almost equal to it, twice pinnately cut, lobes 2-3 pairs,
divided into 3 or 4 linear or linear-lanceolate secondary lobes, 5—15
mm long and 1-2 mm wide; cauline leaves smaller, short-petiolate, at
base usually with small lobes; uppermost leaves sessile, pinnate, with
undivided linear lobes. Capitula 3-7, drooping, 10-12 mm in dia,
hemispherical, one each on 3-12 cm long peduncles, adpressed to
stem, in simple racemose inflorescence. Involucre cup-shaped, involu-
cral bracts oval or oblong-ovate, green and hairy in middle, with brown-
ish-black scarious border. Receptacle glabrous. Peripheral florets pis-
tillate, fertile, 8-12, their corollas narrowly tubular, with 3 teeth, densely
hairy, stigma lobes narrowly linear, glabrous, after anthesis more or
less divergent; disk florets bisexual, numerous (120-156), fertile, in
part immature, their corollas narrowly conical, to 4 mm long, with 5
teeth, yellow, densely hairy or glabrous, punctate-glandular (var.
uralensis Rupr.), anthers on short filaments, linear, acute, connate into
a tube [syngenesious], stigma lobes narrowly linear, after anthesis
exserted from corolla tube, arcuate, apically ciliate. Achenes elon-
gated, prismatic, brown; pappus lacking.

482

457

Stony slopes and outwash gravel in alpine and arctic-alpine zone.—
Arctic: Arctic Europe (northern Urals), Arctic Siberia (Kara River
basin, Kara tundra). General distribution: Scandinavia (Norway).
Described from Kongevold in Norway. Type in Sweden.

Series 7. Globulares Krasch. ex Poljak. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XVII (1955) 426.—Corolla of all florets alike
with 5 teeth, violet; capitula in compact capitate inflorescence.

43. A. globularia Bess. in Nouv. Mém. Soc. Nat. Mosc. III (1834)
64; DC. Prodr. VI, 116; Ldb. Fl. Ross. II, 588; Maxim. in Bull. Acad.
Sc. Pétersb. VIII, 534; Rydb. North Am. Fl. 4, part 3, 261.—A.
norvegica globularia Hall and Clements. Amer. Artem. (1923) 58.—
Aiania globularia (Bess.) Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVII (1955) 426.

Perennial. Root woody, vertical with very short, crowded, perennial,
sterile shoots bearing leaf rosette, forming small tussocks. Stems single
or 2-3, erect or somewhat inclined, 7-13(17) cm high, simple, sulcate,
with 1-2-3 leaves excluding basal, woolly in upper part, glabrous
below or weakly hairy. Leaves grayish from dense, long, straight,
appressed hairs; basal leaves and leaves of sterile shoots long-petiolate,
2-4 cm long and 0.6—1.5 cm wide, usually ternate, lobes undivided or
often divided, with 2-3 lanceolate-linear, 4-10 mm long and 0.5—-1.5
mm wide, short-acuminate lobules; cauline leaves considerably shorter,
undivided, linear-lanceolate or apically short-ternate. Capitula at apices
of stems, subsessile, in compact capitate inflorescence; less often lower
capitula somewhat remote on up to 2.0-2.5 cm long peduncles.
Involucral bracts oblong or oval, more or less densely hairy, with
wide, scarious, brown, unevenly incised margin. All florets with wide,
cup-shaped-conical, dark purple, punctate-glandular corolla, with rather
large, deltoid recurved teeth; peripheral florets pistillate, 9; disk florets
bisexual, to 20; anthers broadly lanceolate, apical appendages of anthers
broadly obtuse, basal appendages smaller, subacute, stigma lobes of
all florets identical, short, linear, apically ciliate, recurved after anthesis.
Achenes oblong, slightly narrowed toward base, tetraquetrous, with
weakly bordered flattened apex; pappus lacking. Flowering August.

Stony seacoasts, rocks, islands, in arctic and subarctic zones.—
Far East: Chukotka, Anadyr, Kamchatka. General distribution: North
America (Alaska). Described from Eschscholz’s collections from Gulf
of Saint Lawrence. Type in Leningrad.

Series 8. Glomeratae Rydb. North Am. FI. 34, part 3 (1916) 260.—
Florets glabrous or weakly hairy; inflorescence usually compact or lax,
corymbose, capitate or racemose.

483

458

44. A. glomerata Ldb. in Mém. Acad. St. Pétersb. V (1812) 564;
Bess. in Nouv. Mém. Soc. Nat. Mosc. III, 63; a. Prodr. VI, 116; Ldb.
Fl. Ross. II, 588; Maxim. in Bull. Acad. Sc. Petersb. VIII (1872) 532
p. p.; Rydb. North Am. FI. 34, part 3, 260, exl. syn.; Hult. Fl. Kamtsch.
IV, 182; Kom. Fl. Kamch. III, 147 p. p.—A. glomerata ssp. curiliensis
(Spreng.) Krasch. in Istor. Fl. i Rast. III (1958) 105.—A. curiliensis
Spreng. Syst. veg. III (1826) 489.—A. corymbosa Fisch. and A.
steveniana ex Bess. op. cit. p. 74.—A. norvegica glomerata Hall and
Clements, North Amer. Artem. (1923) 61.—Ajania glomerata (Ldb.)
P. Pol. comb. nov. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XVII (1955) 426.

Perennial. Tap root woody, with numerous, perennial, woody,
sterile, prostrate or ascending, short shoots in upper part with leaf
rosette, thus forming rather compact tussocks. Fertile stems numerous
or few, (6)10—15(25) cm high, straight or flexuous, in lower part arcuate,
ribbed, simple, weakly leafy, more or less densely hairy above. Leaves
whitish, appressed-hairy; basal leaves and leaves of sterile shoots
petiolate, 1.5—2.0 cm long and 0.7—1.5 cm wide, thrice pinnately parted,
lobes pinnately parted, terminal lobe lanceolate, scarcely acute, 2-6
mm long; cauline leaves few, sometimes 2-3, with 3-6 lobes or
undivided, narrowly lanceolate. Capitula broadly campanulate, to 4
mm wide, at stem apex in more or less compact corymbose-capitate
inflorescence; sometimes lower capitula somewhat remote on long
peduncles (ssp. kuriliensis (Spreng.) Krasch.). Involucral bracts oblong-
elliptical, densely hairy, with involute scarious margin. Receptacle

narrow, glabrous. All florets fertile; peripheral pistillate (to 7), their

corolla tubular, with 3—4 teeth, glabrous; disk florets bisexual, usually
15, their corolla cup-shaped, conical, with 5 teeth, anthers on rather
long filaments, lanceolate-linear, apical appendages of anthers acute,
basal appendages shorter, round; stigma lobes of all florets narrowly
linear, short-ciliate, recurved after anthesis. Achenes to 2 mm long,
narrowly oblong, indistinctly tetraquetrous, with flat scarcely bordered
upper margin.

Seacoasts, in tundra, arctic zone and mountains up to the alpine
zone.—Arctic: Arctic Siberia; Far East: Chukotka, Anadyr, Kamchatka,
Okhotsk. General distribution: North America (Alaska). Type in
Leningrad.

45. A. furcata MB. FI. taur.-cauc. III (1819) 567; Krascheninn. in
Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, IX, 179.—A. trifurcata
Steph. in Spreng. Syst. veg. III (1826) 488; Bess. in Nouv. Mém. Nat.
Mosc. III, 73; DC. Prodr. VI, 119; Ldb. Fl. Ross. II, 591; Maxim. in
Bull. Acad. Sc. Pétersb. VIII (1872) 533; Kom. Fl. Kamch. III, 149;

484

459

Hult. Fl. Kamtsch. IV, 187.—A. heterophylla Bess. op. cit. 74; Ldb.
op. cit. 591; Rydb. North Am. FI. 34, part 3, 262, excl. syn.; Sugawara,
Ill. Fl. of. Sagh. IV, 1831.—Ajania furcata (MB.) P. Poljak. in Bot.
Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVII (1955) 428.—Ic.: Sugaw.
op. cit. tab. 839.

Perennial. Root woody, vertical, with number of very short, woody,
crowded, sterile shoots in upper part with leaf rosettes, forming a
small tussocks. Fertile stem single or two or three, erect or slightly
bent, simple, weakly leafy, dull violet, 1S—25 cm high, densely lanate
in upper part, glabrous below. Leaves weakly hairy, sometimes
subglabrous, with erect, rather long, appressed or somewhat divergent
hairs; basal leaves and leaves of sterile shoots long-petiolate, 3-4 cm
long and to 1.5 cm wide, ternate or palmately divided into 7-12 mm
long and to 1 mm wide, short-acuminate lobes; cauline leaves undivided,
linear, less often ternate. Capitula subglobose, 4—6 mm in dia, in simple
raceme, crowded or almost capitately clustered, upper capitula
subsessile, lower on 5—8 mm long peduncles. Involucral bracts broadly
ovate, densely hairy, with brownish scarious margin. All florets with
colored corolla; peripheral pistillate florets 7, tubular, with 3-4 teeth,
punctate-glandular, glabrous; disk florets bisexual, 15-26, their corollas
cup-shaped-conical, with 5 teeth; anthers lanceolate, apical appendages
of anthers obtuse, basal appendages short, round; stigma lobes of all
florets identical, linear, apically short-ciliate, arcuately curved after
flowering. Achenes oblong, ribbed, with flat-bordered apex; pappus
lacking. Flowering August.

Seacoasts, in river valleys on well-drained terraces, in arctic zone
and alpine tundra belt above timberline—Arctic: Arctic Siberia
(Taimyr), Chukotka, Anadyr; Eastern Siberia: Dauria (Khamer-Daban),
Lena-Kolyma (low-lying areas of Lena and Oleneka rivers); Far East:
Zeya-Bureya (mountains), Okhotsk, Kamchatka. General distribution:
North America (Alaska).

46. A. insulana Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, IX (1946) 177.—A. trifurcata auct. non Steph.: Fedtsch. Fl. des
iles du Commadeur (1906).—A. norvegica ssp. heterophylla Hall and
Clements. North Amer. Spec. of Artemisia (1923), quoad plant. ins.
Command.—Ajania insulana (Krasch.) Poljak. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XVII (1955) 428.

- Root thick, woody, with short perennial, woody, thick, sterile shoots
in upper part, bearing leaf rosette, rather densely crowded forming
small tussock. Fertile stems few, sometimes single, divergent, straight,
sometimes slightly bent, 10-20 cm high, ribbed, arachnoid-hairy, weakly
leafy. Leaves very densely sericeous; basal leaves and leaves of sterile

485

460

shoots petiolate, 2-5 cm long and 1.5-3.0 cm wide, twice ternately
divided, lobes ternate, approximate, terminal lobe 5-15 mm long and
1.5-2.0 mm wide, lanceolate, short-acuminate; cauline leaves ternate
or pinnately parted, or even simple, linear-lanceolate. Capitula in simple,
5-10 cm long raceme, lower capitula on more or less long peduncles,
remote, drooping, upper capitately clustered, hemispherical, 5-8 mm
in dia. Involucral bracts oval, densely hairy, with blackish scarious
margin. Receptacle glabrous. All florets fertile, purple-violet; peripheral
florets pistillate, 8, tubular, with 3—4—5 teeth; disk florets bisexual, 12,
their corollas conical, with 5 teeth, hairy, anthers on short filaments,
lanceolate, apical appendage of anthers obtuse, basal appendages short,
sub-acute; stigma lobes of all florets identical, linear, apically ciliate.
Flowering August.

Stony slopes of [mud] volcanic cones and outwash gravel.—Far
East: Commander Islands. Endemic. Described from Mednyi Island.
Type in Leningrad.

Series 9. Senjavinenses Poljak.—Leaves ternate; capitula in
compact capitate or oblong[cylindrical] inflorescence.

47. A. senjavinensis Bess. in Nouv. Mém. Soc. Nat. Mosc. III
(1832) 65, err. “semavinensis;” DC. Prodr. VI, 116; Ldb. Fl. Ross. II,
588; A. Gray Synopt. Fl. North Am. Fl. 1, part 2, 370; Rydb. North
Am. FI. 34, part 3, 260.—Ajania senjavinensis (Bess.) Poljak. in Bot.
Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVII (1955) 428.

Perennial. Whole plant whitish from dense, long divergent hairs.
Tap root woody, with very short, sterile, woody shoots in upper part,
densely covered with small overlapping leaves, hence assuming habit
of hemispherical or oblong “heads,” 0.3-1.5 cm long and 0.5-1.3 cm
wide, “heads” in turn overlapping to form small compact tussock. Stems
few, 3-6 cm high, straight, simple, leafy. Leaves of sterile shoots 4—
7 mm long, in upper half ternate, lobes lanceolate, acute; cauline leaves
usually 2—5, with 3-5 lanceolate lobes. Capitula on short peduncles,
campanulate, 4 mm long and 3.5 mm wide, in compact, terminal,
capitate, globose or oblong inflorescence, 1-2 cm long and 1.0—1.5 cm
wide. Involucral bracts semitransparent, membranous, on margin
unevenly incised and with acute teeth; outer bracts broadly elliptical,
densely hairy; inner bracts oblanceolate, glabrous, with occasional hairs
only on margin. Receptacle convex, conical, glabrous. All florets fertile;
peripheral florets pistillate (5); their corollas tubular with 3 teeth,
glabrous, yellow; disk florets bisexual, rather numerous, their corollas
cup-shaped-conical, with 5 teeth, anthers on rather long filaments,
lanceolate, apical appendages of anthers obtuse, basal appendage

486

461

shorter, acute; stigma lobes of pistillate and bisexual florets identical,
narrowly linear, apically short-ciliate, usually arcuately bent after
anthesis. Achenes about 2 mm long, linear-oblong, with 4 indistinct
angles, brown, with flat apex, weakly bordered. Flowering August.

Arctic seacoasts.—Far East: Chukotka. General distribution: North
America (western Alaska). Described from Bering Sea from Mertens’
collections. Type in Leningrad.

48. A. leontopodioides Fisch. ex Bess. in Nouv. Mém. Soc. Nat.
Mosc. III (1832) 63; DC. Prodr. VI, 116; Ldb. Fl. Ross. II, 587.—A.
glomerata Rydb. North Am. Fl. 34, part 3 (1916) 260 p. p.—Ajania
leontopodioides (Bess.) in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XVII (1955) 428.

Perennial. Whole plant silky from long, straight, appressed, whitish
hairs. Root woody, with numerous short, ascending, woody leafy shoots
in upper part forming more or less a compact tussocks. Stems rather
numerous or few, (6)10—15(20) cm high, arcuate or flexuous, simple,
weakly leafy. Basal leaves and leaves of sterile shoots petiolate, to
2.0-—2.5 cm long, ternate, lobes usually ternate, their terminal secondary
lobes linear-lanceolate, acute, 2-5 mm long; cauline leaves smaller,
mostly simple, ternate, uppermost sometimes simple, narrowly
lanceolate. Capitula numerous, on short peduncles, campanulate or
hemispherical, 3—4 mm in dia, in compact globose, corymbose-capitate
inflorescence. Involucral bracts membranous, densely hairy; outer bracts
oval, inner oblong-lanceolate. Receptacle small, glabrous, peripheral
florets pistillate, their corollas tubular, with 3-4 teeth; disk florets
bisexual, their corollas broadly cup-shaped conical, deeply notched-
toothed, densely covered with long, straight white, divergent hairs.
Achenes oblong-linear, weakly ribbed, with long, white divergent hairs.

Stony seacoasts—Far East: Kamchatka, Sakhalin (Kuril Islands).
General distribution: Reported for North America (Alaska). Type in
Leningrad.

Series 10. Angaro-ochotenses Krasch. ex Poljak.—Perennials,
densely caespitose; leaves twice pinnately cut or twice ternate.

49. A. lagopus Fisch. ex Bess. in Nouv. Mém. Soc. Nat. Mosc. III
(1832) 69; DC. Prodr. VI, 117; Ldb. Fl. Ross. Il, 589.—Maxim. in
Bull. Acad. Sc. Pétersb. VIII, 34.—A. meyeriana Bess. in Linnaea,
XV (1841) 96.

Perennial. Whole plant silky-woolly from whitish (ochraceous when
dry), long, straight or flexuous, appressed and divergent hairs. Root
woody, with perennial woody, procumbent or ascending sterile shoots,

487

462

with young leafy shoots forming compact tussocks. Fertile stems few,
sometimes single, (7)12—20(25) cm high, straight, at base arcuately
curved, cylindrical or weakly ribbed, simple, leafy. Leaves of sterile
shoots 2—3 cm long, long-petiolate, petioles gradually thickening toward
base, lamina round, 0.7—1.2 cm wide, twice pinnately cut, with strongly
approximate lobes, terminal lobes lanceolate-linear, acute, 1.5-3.0 mm
long; lower and middle cauline leaves short-petiolate, upper leaves
sessile, simple pinnate. Capitula sub-globose, 4-5 mm long and 3-4
mm in dia, subsessile or on short peduncles, clustered in terminal,
compact, linear-oblong, 3-5 cm long, spicate-racemose inflorescence,
but sometimes lower capitula somewhat remote, and then inflorescence
elongated (ssp. abbreviata Krasch.), up to 8-12 cm long. Receptacle
convex, scatteredly hairy. Involucral bracts lanceolate, acute; outer
bracts woolly, inner ones shorter than outer bracts, weakly hairy, with
wide scarious border. Peripheral florets pistillate, with narrowly tubular,
punctate-glandular, glabrous corolla. Disk florets bisexual, their corollas
conical, with hairs on teeth. Achenes narrowly oblong, somewhat
narrowed toward base, ribbed. Flowering August.

Sandy seacoasts and rocks.—Far East: Okhotsk. Endemic.
Described from Okhotsk coast from Krause’s collections. Type in
Leningrad.

50. A. triniana Bess. in Nouv. Mém. Soc. Nat. Mosc. III (1832)
67; DC. Prodr. VI, 117; Ldb. Fl. Ross. II, 589; Krascheninn. in Sov.
Bot. No. 5 (1943) 14.

Perennial. Whole plant whitish-tomentose from long, usually
appressed hairs. Root woody, vertical, with rather numerous, branched,
perennial, prostrate shoots densely caespitose, with short, young, densely
leafy shoots. Fertile stems rather numerous, sometimes few, including
inflorescence 3—5—10 cm high, usually densely leafy, simple. Leaves
of sterile shoots and lower cauline leaves petiolate, 1.0—1.4(1.8) cm
long, petioles thickening toward base, lamina almost round, 0.6—1.0
cm wide, twice ternate or twice ternate-palmately divided, lobes often
3, short-petiolate, ternately palmate or pinnate, secondary lobes 1-3
mm long, lanceolate or linear-lanceolate, subacute; middle and upper
cauline leaves sub-sessile or sessile, mostly pinnately cut. Capitula
hemispherical, 4-5 mm long and 3.5-4.5 mm in dia, on very short
peduncles, clustered in compact, terminal, ovate or oblong-ovate, 1—2
cm long inflorescence. Outer involucral bracts oblong-lanceolate, with
narrowly herbaceous midrib and scarious margin, brownish, usually
densely covered with long, white, semi-squarrose hairs; inner bracts
oblong, glabrous, hairy only along margin. Receptacle convex,

488

463

glabrous. Peripheral florets pistillate, 7, their corollas narrowly cup-
shaped-conical, with 4 teeth, punctate-glandular, glabrous; disk florets
bisexual, 11-12, cup-shaped conical; anthers on long filaments, lan-
ceolate-linear, apical appendages of anthers acute-angled, basal ap-
pendages shorter, acute; stigma lobes of all florets linear, apically
short-ciliate, straight, after anthesis recurved.

Sandy coasts, well-drained high terraces of rivers in arctic zone.—
Arctic: Arctic Siberia (low-lying areas of the Lena, Olenyek and
Khatanga rivers. Endemic. Type in Leningrad.

Series 11. Paniculigerae Poljak.—Artemisia subsect. Glochistigma
Kitamura in Acta Phyt. et Geobot. VIII (1939) 63.—Capitula in
racemose-paniculate inflorescence; stigma lobes with narrowly linear
apical appendage.

51. A. keiskeana Miquel, in Ann. Mus. Lugd.-Bat. II (1866) 176;
Maxim. in Bull. Acad. Sc. Pétersb. VIII, 534; Kom. Fl. Manchzh. III,
669; Nakai Fl. Korean. II, 29; Nakai in Bot. Mag. Tokyo, XXVI, 101,
incl. f. typica, hirtella and hypoleuca: Pampan. in Nouv. Giorn. Bot.
Ital., n.s. XXXIV, 670; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II,
1042; Hara, Enum. Spermatoph. Japonic. II, 119.—A. vulgaris
stolonifera lusus b. glabrescens Rgl. Fl. Ussur. (1862) 95.—A. japonica
F. Schmidt. Fl. Sachal. (1868) 149, planta Albrechti e Hokadate, non
Thbg.

Perennial. Rhizome strong, branched, with underground or aerial
creeping shoots. Stems erect or bent, 20-50 cm high, uniformly leafy,
with divergent hairs, in part subglabrous, with short branches in upper
part or almost simple. Leaves sometimes with small acute auricles,
green above, usually glabrous, pale green beneath, weakly hairy; lower
cauline leaves and leaves of sterile shoots 5—6(8) cm long, usually
short-petiolate, cuneate or ovate, broadly toothed with 7 teeth, cuspidate;
middle cauline leaves always sessile, smaller, with 3-5 teeth; uppermost
leaves broadly lanceolate, entire or with a pair of very short, acute
teeth. Capitula on long peduncles, bent or drooping, subglobose, 2.5—
3.0 mm long, in short, secund, lax racemes forming narrowly racemose-
paniculate inflorescence. Involucral bracts glabrous, convex, green along
midrib, with wide scarious margin; outer bracts oval, inner larger, with
attenuate obtuse apex. Peripheral florets pistillate, 7, their corollas
narrowly tubular, punctate-glandular, pilose; stigma lobes short, broadly
linear, scarcely reduced; disk florets 14, bisexual, their corollas conical,
punctate-glandular, anthers linear, apical appendages of anthers short,
obtuse, basal appendages smaller, subobtuse; stigma lobes apically
broadened, bearing quite long, narrowly linear appendage. Achenes

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464

plano-convex, ovate, to 1.5 mm long, dark brown, glabrous, lacking
pappus. Flowering August.

Forest zone, on slopes of [mud] volcanic cones and ridges, deciduous
and deciduous-open coniferous forests, partly on rock outcrops.—
Far East: Zeya-Bureya, Ussuri (southern Sikhote-Alin). General
distribution: Northern China, Korea, Japan. Described from Japan.

Series 12. Annuae Rydb. in North Amer. FI. 34, part 3 (1916)
259.—Plants glabrous or scatteredly short hairy; leaves twice or thrice
pinnately divided into oblong-lanceolate, toothed lobes with subacute
teeth, less often lobes entire; receptacle glabrous.

52. A. tournefortiana Rchb. Icon. ex. Cent. 1 (1823) 6, tab. 5;
Bess. in Nouv. Mém. Soc. Nat. Mosc. III, 82; DC. Prodr. VI, 119;
Ldb. Fl. Ross. Ii, 592; Boiss. Fl. or. III, 372; Hook. Fl. Brit. Ind. III,
324; Fedtsch. Perech. Rast. Turk. IV, 200; Pampan. in Nouv. Giorn.
Bot. Ital. n. s). XXXIV (1927) 705; Grossh. Fl. Kavk. IV, 138.—
Pampan. in Lavori Inst. bot. Univ. Cagl. XL, 570.—A. annua Trautv.
Enum. pl. Song. (1868) No. 616; non L.

Annual. Plants glabrous or with scattered, bifid, forked, appressed
hairs. Stem 120-190 cm high and up to 7-14 mm thick, brownish-
violet, finely ribbed, almost simple, leafy. Leaves (except bracteal
leaves) petiolate; lower leaves 7-16 cm long and 4—6 cm wide, lamina
ovate-oblong, twice pinnately cut, their lobes short-petiolate, oblong-
lanceolate, deeply pinnately divided or almost divided into lanceolate,
subacute, toothed lobules; middle and upper cauline leaves shorter,
less incised, bracteal leaves simple pinnate, serrate dentate or even
sometimes entire, linear-lanceolate. Capitula numerous, broadly ovate
or subglobose, to 2 mm wide, upright, densely clustered on short or
long, weakly divergent shoots forming usually narrow, rather compact
paniculate inflorescence. Involucral bracts with herbaceous midrib,
glabrous, broadly scarious; outer bracts oval, inner bracts lanceolate
or elliptical. Receptacle convex, glabrous, tuberculate-glandular, stigma
lobes exserted, narrowly linear, straight or slightly recurved. Disk florets
bisexual 10, their corollas conical, purple-pink, in lower part punctate-
glandular; anthers narrowly linear, apical appendages of anthers
elongated, cuneate, basal appendages short, acute, their stigma lobes
short, not exserted from tube, linear. Achenes to 1 mm long, oblong-
narrowly ovoid, somewhat flattened, finely sulcate, apically with flat,
round areole. Flowering August.

Weed, on riverbanks, irrigation canals, irrigated lands in desert
and steppe zones, in mountains, along valleys and slopes.—Caucasus:
Southern Transcaucasia; Soviet Central Asia: Lake Balkhash Region,

490

465

Dzhungaria-Tarbagatai, Tien Shan, Syr-Darya, Pamiro-Alai, Amu-
Darya, Kara-Kum, (Amu-Darya delta), mountainous Turkmenia.
General distribution: Armenia and Kurdistan, Iran, Indo-Himalayas,
Dzhungaria-Kashgaria, Tibet. Described from “Orient.” Type in Berlin.

Economic Importance. S.N. Kudryashev (1934) reports that the
aerial part of the plant contains about 0.01% essential oil; according
to the data of Vyshensky (1938)—0.14—0.34%. According to the data
of M.I. Goryaev et al. (1954), from plants collected 15 km from the
village of Lugovaya (Dzhambul Region), the yield of essential oil was
0.053% in absolutely dry matter, but 0.035% on an air-dry basis. The
oil is a viscous liquid, yellow-orange in color, with a pleasant smell.
n> = 1.5080.

.53. A. annua L. Sp. pl. (1753) 847; Bess. in Nouv. Mém. Soc.
Nat. Mosc. III, 81; DC. Prodr. VI, 119; Ldb. Fl. Ross. II, 592; Boiss.
Fl. or. HI, 371; Maxim. in Bull. Acad. Sc. Pétersb. VIII, 529; Hook.
Fl. Br. Ind. IJ, 323; Kom. in Fl. Manchzh. III, 659; Nakai, Fl. Korean.
II, 30; Fedtsch. Perech. Rast. Turk. IV, 200; Rydb. North Am. FI. 34,
part 3, 259; Pampan. in Nouv. Giorn. Bot. Ital. n. s. XXXIV, 637; Hall
and Clem. Artem. (1923) 102; Kom. and Alis. Opred. Rast. Dalnevost.
Kr. II, 1036; Krascheninn. in Fl. Yugo-Vost. Evrop. Ch. SSSR, VI,
357; Grossh. Fl. Kavk. IV, 138; Krascheninn. in Kryl. Fl. Zap. Sib.
XI, 2816; Poljak. in Majevski. Fl. 586.—A. chamomilla Winkl. in Tr.
Peterb. Bot. Sada, X, 87; Fedtsch. loc. cit.—Ic.: Amm. Stirp. rar. t,
193, f. 23; Gmel. FI. sib. II, tab. 125.— Exs.: GRF No. 3152.

Annual. Plant aromatic, green, glabrous or with scattered, small,
approximate hairs. Stems erect, ribbed, brownish or violet-brown, 30—
100 cm high. Leaves alveolate-punctate-glandular; lower leaves
petiolate, 3-5 cm long and 2-4 cm wide, ovate, thrice pinnatley cut,
their lobules oblong-lanceolate, short-acuminate, entire or with 1-2
teeth, 1-2 mm long and 0.5 mm wide; middle and cauline leaves twice
pinnately cut; upper leaves sessile smaller and less compound;
uppermost leaves bracteal, simple or with fewer lateral lobes. Capitula
globose, 2.0-2.5 mm in dia, numerous, divergent or drooping, on short
peduncles, approximate on short branches, usually in long pyramidal
paniculate inflorescence. Involucre glabrous. Outer involucral bracts
linear-oblong, green; inner oval or almost round, with wide scarious
border, lustrous. Receptacle convex, glabrous. Peripheral florets
pistillate, 10-20, filiform, punctate-glandular; their stigma lobes
narrowly linear, obtuse, exserted from corolla tube; disk florets bisexual,
12-30, their corollas cup-shaped-tubular, glabrous; anthers narrowly
linear, apical appendages of anthers long, acute, basal appendages very
short, subacute; style shorter than stamens, stigma lobes linear, straight,

491

466

weakly divergent, apically ciliate. Achenes 0.8—0.6 mm long, oblong-
ovate, flattened, with small round areola at apex, scarcely bordered.
Flowering August to September.

Weedy places near dwellings, kitchen-gardens, gardens.—Euro-
pean part: Upper Dniester, Upper Dnieper, Volga-Don, Trans- Volga,
Lower Don, Black Sea Region, Bessarabia, Crimea; Caucasus: Eastern
and Western Transcaucasia, Talysh; Soviet Central Asia: Aralo-Caspian
Region, Lake Balkhash Region, Dzhungaria-Tarbagatai, Tien Shan, Syr-
Darya, Pamiro-Alai, Amu-Darya, mountainous Turkmenia. General
distribution: Central Europe, Mediterranean, Balkans-Asia Minor,
Armenia and Kurdistan, Iran, Dzhungaria-Kashgaria, China, Japan,
Mongolia, North America (introduced). Amman’s and Gmelin’s figures
are the type for this species.

Economic Importance. The yield of essential oil is 0.1-0.64%
(according to Goryaev). According to the data of Joshikazu Imade
(1937), this oil contains cineole and a substance with the composition
C,,H,,O; this substance was analyzed in detail by Asahina and Ioshitomi
(1917) and along with artemisia ketone, its isomer, isoartemisia ketone,
was also found in the oil. This ketone could be isolated from the
mother liquor left behind after obtaining the semicarbozone of artemisia
ketone. Seisi Takagi (1928), extending his investigations of the Japanese
A. annua, added two more new components, viz. cadinene and
caryophyllene, to the four components already known in the oil. Based
on the investigations of Rutovsky and Vinogradov (1929), the oil
contains Q-pinene, cineole, camphene, artemisia ketone and isoartemisia
ketone, a small quantity of borneol, acetic and butyric acids,
cumaldehyde (presumably semicarbozone) and phenol (probably
eugenol). A small quantity of alkaloids was found in the underground
parts (Lazurevsky and Sadykov, 1939; Massagetov, 1947). According
to field observations (Yunatov, 1954), cattle do not graze on it when
green.

The presence of alkaloids is confirmed by the data of M.I. Goryaev,
G.K. Kruglykhina, and E.I. Satdarova (1959).

Series 13. Auratae Poljak.—Plants glabrous; leaves pinnatisect or
twice pinnately cut with linear-lanceolate or more often filiform-linear
lobes.

54. A. palustris L. Sp. pl. (1753) 846; Bess. in Nouv. Mém. Soc.
Nat. Mosc. III, 184; DC. Prodr. VI, 120; Ldb. FI. Ross. II, 593; Maxim.
in Bull. Acad. Sc. Pétersb. VIII, 529; Komar. Fl. Manchzh. III, 661;
Pampan. in Nouv. Giorn. Bot. Ital. n. s. XXXIV, 683; Kom. and Alis.
Opred. Rast. Dalnevost. Kr. II, 1036; Krascheninn. in Kryl. Fl. Zap.

492

467

Sib. XI, 2817; Grub. Konsp. Fl. Mong. 267.—Ic.: Gmel. Fl. sib. II,
tab. 55.—Exs.: GRF No. 3153a, b, c.

Annual. Plant glabrous, with slender vertical root. Stem (4)10—50
cm high, erect, finely ribbed, green or violet-brownish, branched
sometimes from base itself with rather long, upward spreading branches,
less often simple. Leaves green, except lowermost sessile, with auricles,
mostly 2-5 cm long, 1.5-3.0 cm wide, simple, pinnate or almost twice
pinnately cut into narrow filiform-linear, somewhat thick lobes, 2-3
cm long and 0.2-0.5 mm wide, with scarcely visible punctate glands;
bracteal leaves smaller, more often simple pinnate, divergent. Capitula
subglobose, 2-3 mm in dia, 2-10, clustered in compact, lax or more
or less approximate heads or short racemes forming paniculate
inflorescence. Involucral bracts broadly oval, spatulately elongated,
lustrous; outer bracts with green herbaceous midrib and wide, scarious,
transparent or brownish margin; inner membranous. Receptacle convex-
conical, glabrous. Peripheral florets pistillate, 11-13, filiform-tubular;
their stigma lobes narrowly linear, obtuse, divergent, exserted from
corolla tube; disk florets 28-30, their corollas conical, glabrous,
punctate-glandular; anthers linear, apical appendages of anthers acute,
basal appendages short, subobtuse; stigma lobes short, linear, apically
ciliate, after anthesis divergent. Achenes 0.5-0.7 mm long, oblong-
ovate, flattened, brown, with flat round areola at apex. Flowering
August.

Steppe meadows, sandy banks of rivers and lakes, gravel beds,
abandoned fields, pastures and roadsides.—Western Siberia: Altai
(Chuya steppe); Eastern Siberia: Angara-Sayans, Dauria; Far East:
Zeya-Bureya (western part) General distribution: Mongolia, northern
China. Described from Siberia. Type in London.

55. A. aurata Kom. in Tr. Peterb. Bot. Sada, XVIII (1901) 422;
Kom. Fl. Manchzh. III, 661; Nakai Fl. Korean. II, 30.—Kom. and
Alis. Opred. Rast. Dalnevost. Kr. II, 1036.

Annual. Plant glabrous, with slender roots. Stem 15—60(80) cm
high, erect, strongly branched, with long, three-angled, acute-scabrous,
arcuate branches. Lower cauline leaves long-petiolate, 6-12 cm long,
twice pinnately cut, lobes narrowly linear-lanceolate, acute, simple or
with linear teeth; middle cauline leaves smaller, short-petiolate, with
narrowly linear or almost filiform lobes; upper leaves sessile, simple,
pinnate; uppermost leaves bracteal, undivided, narrowly linear-
lanceolate or with 1-2 pairs of linear, acute lobes. Capitula sessile,
subglobose, 2-3 mm long, in heads of 2—5 forming a lax panicle.
Involucral bracts broadly oval or round, with or without narrow green
keel, membranous, lustrous. Receptacle convex, conical, glabrous.

493

468

Peripheral florets pistillate, 13-14, their corollas narrowly tubular,
toward base slightly broadened, glabrous, punctate-glandular; stigma
lobes exserted from tube, linear, obtuse, divergent; disk florets bisexual,
14-25, their corollas to 1.5 mm long, conical, golden-yellow, anthers
linear; stigma lobes linear, straight, short, apically ciliate, after anthesis
somewhat divergent. Achenes very small, very similar to those of the
preceding species.

South-facing stony slopes, in crevices of rocks, in the zone of
coniferous-deciduous forests.—Far East: Ussuri. General distribution:
Japan, north China, north Korea. Described from northeastern China.
Type in Leningrad.

Note. This species is very similar to A. palustris L. from which it
is distinguished by a more complex dissection of the leaves.

Section 4. Absintium DC. in Lam. and DC. FI. Franc. ed. 3 (1815)
189.—Herbaceous perennials or annuals, or even low semishrubs
(shrubs), more or less densely covered with white, appressed or erect,
dichotomous or Malpighian[sic.] hairs, less frequently almost glabrous;
receptacle pubescent, with, very rarely without, floral [sic.] hairs; leaves
usually twice or thrice pinnately cut.

Section 1. Argyrophyllae Poljak.—Leaves twice pinnately cut or
twice ternate; capitula hemispherical, globose or even ovoid, 2-4 mm
in dia; stem erect or ascending.

56. A. caucasica Willd. Sp. pl. III (1804) 1823; Boiss. Fl. or. III,
374; Grossh. Fl. Kavk. IV, 140.—A. alpina Pall. in Wiild. Sp. pl. II,
1824; MB. FI. taur.-cauc. II, 297.—A. lanata DC. Prodr. VI (1837)
123; Ldb. Fl. Ross. II, 596; A. monticola C. Koch in Linnaea, XXIV
(1851) 347; Grossh. Fl. Kavk. IV, 140.—A. rupestris Pall. Ind. Taur.
(1801); MB. Casp. app. 210 (non L.).—A. chevsurica Somm. and Lev.
in Bull. Soc. Bot. Ital. (1898) 130.—A. argentatum Klok. in Bot. Mat.
Gerb. Akad. Nauk SSSR, XVI (1954) 366.—A. grossheimii Krasch. ex
Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVII (1955)
410 p. p.; Grossh. Opred. (1949) 463, nomen.—Absinthium alpinum
Bess. in Bull. Soc. Nat. Mosc. (1829) 240.—Exs.: Herb. Fl. Cauc. No.
496.

Perennial. Whole plant silvery-white from dense, silky, appressed
hairs. Root thick, vertical, with perennial, branched, short, woody
shoots, which together with young, sterile, very short, densely leafy
shoots form rather compact tussocks. Fertile stems few or rather
numerous, (8)12—15(20) cm high, erect, simple, weakly leafy. Leaves
of sterile shoots and lower cauline leaves petiolate, 1.0—1.8 cm long,

494

469

lamina round or broadly round, twice pinnately cut or ternate, terminal
lobes 4—6 mm or 5-10 mm long (var. grossheimii Poljak.), narrowly
lanceolate-linear, short-acuminate; middle and upper cauline leaves
sessile, simple pinnatisect or serrate, sometimes ternate. Capitula in
simple raceme, on short peduncles, subglobose, 4-5 mm long and 3.0—
4.5 mm in dia, solitary or few clustered in rather narrow, interrupted
racemose-panicle or raceme. Involucral bracts densely pilose, with wide
scarious margin; outer involucral bracts elliptical, inner lanceolate.
Receptacle hairy. Peripheral florets pistillate, 5-8; their corollas
narrowly tubular, hairy in upper part, stigma lobes linear, truncate;
disk florets rather numerous (18-20), bisexual, their corollas conical,
apically densely hairy; anthers linear, apical appendages of anthers
narrowly linear, subacute, basal appendages much smaller, subacute,
stigma lobes linear, obtuse, apically ciliate, after anthesis recurved.
Achenes to 1.2 mm long, oblong-ovoid, somewhat flat, dark brown,
smooth, with round convex apex. Flowering August.

Steppe, outlier hillocks and low hills.—European part: Black Sea
Region, Crimea, Lower Don; Caucasus: Ciscaucasia, Dagestan, Eastern,
Western and Southern Transcaucasia. General distribution: Armenia
and Kurdistan. Described from Caucasus. Type in Berlin.

Note. A doubtful species displaying the closest affinity with A.
laxa Lam., found in the Alps. It is distinguished from the latter by
minor characters such as denser pubescence of the corolla and the
involucral bracts. As regards A. argentata Klok., characters such as
little variation in the size and shape of the leaf lobes, their number, as
well as the size of the capitula and the density of pubescence on the
vegetative organs, enables us to separate this plant as an ecological
form.

57. A. splendens Willd. Sp. pl. III (1804) 1822; DC. Prodr. VI,
123; Ldb. Fl. Ross. II, 596; Boiss. Fl. or. III, 375; Grossh. Fl. Kavk.
IV, 139, Opred. Rast. Kavk. 463.—A. peduncularis MB. FI. taur.-
cauc. III (1819) 566.—A. pedunculata Steudél. Nomencl. (1822).— ?
A. mutellina S.G. Gmel. Reise. I (1770) 159.—Absinthium splendens
(Willd.) Bess. in Bull. Soc. Nat. Mosc. (1829) 238.—A. peduncularis
Stev. in Mém. Soc. Nat. Mosc. IV (1835) 63.

Perennial. Tap root woody, with strongly condensed, ascending,
woody, perennial, densely leafy shoots usually forming compact tussock.
Fertile stems few, sometimes single, (10)15—25 cm high, erect or bent,
ascending, brownish, obtusely ribbed, simple, weakly leafy, more or
less pubescent. Leaves on both sides silvery from dense, silky, appressed
hairs; lower cauline leaves long-petiolate, highly variable in size, 2.0—
4.5(6.0) cm long and 1.5—2.5(4.0) cm wide, twice pinnate or twice

497

470

pinnate-ternate, or twice ternate, their terminal lobes linear, acute, 4—
10(12) mm long; cauline leaves shorter, less compound; uppermost
leaves linear. Capitula in simple raceme, low, remote, on long (to 2—
3 cm long) peduncles; upper capitula on short peduncles, more or less
approximate, hemispherical, narrowed toward base, 5-7 mm in dia.
Involucral bracts with wide scarious margin; outer bracts oval, dark
brown along midrib, hairy; inner bracts narrower, weakly hairy.
Receptacle convex, hemispherical, with long hairs. Peripheral florets
pistillate, 9-12, their corollas filiform-tubular, with 2 teeth, stigma
lobes narrowly linear, scarcely acuminate, exserted from tube; disk
florets 33-56, bisexual, their corollas tubular-conical, with erect hairs
in upper part; anthers linear, apical appendages of anthers long, acute-
subulate, basal appendages smaller, subacute; stigma lobes narrowly
linear, truncate, ciliate, recurved after flowering. Achenes narrowly
prismatic, angled, flat, dark brown, with convex areola at apex, scarcely
bordered. Flowering August.

On loamy, rubbly, meadow, high-mountain slopes, river deposits,
old moraines, at upper limit of forest.—Caucasus: Ciscaucasia,
Dagestan, Eastern and Southern Transcaucasia. General distribution:
Armenia and Kurdistan. Described from Armenia. Type in Berlin.

58. A. frigida Willd. Sp. pl. III (1804) pag. 1838; DC. Prodr. VI,
124; Ldb. Fl. Ross. II, 597; Maxim. in Bull. Acad. Sc. Pétersb. VIII,
537; Kryl. Fl. Alt. III, 656; Kom. Fl. Manchzh. III, 680; Fedtsch.
Perech. Rast. Turk. IV, 203; Rydb. North Am. FI. 34, part 3, 258;
Pampan. in Nouv. Giorn. Bot. Ital. n. s. XXXIV, 655; Krascheninn. in
Fl. Yugo-Vost. Evrop. Ch. SSSR, VI, 361; Kryl. Fl. Zap. Sib. XI,
2797.—A. frigida v. typica v. intermedia Trautv. in Bull. Soc. Nat.
Mosc. XXXIX, 2 (1866) 358.—A. frigida ssp. willdenoviana (Bess.)
Krasch. in Kryl. Fl. Zap. Sib. XI (1949) 2798.—A. frigida ssp.
gmeliniana (Bess.) Krasch. Ibid. 2798.—A. frigida ssp. parva Krasch.
Ibid. 2798.—A. frigida v. argyrophylla Trautv. op. cit. 358; Kryl. FI.
Alt. III, 656.—A. jeniseensis Spreng. Syst. veg. III (1826) 486.—A.
argyrophylla Ldb. Fl. Alt. TV (1833) 66; DC. Prodr.VI, 123; Ldb. FI.
Ross. II, 598; Kryl. Fl. Zap. Sib. XI, 2799.—Absinthium frigidum
Bess. and A. frigidum willdenoviana Bess. in Bull. Soc. Nat. Mosc.
(1829) 251, 256.—Ic.: Ldb. Ic. pl. Fl. Ross. V (1834) tab. 463.—Exs.:
GRF Nos. 66, 3172a, b, c.

Perennial. Whole plant densely covered with bifid, slender,
appressed, silky hairs. Root woody, with perennial, woody spreading
branches, which together with young sterile leafy shoots form more or
less dense tussocks. Fertile stems rather numerous or few, 15-30 cm
or 5-10 cm (var. parva (Krasch.) Poljak.) or even 30-50 cm (var.

471

Plate XXIII.
1 — Artemisia abrotanum L., upper part of plant, bisexual disk floret, achene.
2 —A. gmelini Web. ex Stechm., upper part of plant, peripheral pistillate floret,

495

pistil, stamen, bisexual disk floret.

498

472

gmeliniana (Bess.) Poljak.) high, basally ascending or erect, usually
strong, leafy, densely hairy, later glabrous or subglabrous in lower
half. Leaves short-petiolate or sessile (var. argyrophylla (Ldb.) Kryl.)
almost twice pinnate or sometimes ternate, most often 1-2 cm long and
7-15 mm wide; lower pinnae, 2—3-fid or undivided, others crowded at
leaf apex, pinnately divided or ternate, with lanceolate-linear, acute lobes,
2-5 mm long and 0.5—1.0 mm wide; upper bracteal leaves palmately cut
or ternate. Capitula subglobose or globose, 3-4(5) mm wide, divergent or
drooping, in narrowly paniculate or racemose inflorescence. Involucral
bracts with white scarious border; outer bracts oblong-elliptical, subobtuse,
densely hairy, inner bracts linear-oblong, hairy only along margin.
Receptacle convex, hairy. Peripheral florets pistillate, 9-12, their corollas
narrowly or almost filiform-tubular, with 2 teeth, glabrous, punctate-
glandular; stigma lobes filiform-linear, exserted from corolla tube, weakly
divergent; disk florets bisexual, 22-36, their corollas conical, yellow, less
often purple-pink, glabrous, sometimes hairy (f. argyrophylla (Ldb.)
Poljak.). Anthers linear, apical appendage of anthers linear-subulate,
elongate, their basal appendages short, round; stigma lobes scarcely ©
exserted from corolla, linear, apically ciliate, arcuate after flowering.
Achenes 1 mm long, pyriform, somewhat flat, with scarcely visible fine
ribs, brown, apically flat with border on one side. Flowering August.

Steppe zone on rubbly stony slopes of [mud] volcanic cones, low
hills, sometimes on sand dunes and terraces, edge of pine forests or,
less often, in old fields and steppe meadows.—European part: Volga-
Kama, Trans-Volga; Soviet Central Asia: Aralo-Caspian Region
(northern part), Lake Balkhash Region, Dzhungaria-Tarbagatai; Western
Siberia: Upper Tobol, Irtysh, Altai, Angara-Sayans, Dauria, Lena-
Kolyma. General distribution: Mongolia, North America. Described
from Siberia, probably from Gmelin’s collection. Type in Berlin.

Note. As in A. ‘rutifolia Steph., in A. frigida Willd. the corolla is
hairy or glabrous in the upper part, but the bracteal hairs (“receptacle
hairy”) sometimes are almost absent.

Economic Importance. According to Rabak (1912), the plant,
particularly at and after anthesis, contains an essential oil whose yield
varies from 0.26 to 0.41% of the fresh weight of the plant; but when
the plant is dried drops to 0.07%. According to Goryaev (1952), the
oil is yellow in color; its constants are: D??~0.927-0.940; aD—from
23°40' to 25°10'; nD—1.4716; acid number-1.2; ester number-31.8;
ester number after acetylation—139-143. It dissolves in 80% alcohol
in the ratio of 1:2; in 90% alcohol—1:0.5. American oil contains 38-
48% \|-borneol in bound form and to 35% in free form; 18-20% cin-
eole; 8-10% l-fenchone; capryl and evant acids, esters of valerian,
evant, undecyl and formic acids. According to reports of perfumers

499

473

(Rabak, 1912; Goryaev, 1952), the oil is suitable for imparting fra-
grance to soaps. According to Yurensky (1850), the plant is a nutritive
fodder for sheep. A. frigida Willd. is highly valued as fodder in Canada
and the USA during winter. A cold infusion of the herb is used as a
diuretic and mild purgative (Hall and Clements, 1923).

59. A. austriaca Jacq. in Murr. Syst. (1784) 744; Bess. in Nouv.
Mém. Soc. Nat. Mosc. III, 47; DC. Prodr. VI, 112; Ldb. Fl. Ross. hy,
583; Boiss. Fl. or. III, 369; Kryl. Fl. Alt. III, 649; Fedtsch. Perech.
Rast. Turk. IV, 199; Hegi, Ill. Fl. VI, 2, 642; Krascheninn. in FI.
Yugo-Vost. Evrop. Ch. SSSR, VI, 360; Krascheninn. in Kryl. Fl. Zap.
Sib. XI, 2800; Grossh. Opred. Rast. Kavk. 464; Poljak. in Majevski,
Fl. 586.—A. repens Pall. ex Willd. Sp. pl. III (1800) 1840; Less. in
Linnaea, IX, p. 189.—A. nivea Redowsky ex Willd. Enum. pl. II (1813)
863.—A. frigida Eichw. Casp. cauc. (1833) 31.—A. frigidae var. folis
magis dissectis Ldb. Fl. Alt. IV (1833) 66.—Ic.: Hegi, Ill. Fl. VI, 2,
p. 642.—Exs.: GRF No. 1522.

Perennial. Whole plant whitish from dense silky hairs. Root with
slender, woody, creeping or obliquely ascending rhizomes. Stems few,
basally ascending or erect, 20-60 cm high, branched from base or
above with spreading branches, with densely leafy sterile shoots. Basal
leaves petiolate, 1.5-4.0 cm long and 1-3 cm wide, oval, twice pinnately
cut, lobes 1-2 pairs, palmately- or pinnately- cut into linear, acuminate,
2-12 mm long and 0.5-1.0 mm wide secondary lobes, other leaves
sessile, with basal lobes; upper leaves ternate; uppermost leaves
bracteal, undivided, linear. Capitula hemispherical or ovoid, about 2
mm in dia, divergent or drooping, on long peduncles, more or less in
broadly paniculate inflorescence. Involucral bracts hairy; outer bracts
almost linear or lanceolate-linear; inner oval, with scarious border.
Receptacle glabrous or with occasional floral[sic., receptacular bristles]
hairs (var. neglecta Krasch.) or with rather numerous hairs (var.
barabensis Serg.). Peripheral florets usually 5—7, their corollas very
small, almost filiform, broadened toward base; stigma lobes filiform,
divergent; disk florets bisexual, 7-8; their corollas narrowly linear,
apically hairy, anthers linear, apical appendage of anthers acute, basal
appendage short, subacute. Achenes about 1 mm long, oblong, flat,
with small convex areola at apex. Flowering August.

Alkaline meadows, sandy-loam steppe, sands near pine forests,
often in old fields, pastures, roadsides, near dwellings, in forest and
steppe zones.—European part: Karelia-Lapland, Dvina-Pechora,
Ladoga-Ilmen, Baltic Region, Upper Dnieper, Upper Volga, Volga-
Kama, Volga-Don, Trans-Volga, Lower Volga, Lower Don, Black Sea
Region, Middle Dnieper, Upper Dniester; Caucasus: Ciscaucasia,

500

474

Dagestan, Eastern Transcaucasia; Soviet Central Asia: Aralo-Caspian
Region (northern part), Lake Balkhash Region, Dzhungaria-Tarbagatai,
Tien Shan; Western Siberia: Upper Tobol, Irtysh, Ob River Area (south-
ern part), Altai; Far East: Ussuri (introduced). General distribution:
Central Europe, Western Mediterranean, Balkans-Asia Minor, Arme-
nia and Kurdistan, Iran. Described from Austria. Type in Vienna.

Note. I.M. Krascheninnikov (FI. Zap. Sib., XI, p. 2800) distin-
guishes var. neglecta Krasch. by the presence of short, sparse hairs on
the receptacle, but later (Mat. po Istor. fl. i Rast. III (1958) 91) he
reports that var. barabaensis Serg. is also distinguished by these very
same characters.

Economic Importance. According to the investigations of
Kazakevich and Sobolevskaya (1923), the yield of essential oil is 0.46%
(of weight of dry matter); at maturity the oil has the following constants:
D,°— 0.9319; D—3.44°; n,”'—1.47; acid number 6.33; ester number
47.66; ester number after acetylation—118.95. It is soluble in 70%
alcohol in the ratio of 1:1.8; in 80% alcohol 1:0.5; in 90% alcohol it
is entirely soluble. The oil contains: cineole—30%, a and B—
thujones—29%, thujyl alcohol—2% and thujylacetate—6%. According
to the opinion of Goryaev (1952), the oil apparently contains a
considerable quantity of azulene. Its reaction to aldehydes is positive,
to phenols and ketone, negative. According to the data of Dionisev
and Chernomaz (1938), the oil contains 0.667% alkaloids and 0.98%
glucosides, which apparently is poisonous to animals (Larin, 1956).

60. A. caespitosa Ldb. Fl. Alt. IV (1833) 80; Bess. in Bull. Soc.
Nat. Mosc. IX, 68; DC. Prodr. VI, 118; Ldb. Fl. Ross. II, 590; Kryl.
Fl. Alt. Ill, 651; Krascheninn. in Kryl. Fl. Zap. Sib. XI, 2801.—Ie.:
Ldb. Ic. pl. Fl. Ross. V, tab. 472.—Exs.: GRF No. 3155.

Perennial. Whole plant whitish or greenish-gray from fine and
long, silky, approximate hairs. Root slender, vertical, bearing short
(0.5—2.0 cm), strongly branched and spreading, perennial, sterile shoots
producing arise fertile, simple, erect or somewhat spreading 3-15 cm
high stems. Basal leaves small, undivided, linear-oblong, withering
before flowering; leaves above them petiolate, 1.0—1.5 cm long, ternate,
lobes lanceolate-linear, acute, 2-5 mm long; middle cauline leaves
divided into 3-5 lobes, sometimes pinnately cut, often with 2 lobes at
base; upper bracteal leaves ternate or undivided, lanceolate-linear. Capitula
sessile, globose-ovoid, 2-3 mm in dia, crowded or in depauperate
simple raceme or additionally on short lateral branches. Involucral
bracts oval-elliptical, convex, light greenish along midrib, with wide
scarious margin; outer bracts hairy, somewhat wider than sub-glabrous
inner bracts. Receptacle convex, glabrous. Peripheral florets pistillate,

501

475

9; their corollas narrowly tubular, with 4 teeth, glabrous, punctate-
glandular, in upper part purple-pink, stigma lobes exserted from corolla
tube, linear, obtuse, arcuate; disk florets bisexual, 14, their corollas
conical, in upper part purple, glabrous; anthers on shorter filaments,
linear, apical appendage of anthers long, acute, basal appendages short,
round; Stigma lobes after anthesis weakly exserted from corolla tube,
arcuate. Achenes up to 1.2 mm long, somewhat flat, oblong-ovoid,
dark brown, with flat{smooth] round apex. Flowering August.

Alkaline and poor rubbly soil, desert-steppe zone.—Western
Siberia: Altai (Chuya steppe). General distribution: Mongolia.
Described from Altai. Type in Leningrad.

61. A. cuspidata Krasch. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser.
1, 3 (1936) 349.

Perennial. Plant more or less densely caespitose, whitish from dense
long silky appressed hairs; sterile shoots rather numerous, 5-15 mm
high; fertile stems 1-2, erect, 6-10 cm high, simple, more or less
densely leafy. Leaves of sterile shoots and lower cauline leaves short-
petiolate, 5-10 mm long, ternate, lobes 4-7 mm long, linear or
lanceolate-linear, thin-acuminate; bracteal leaves shorter, undivided or
ternate, with short lateral lobes. Capitula hemispherical, 4-5 mm in
dia, 4-14, upward-spreading or divergent, on short peduncles; upper
capitula approximate, lower remote, in spicate or in lower part in
almost racemose inflorescence. Involucre with dense, appressed and
long hairs; outer bracts linear, inner linear-lanceolate, with wide,
scarious border. Receptacle hairy[setose]. Corolla of bisexual disk
florets tubular-conical, yellow, glabrous. Flowering August.

Stony steppe slopes, calcareous rocks, in forest zone.—Eastern
Siberia: Dauria (Olkhon Island and east bank of Lake Baikal), Lena-
Kolyma (Aldan River near mouth of Maya River). Described from
Lake Baikal. Endemic. Type in Leningrad.

62. A. schmidtiana Maxim. in Bull. Acad. Sc. Pétersb. VIII (1872)
537; Pampan. in Nouv. Giorn. Bot. Ital. n.s. XXXIV (1927) 699.—A.
sericea F. Schmidt, Fl. Sachal. in Mém. Acad. Pétersb. VII, XII, No.
2 (1868) 150; Sugaw. Ill. Fl. Saghal. IV, 1829; Hara, Enum.
Spermatoph. Japonic. 125.—Ic.: Sugaw, op. cit. tab. 838.

Perennial. Semishrub to 60 cm high, silvery from dense, slender,
appressed, silky hairs; stem at base subglabrous, densely leafy above,
branched, with straight, long, more or less divergent branches. Leaves
petiolate, 4-5 cm long, twice pinnately cut, lobes 2 pairs, remote,
linear, 1.5-2.0 cm long and about 1 mm wide, with 1-2 narrowly
linear, S—12 mm long lobules, apical bracteal leaves undivided, narrowly

502

476

linear, up to 1.5 cm long. Inflorescence densely leafy, pyramidal-pan-
iculate. Capitula on long leafy peduncles, hemispherical, 5-6 mm in
dia. Involucral bracts silvery-woolly; outer bracts linear; inner larger,
oblong, with whitish-scarious margin. Receptacle convex, hemispheri-
cal, tuberculate, setose. Peripheral florets pistillate, 3-8, narrowly
tubular, with 2 teeth, almost to tip covered with long white hairs;
stigma lobes exserted from corolla tube, linear, obtuse, arcuate; disk
florets bisexual, 21-33, their corollas conical, punctate-glandular, yel-
low, hairy in upper part, anthers lanceolate, apical appendages of anthers
elongate, basal appendages shorter, subobtuse; style usually shorter
than tube, stigma lobes linear, long, recurved at the end of vegetative
growth of plant. Achenes oblong, indistinctly ribbed, somewhat flat,
apex flat[smooth], scarcely bordered. Flowering September.

Sandy coasts and dry stony slopes.—Far East: Southern Sakhalin,
Kuril Islands (Sikotan). General distribution: Japan. Described from
Japan. Type in Leningrad.

Economic Importance. In Japan, sometimes cultivated as an or-
namental plant for its silvery leaves.

63. A. sericea Web. ex Stechm. Artem. (1775) 16; DC. Prodr. VI,
122; Ldb. Fl. Ross. II, 595; Maxim. in Bull. Acad. Sc. Pétersb. VIII,
537; Kryl. Fl. Alt. III, 653; Fedtsch. Perech. Rast. Turk. IV, 202;
Krascheninn. in Fl. Yugo-Vost. Evrop. Ch. SSSR, VI, 361; Kryl. Fl.
Zap. Sib. XI, 2795; Poljak. in Majevski, Fl. 585; Bobrov in Bot. Mat.
Gerb. Bot. Inst. XX, 16.—A. holosericea a. grandiflora and B.
parviflora Ldb. FI. Alt. IV (1833) 63; DC. Prodr. VI, 123.—A. sericea
var. gmeliniana and var. ledebouriana and var. pallasiana Bess. in
Bull. Soc. Nat. Mosc. IX (1836) 14.—A. sericea Stechm. var. steveniana
Bess. ibid. 14.—A. sericea Web. var. nitens (Stev.) DC. Prodr. VI
(1837) 122.—A. nitans (Bess.) Stev. ex Krasch. in Spisok Rast. Gerb.
Fl. SSSR, XI (1949) 50.—A. sericea var. grandiflora DC. and var.
parviflora DC. Prodr. VI (1837) 122; Ldb. Fl. Ross. II, 2, 595.—A.
czekanovskiana Trautv. in Tr. Peterb. Bot. Sada, V, 1 (1877) 72.—
A. sericea var. nitens f. parviflora Pamp. Nuov. Giorn. Bot. Ital. n. s.
XXXVI (1930) 700.—Absinthium sericeum Bess. in Bull. Soc. Nat.
Mosc. VIII (1829) 237.—A. grandiflorum Bess. ibid. 232.—A. nitens
Stev. ex Bess. ibid. 235.—Ic.: Gmel. FI. sib. II, 131, t. 64, fig. 1; Ldb.
Ic. pl. Fl. Ross. V, 459. imperfect—Exs.: GRF Nos. 3174a, 3175,
317G6a, 3292 32938:

Perennial. Root vertical, woody, bearing sterile densely leafy shoots
up to 20-35 cm high. flowering stems 40-70 cm high, few or single,
straight or at base ascending, brownish, pubescent in upper part,
subglabrous below, simple or branched above, leafy. Leaves on both

503

477

sides covered with slender, silky, appressed hairs but more densely
beneath and hence whitish-silvery; lower leaves withering early, smaller
than middle, petiolate, ternate or palmately cut, with undivided leaflets;
middle leaves 3-5 cm long and 2—4 cm wide, twice pinnately cut, with
1-2 pairs of lobes, terminal lobe pinnately cut but sometimes palmately
cut, undivided, linear or lanceolate-linear, 10-17 mm long and 1-2
mm wide, acute, with narrow, thick stripe and distinct midrib beneath
along margin, upper leaves sessile, simple pinnate or ternate. Capitula
hemispherical, 4-6(9) mm in dia, drooping, in narrowly paniculate-
racemose inflorescence, less frequently in deparperate panicle with
long lateral branches, or in simple elongate raceme. Involucral, bracts
whitish-hairy; outer bracts oblong-ovate or lanceolate, acuminate, with
narrow and brownish scarious margin; inner bracts longer than outer,
elliptical, with wide scarious-whitish margin. Receptacle convex,
hemispherical. Peripheral florets pistillate, 12-14; their corollas
narrowly tubular, toward base somewhat broadened, punctate-glandular;
stigma lobes exserted from corolla tube, linear, obtuse, arcuate; disk
florets bisexual, numerous (40-65); their corollas conical, hairy above,
glabrous below; anthers linear, apical appendages of anthers long, acute,
basal appendages short, round. Achenes to 1.2 mm long, oblong-conical,
angular-ribbed apically flat}smooth], along border unevenly toothed.
Flowering August.

Steppe, sometimes weakly saline meadows, along slopes of hillocks
and mountains, often rubbly and stony, in southern part of forest and
forest-steppe zones.—European part: Volga-Kama, Volga-Don, Trans-
Volga; Western Siberia: Upper Tobol, Irtysh, Altai; Eastern Siberia:
Angara-Sayans, Yenisei, Dauria, Lena-Kolyma; Soviet Central Asia:
Lake Balkhash Region (northern part). General distribution: Northern
Mongolia. Described from Siberia. The cited figure by Gmelin is the
type.

Note. The following forms can be distinguished 1) f. pallasiana
(Bess.) Poljak.—with small capitula, mainly associated with steppe
slopes and thickets of steppe shrubs in the central Urals; 2) f. nitens
(Stev.) Poljak.—with larger capitula, wide, ovate-lanceolate inner
involucral bracts, and a lax, weakly branched, often racemose panicle,
distributed mainly in mountainous Central and Eastern Siberia; 3) f.
czekanovskiana (Trautv.) Poljak.—with large many-flowered capitula
usually in a simple racemose inflorescence, distributed in the northern
part of Yakutia, evidently in the forest-tundra.

64. A. aschurbajevii Winkl. in Tr. Peterb. Bot. Sada XI (1892)
332; Fedtsch. Fl. Pam. (1903) 117; Fedtsch. Perech. Rast. Turk. IV,
203.—A. sericea var. turkestanica Winkl. op. cit. 203.—Exs.: GRF
No. 3294.

504

478

Perennial. Rhizome not thick, woody, creeping, bearing numerous,
short, sterile, densely leafy shoots, loosely caespitose; stems few,
sometimes numerous, (15)20—30(50) cm high, erect or at base arcuate,
ascending, weakly ribbed or subcylindrical, simple, leafy, more or less
hairy. Leaves on both sides densely covered with fine, silky, appressed
hairs; lower cauline leaves and leaves of sterile shoots petiolate, 1.5—
2.5(3.5) cm long and 1.2—2.0(3.2) cm wide, lamina round or reniform,
twice ternate or pinnately-ternate, lobes petiolulate, 3-5, usually
ternately divided, terminal lobes lanceolate, obtusely acuminate, more
often 5-10 mm long; middle cauline leaves short-petiolate or sessile;
uppermost bracteal leaves, undivided, linear-lanceolate. Capitula in
simple raceme, half as long as stem, on short peduncles; all capitula more
or less remote or only in lower part of inflorescence but approximately
above, hemispherical, 5—7 mm in dia, divergent or drooping. Involucral
bracts elliptical, herbaceous[sic.,? carinate] along midrib, hairy, with
wide, dark brown, scarious, irregularly incised margin. Receptacle
convex, hemispherical, hairy. Peripheral florets pistillate, 10, narrowly
tubular; their stigma lobes distinct, linear, recurved; disk florets
numerous (40-55), their corollas conical, in upper part hairy, below
glabrous, sometimes entirely hairy, anthers linear, apical appendages
of anthers long, acute, subulate, basal appendages considerably shorter,
subacute, style as long as corolla tube, stigma lobes erect, linear,
apically ciliate, recurved at maturity of achene. Achenes about 1 mm
long, oblong, indistinctly ribbed. Flowering July to August.

Mountain-meadow and meadow-steppe slopes of mountains, at
timberline.—Soviet Central Asia: Dzhungaria-Tarbagatai, Tien Shan,
Pamiro-Alai. Described from Pamir. Type in Leningrad.

Note. A. aschurbajevii C. Winkl. is very similar to A. sericea
Web. and is distinguished by a somewhat lower height of the stem,
simple racemose inflorescence, smaller size of the lamina, and by the
ecology and range.

Series 2. Crossostephioides Poljak.—Leaves linear-lanceolate,
toward base narrowly cuneate, in upper part undivided or with 3 short
or long teeth.

65. A. lagocephala (Bess.) DC. Prodr. VI (1837) 122; Turcz. FI.
baic.-dahur. II, 69; Maxim. in Bull. Acad. Sc. Pétersb. VIII, 531; Kom.
and Alis. Opred. Rast. Dalnevost. Kr. II, 1042.—A. lagocephala var.
kruhaiana (Bess.) Glehn in Maxim. op. cit. 532; Krascheninn. in Spisok.
Rast. Gerb. Fl. SSSR, XI, 48.—A. lagocephala var. besseriana (Bess.)
Pamp. f. a. integrifolia (Ldb.) Pamp. in Nuov. Giorn. Ital. Bot. n. s.
XXXIV (1927) 676.—A. besseriana var. triloba and var. integrifolia

505

479

Ldb. FI. Ross. II (1846) 590.—A. lithophila Turcz. Fl. baic.-dahur. II
(1856) 69, nom.—A. kruhsiana Bess. in Nouv. Mém. Soc. Nat. Mosc.
III (1834) 22; DC. Prodr. VI; Ldb. Fl. Ross. II, 576.—Absinthium
lagocephalum Fisch. ex Bess. in Bull. Soc. Nat. Mosc. VIII (1829)
233.—Ic.: Gmel. Fl. sib. II, tab. 61, f. 1, 2.—Exs.: GRF No. 3291a, b.

Perennial. Semishrub (20)30—60(70) cm high. Root woody, thick,
cordlike, with more or less numerous, strongly reduced, ascending,
woody, perennial, sterile shoots in upper part covered with brownish-
gray bark, with leaf rosettes at apex. Fertile stems rather numerous,
erect, ribbed, short-hairy, leafy, simple or branched in upper part, with
short or somewhat long, upright branches. Leaves short-petiolate or
subsessile, dark green and weakly hairy or subglabrous above, whitish,
finely tomentose beneath from appressed hairs; lower cauline leaves
and leaves of nonflowering shoots 3—7 cm long and 3-12 mm wide,
undivided, narrowly lanceolate, acuminate or undivided, obcuneate,
usually with 3-5 short, subacute or linear, longer, subobtuse teeth;
cauline leaves lanceolate or linear-oblong, short-acuminate or obtuse,
round, gradually cuneately narrowed toward base; bracteal leaves small,
not exceeding lateral shoots. Capitula hemispherical, narrowed toward
base, 4—7 mm in dia, on more or less long peduncles, divergent or
drooping, in narrow racemose inflorescence. Involucral bracts oblong-
lanceolate, with wide, scarious, incised-fimbriate margin; outer bracts
densely hairy; inner subglabrous or weakly hairy. Receptacle convex,
hemispherical, hairy, sometimes glabrescent. Peripheral florets pistillate,
9-14; their corollas narrowly tubular, with 2-3 teeth, punctate-glandular,
pubescent or glabrous, stigma lobes 2 (very rarely 3), narrowly linear,
obtuse, after anthesis weakly recurved; disk florets bisexual, 28—55-
80; their corollas up to 3 mm long, narrowly cup-shaped, conical, with
hairy teeth, glabrous, punctate-glandular, anthers lanceolate, apical
appendages of anthers long, acute, basal appendages small, obtuse,
stigma lobes linear, obtuse, ciliate. Achenes up to 2 mm long, oblong,
somewhat flat, indistinctly ribbed, with flat{smooth] apex, and short,
indistinctly toothed border. Flowering August. (Plate XXVI).

Stony alluvium and rocky outcrops in alpine tundra.—Arctic: Arctic
Siberia, Chukotka; Eastern Siberia: Angara-Sayans (eastern part),
Dauria, Lena-Kolyma; Far East: Zeya-Bureya, Ussuri, Okhotsk.
Endemic. Described from Eastern Siberia. Type in Kiev.

Note. The American species A. alaskana Rydb. is very similar to
this species.

For our species, it is possible to distinguish var. kruhsiana (Bess.)
Glehn., which is characterized by toothed incised basal leaves, partly
by the absence of receptacular hairs, and partly by the simple recemose
inflorescence; it grows in forest-tundra and partly in tundra.

506

480

Series 3. Rutifoliae Poljak.—Leaves twice pinnately cut or twice
ternately divided.

66. A. rutifolia Steph. ex Spreng. Syst. Veg. III (1826) 488; Bess.
in Nouv. Mém. Soc. Nat. Mosc. III, 22; DC. Prodr. VI, 106; Ldb. FI.
Ross. II, 576; Maxim. in Bull. Acad. Sc. Pétersb. VIII, 531;
Krascheninn. in Kryl. Fl. Zap. Sib. XI, 2789.—A. turczaninowiana
Bess. op. cit. 23; DC. Prodr. op. cit. p. 106; Ldb. op. cit. 577; Boiss.
Fl. or. III, 369; Maxim. in Bull. Acad. Sc. Pétersb. VIII, 531; Kryl. FI.
Alt. III, 641; Fedtsch. Perech. Rast. Turk. IV; 196; Pampan. in Nouv.
Giorn. Bot. Ital. n. s. XXXIV [sic., recte XXXIV], 706.—A.
turczaninoviana §. dasyantha Schrenk Enum. pl. novar. (1841) 50.—
A. turczaninowiana var. falconeri (Clarke) O. Fedtsch. Second
Addendum to FI. Pam. (1901) No. 82.—A. turczaninowiana var. altaica
Kryl. op. cit. 641.—A. falconeri Clarke in Hook. FI. Br. Ind. III (1882)
320.

Perennial. Semishrub, 25-80 cm high, with stocky, rather thick
and strongly branched, woody stem covered with brownish-gray bark
bearing numerous strongly leafy, erect or in part somewhat spreading
branches that together with the dead remnants of previous year form
more or less thick turf. All leaves petiolate, lacking lobes at base,
grayish from fine felt of silky appressed hairs on both sides like young
branches; lamina almost round or reniform, 0.6—2.0 cm long and 0.8—
3.0 cm wide, twice pinnately cut or twice ternate, primary lobes 5,
ternate or pinnately divided, secondary lobes linear-oblong, subobtuse,
2-12 mm long and 0.5-1.5 mm wide; lowermost as well as upper
leaves less compound, sometimes ternately divided; upper bracteal
leaves undivided, linear. Capitula hemispherical, 3.5—-5.0 mm in dia,
pedunculate, divergent or drooping, in racemose or paniculate
inflorescence; common receptacle glabrous or hairy. Involucre cup-
shaped; outer involucral bracts oblong, whitish-tomentose, inner
elliptical or oval, subglabrous, with wide scarious margin. Peripheral
florets pistillate, 5-15, their corollas narrowly tubular, toward base
gradually broadened, glabrous, punctate-glandular; stigma lobes
somewhat exserted from corolla tube, linear, recurved; disk florets
bisexual, 12-28, their corollas conical, glabrous or hairy, punctate-
glandular; anthers linear, apical appendages of anthers long, scarcely
acuminate, basal appendages short, acuminate; stigma lobes linear,
apically broadened, ciliate, after flowering recurved. Achenes oblong-
prismatic, angular-ribbed, with flat{smooth] apex and uneven border.
Flowering August. (Plate XXII, Fig. 2).

Mountain-steppe, stony slopes of mountains.—Western Siberia:
Altai; Eastern Siberia: Angara-Sayans, Dauria; Soviet Central Asia:

507

481

Dzhungaria-Tarbagatai, Tien Shan, Pamiro-Alai Region. General
distribution: Iran-Afghanistan (Afghanistan), western Himalayas,
Dzhungaria-Kashgaria, Mongolia, Tibet. Described from Siberia. Type
in Leningrad.

Economic Importance. According to the data of Bazilevskaya
(1936), the leaves and capitula contain 0.85% essential oil in the dry
matter, but according to the data of S.N. Kudryashev (1936)—0.21-
0.36%. According to the data of M.I. Goryaev et al. (1954), the plants
collected in 1950 from Dzhungarian Alatau in the vicinity of the village
of Tsaritsyno yielded 0.48% oil of the dry weight of the herb. The oil
is light green in color, with flowery odor and partially crystalizes on
cooling; it shows a weak reaction to alkaloids and a negative reaction
to phenols, cineole and azulene-forming sesquiterpenes. Constants of
the oil are: nD—1.4668; acid number 6.5; ester number 129.5;
saponification number 135.49. Oil obtained from plants collected in
1951 had the following constants: D*°—0.9278; n,'*—1.4695; acid
number 2.02; ester number 30.51; saponification number 32.56; ester
number after acetlyation 93.76. The oil was found to contain 8.4% of
bound alcohols and 17.4% of free alcohols. Its reaction to phenol was
negative; reaction to carbonyl compound was positive; and the reaction
to azulene-forming sesquiterpenes was negative.

According to P.S. Massagetov’s (1947) analysis, aerial parts of
the plant contain many alkaloids.

Series 4. Persicae Poljak.—Leaves thrice pinnately cut.

67. A. persica Boiss. Diagn. ser. I, 6 (1855) 91; FI. or. II, 373;
Pampan. in Nouv. Giorn. Bot. Ital. n. s. XXXIV, 685; Pampan. in
Lavori Inst. bot. Univ. Cagliari, XL, 577.—A. togusbulakensis B.
Fedtsch. in Tr. Bot. Muz. 1 (1902) 143; Fedtsch. Perech. Rast. Turk.
IV,,.497;

Perennial. Semishrub, 25-70 cm high, grayish-green from dense
crisped hairs. Root woody, thick. Stem perennial, woody, thick, strongly
reduced, branching from base, erect or ascending, covered with grayish-
brown, cracking bark; flowering stems light greenish, erect, long, ribbed,
leafy, simple or with more or less long, divergent or upright shoots.
Lower and middle cauline leaves usually short-petiolate, 0.8—1.5 cm
or more often 2.0—3.5(4.5) cm long (f. hissarica Poljak.), lamina round,
thrice pinnately cut, terminal lobes narrow-lanceolately linear or linear,
short acuminate or obtuse, upper cauline leaves sessile, smaller, and
less divided. Capitula more or less remote, on short or long peduncles,
in simple raceme or narrow panicle (f. hissarica Poljak.), hemispherical,
3-5 mm in dia. Involucral bracts densely hairy, outer bracts herbaceous,

508

482

lanceolate-linear, inner broadly ovate, herbaceous [sic.? carinate?] along
midrib, with wide scarious margin. Receptacle convex, hairy or
subglabrous. Peripheral florets pistillate, 10; their corollas narrowly
tubular, with 2 teeth, punctate-glandular, glabrous, stigma lobes exserted
from corolla tube, linear, obtuse, divergent, recurved; disk florets
bisexual, numerous (30-40), their corollas narrowly cupshaped, glabrous
or more often weakly hairy; anthers linear-lanceolate, apical appendages
of anthers acute, basal appendages short, acute; style included in corolla
tube, stigma lobes linear, erect. Achenes 1 mm long, oblong-conical,
angular-ribbed, apically with flat areola, with uneven border. Flowering
August. (Plate XXI, Fig. 2).

Rubbly slopes of mountains, sometimes on talus, gravel-beds,
reaching 3,000 m and above.—Soviet Central Asia: Tien Shan (western
part), Pamiro-Alai Region. General distribution: Iran, Indo-Himalayas,
western Tibet. Described from Iran. Type in Geneva.

Economic Importance. According to the investigations of M.Kh.
Khodzhinov and I.P. Tsukervanik (1948), the plants collected from the
Chimgan District contained: about 12% a-pinene, about 16% camphors,
about 23% camphene, about 33% |-borneol and |-bornylacetate. In the
plants collected from the Bostandyk Region, according to the data of
M.I. Goryaev, T.E. Serkibaeva, and L.A. Iganatova (1953), the yield
of essential oil was 0.35% of the air-dried material. The oil was found
to contain a-pinene—7.2%, borneol—18.3%, esters of borneol, mainly
bornylcapronate—25.5%, camphene—14% and phenols—1.43%.
According to the data of Goryaev and R.N. Sazonova (1959), this
essential oil contains phenols, ketones, and azulene-forming
sesquiterpenes.

Series 5. Hololeucae Poljak.—Plants caespitose, whitish from dense
appressed arachnoid hairs.

68. A. hololeuca MB. ex Bess. in Nouv. Mém. Soc. Nat. Mosc.
III (1834) 46; DC. Prodr. VI, 112; Ldb. Fl. Ross. II, 583; Krascheninn.
in Spisok Rast. Gerb. Fl. SSSR, X, 85; Poljak. in Majevskii, Fl. 585.—
Exs.: GRF No. 3167.

Perennial. Whole plant whitish from appressed arachnoid hairs.
Root woody, thick, with short, perennial, woody, ascending, densely
leafy shoots in upper part, loosely caespitose. Fertile shoots few, (5)20-
30(35) cm high, ribbed, flexuous, ascending, weakly leafy, branched.
Leaves of sterile shoots and lower cauline leaves long-petiolate, 2.5—
6.0 cm long, ovate or oblong, twice pinnately cut, primary lobes 2—5
pairs, remote, secondary lobes 2-6 mm long and about 1 mm wide,
oblong-linear, flat, scarcely acuminate, with sharply raised midrib and

509

483

thick margin; middle and some upper cauline leaves simple pinnate;
uppermost bracteal leaves undivided, linear. Capitula in narrowly
pyramidal panicle, on long peduncles, sometimes sessile, drooping,
campanulate, 2.5—4.0 mm in dia. Involucral bracts whitish hairy, ovate
or ovate-oblong, with scarious margin; outer bracts acute, shorter than
inner subobtuse. Receptacle convex or flat, sometimes hairy. Peripheral
florets pistillate, fewer (usually 4—6), their corollas narrowly tubular,
punctate-glandular; stigma lobes exserted from [corolla] tube, linear,
truncate, arcuate; disk florets bisexual, 20-24, their corollas conical,
glabrous, punctate-glandular; anthers linear-lanceolate, apical
appendages of anthers acute, basal appendages shorter, subobtuse; style
long; stigma lobes linear, ciliate at apex, after flowering arcuate.
Achenes to 1.5 mm, oblong ovate, somewhat flat, smooth, with flat
round areola at apex, scarcely bordered. Flowering August.

Chalk outcrops, sometimes in river valleys on sand.—European
part: Volga-Don, Middle Dnieper, Lower Don. Endemic. Described
from the Voronezh and Kharkov regions. Type in Leningrad.

Series 6. Rupestres Poljak.—Plants caespitose; stems prostrate,
ascending; leaves glabrous or weakly hairy beneath; terminal lobe
cartilaginous, short-acuminate; receptacle hairy.

69. A. ruprestris L. Sp. pl. (1753) 841; Ldb. Fl. Alt. IV, 67;
Bess. in Bull. Soc. Nat. Mosc. IX, 20 (incl. var. v. willdenoviana
Bess. and var. thuringiaca Bess. and var. oelandica Bess.); DC. Prodr.
VI, 124; Ldb. Fl. Ross. II, 597; Maxim. in Bull. Acad. Sc. Pétersb.
VIII, 537; O. Fedtsch. Fl. Pam. 117; Kryl. Fl. Alt. III (1904) 652 p.
p.; Fedtsch. Perech. Rast. Turk. IV, 201; Hegi, Ill. Fl. VI, 2, 656;
Pampan. in Nouv. Giorn. bot. Ital. n. s. XXXIV (1927) 686; Kraschen.
in Spisok Rast. Gerb. Fl. SSSR, X (1936) 88; Kraschen. in Fl. Yugo-
Vost. Evrop. Ch. SSSR, VI, 362; Kryl. in Fl. Zap. Sib. XI, 2794.—A.
dentata Willd. Sp. pl. II (1800) 1826.—A. viridis Willd. ibid. 1829.—
Kryl. Fl. Zap. Sib. XI, 2794.—A. viridifolia Ldb. Ind. Sem. Horti
Dorpat. (1832) 2; Spreng. Syst. vet. III, 493; Ldb. op. cit. 597.—A.
ruprestris f. salsuginea Kryl. and f. alpina Kryl. op. cit. (1904) 652.—
Absinthium viridifolium rupestre and viride Bess. in Bull. Soc. Nat.
Mosc. VIII (1829) 246.—Exs.: GRF Nos. 1368, 3173a, b, c, 3295.

Perennial. Stems woody at base, more or less branched, prostrate
or spreading developing few short, annual, densely leafy, basally
ascending nonflowering shoots and besides, erect, brownish-violet,
fertile, simple shoots, 20-50 cm or 10-25 cm high (var. viridis DC.),
more or less densely hairy in upper parts. Leaves green, glabrous or
with scattered (var. viridis DC.), usually bifid hairs; lowermost leaves

510

484

petiolate, others sessile, 1.5-5.0 cm long and 1.0—2.5 cm wide, twice
pinnately cut, lower lobes rather small, undivided, middle and upper
ternate or pinnately cut, terminal lobes lanceolate-linear or with
cartilaginous, short-acuminate apex, 1-6 mm long and 0.5—1.0 mm
wide, upper bracteal leaves simple pinnate. Capitula globose, 4-7 mm
in dia, inclined or drooping, in narrowly racemose-spicate inflorescence.
Involucre more or less hairy; outer involucral bracts oblong-linear,
sometimes pinnately divided, inner bracts somewhat shorter, elliptical,
with scarious margin. General receptacle convex, hemispherical, with
floral hairs. Peripheral florets pistillate, 16, their corollas narrowly
tubular, punctate-glandular, stigma lobes somewhat exserted from tube,
linear, obtuse, after flowering somewhat recurved or divergent; disk
florets bisexual, numerous (to 66); their corollas conical, glabrous,
punctate-glandular; anthers as long as filaments, oblong-linear, apical
appendages of anthers narrowly linear, roundish, basal appendages short,
subacute, style shorter than corolla; stigma lobes short, linear, ciliate.
Achenes small, oblong-ovoid, finely sulcate, with flat areola at apex,
scarcely broadened. Flowering August.

Steppe zone, saline and alkaline meadows, sometimes in nonsaline
steppe meadows in deciduous forests, stony slopes and limestone
outcrops.—European part: Baltic Region (Oesel[Saaremaa], Gapsal,
Moon islands, eastern part of Estonia), Volga-Kama; Western Siberia:
Upper Tobol, Irtysh, Altai; Eastern Siberia: Anagara-Sayans, Dauria (basin
of Barguzin River); Soviet Central Asia: Dzhungaria-Tarbagatai, Tien Shan,
Pamiro-Alai Region. General distribution: Atlantic Europe, Scandinavia
(Gotland Island). Described from Gotland Island. Type in London.

Series 7. Obtusliobae Poljak.—Stems erect or partly ascending,
branched; terminal lobes of leaves usually obtuse, round; receptacle
hairy or glabrous.

70. A. obtusiloba Ldb. Fl. Alt. IV (1833) 68, 70 excl. B.
fruticulosa; Bess. in Bull. Soc. Nat. Mosc. IX, 9; DC. Prodr. VI, 121;
Ldb. FI. ross. II, 594; Kryl. Fl. Alt. III, 682; Kraschen. in Kryl. FI.
Zap. Sib. XI, 2792.—A. glabella Kar. and Kir. in Bull. Soc. Nat.
Mosc. XIV (1841) 441; Ldb. FI. Ross. II, 577; Kryl. Fl. Alt. ITI, 641;
Fedtsch. Perech. Rast. Turk. III, 196; Kraschen. in Spisok Rast. Gerb.
Fl. SSSR, XI, 44; Kryl. Fl. Zap. Sib. XI, 2793.—Ic.: Ldb. Ic. pl. Fl.
Ross. V, tab. 465, 466.—Exs.: GRF No. 3281.

Perennial. Root thick, woody, with erect or ascending, branched,
perennial, woody, short, sterile shoots and 30-40 cm high flowering
shoots. Leaves bright green, punctate-glandular, densely pilose or
subglabrous at beginning of vegetative growth, except lower leaves,

513

485

sessile, 6-20 mm long and 6-17 mm wide, with ternate or palmately
parted auricles, lamina palmately or lyrately pinnate, lower lobes
petiolate, undivided, upper closer to leaf apex, usually ternate or
palmately cut, secondary lobes oblong-lanceolate or linear, spatulate;
upper and bracteal leaves pinnately cut or simple, linear. Capitula
globose, 2—3(4) mm in dia, pedunculate or subsessile, inclined or
drooping, in more or less narrowly paniculate inflorescence. Outer
involucral bracts oblong, inner bracts wider, scarious along margin
and often hairy. Receptacle convex, hemispherical, hairy or glabrous
(var. glabella (Kar. and Kir.) Poljak). Peripheral florets pistillate, 12;
their corollas narrowly tubular, stigma lobes exserted from tube, linear,
divergent; disk florets bisexual, 20-24; their corollas conical, with
occasional hairs at apex, sometimes lacking them, punctate-glandular,
anthers linear, apical appendages of anthers narrowly linear, subacute,
basal appendages shorter, acuminate, stigma lobes short, linear, obtuse,
ciliate. Achenes to 0.8 mm long, oblong-conical, somewhat flat, brown,
with roundish areola at apex. Flowering August.

Stony, desert-steppe slopes of [mud]volcanic cones and
mountains.— Western Siberia: Irtysh (southeastern part), Altai (south);
Soviet Central Asia: Lake Balkhash Region. General distribution:
Mongolia (Mongolia Altai). Described from Katun River valley, near
mouth of Chuya River. Type in Leningrad.

Series 8. Subviscosae Poljak.—Stems for most part erect; leaves
somewhat sticky, their terminal lobes short-acuminate; receptacle hairy
or glabrous.

71. A. subviscosa. Turcz. in Bull. Soc. Nat. Mosc. I (1838) 649,
nomen; Bes. in Bull. Soc. Nat. Soc. Nat. Mosc. IX, 24; Ldb. Fl. Ross.
II. 594; Turcz. Fl. baic.-dahur. II, 68; Maxim. in Bull. Acad. Sc. Pétersb.
VIII, 537; Pampan. in Nuov. Giorn. Bot. Ital. n. s. XXXIV, 705.—A.
capillifolia Fisch. ex DC. Prodr. VI (1837) 122.—Exs.: GRF No. 1168.

Perennial. Whole plant, except lower part of stem more or less
densely covered with appressed bifid hairs and punctate glands. Root
thick, woody, with perennial, short, leafy, sterile shoots and numerous
fertile shoots. Stems in lower part glabrous, brown, 25-35(45) cm
high. Lower and, in part middle leaves petiolate, 1.5—-2.3 cm long,
twice pinnately cut, lobes fewer, ternate, terminal lobe linear, short-
acuminate, 3-6 mm long and 0.4-0.8 mm wide; upper leaves sessile,
bracteal, ternate, palmate or undivided, linear. Capitula subglobose,
3-4 mm in dia, on very short peduncles, inclined, approximate or
somewhat remote, in narrowly racemose-paniculate or almost simple
racemose inflorescence. Involucral bracts lanceolate, hairy; outer bracts

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Plate XXIV.

486

511

Artemisia adamsii Bess., habit, lower cauline leaf, capitulum, involucral bract,

peripheral pistillate floret, bisexual disk floret.

514

487

herbaceous, inner bracts slightly shorter, with narrow herbaceous spine
and wide scarious border. Receptacle convex, hairy. Peripheral florets
pistillate, 10; their corollas narrowly tubular, punctate-glandular,
sometimes weakly hairy; stigma lobes exserted from tube, linear, obtuse,
divergent; disk florets bisexual, 20—22; their corollas conical, glabrous,
yellow or pinkish, punctate-glandular; anthers as long as filaments,
linear, apical appendages of anthers subulate, basal, appendages short,
subacute; stigma lobes short, linear, ciliate, divergent at apex after
anthesis. Achenes about 1 mm long, oblong conical, somewhat flat, sulcate,
with a round areola at apex, scarcely bordered. Flowering August.

Dry open slopes and plains.—Dauria (near Lake Baikal). Endemic.
Described from Baikal. Type in Leningrad.

72. A. martjanovii Krasch. ex Poljak. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XVII (1955) 407.—Exs.: GRF No. 1168, sub
nom. A. subviscosa Turcz.

Perennial. Root thick, woody, with short, erect, sterile, leafy shoots
and rather numerous flowering shoots, glabrous below, hairy above;
stem 20-35 cm high. Leaves rugose-punctate, with bifid appressed
hairs; lower leaves petiolate, withering early 1.5—2.5 cm long, twice
pinnately cut, lobes few, ternate, terminal lobule linear, short-acuminate,
3-6 mm long and 0.2-0.5 mm wide; middle and upper leaves usually
sessile, simple pinnately cut. Capitula subglobose, 4-5 mm in dia, on
short peduncles, divergent or drooping, approximate or somewhat
remote, in narrowly paniculate inflorescence. Involucral bracts more
or less densely hairy, with wide scarious margin; outer bracts elliptical,
inner bracts ovate. Receptacle glabrous. Florets mostly fertile; peripheral
florets pistillate, 10, their corollas narrowly tubular, in lower part
slightly enlarged, punctate-glandular; stigma lobes half exserted from
tube, linear, obtuse, somewhat spreading; disk florets bisexual,
numerous (to 45), corolla conical, glabrous; anthers linear, apical
appendages of anthers acute, basal appendages short, subacute; stigma
lobes short, linear, after anthesis strongly recurved. Flowering August.

Stony slopes of lower mountains and sandy soils in steppes.—
Eastern Siberia: Angara-Sayans (Minusinsk District). Endemic.
Described from vicinity of Minusinsk. Type in Leningrad.

Note. The species is similar to A. subviscosa Turcz. but differs
from it by the glabrous peduncles, large involucres, and large number
of florets.

73. A. altaiensis Krasch. in Kryl. Fl. Zap. Sib. XI (1949) 2792.—
A. obtusiloba £. fruticosa Ldb. Fl. Alt. IV (1833) 69; DC. Prodr. VI,
121, Ldb. Fl. Ross. II, 594.—A. obtusiloba Kryl. Fl. Alt. III (1904)
682, partim. non Ldb.—Ic.: Ldb. Icones pl. Fl. Ross. V, tab. 466.

515

488

Plant moderately hairy with bifid appressed hairs; root 0.4—2.5 cm
thick, at collar woody, many-headed, bearing perennial, woody, more:
or less strongly and irregularly branched and spreading, short shoots;
fertile shoots numerous, 5-20 cm high, erect. Leaves bright green,
punctate-glandular besides, long, bifid hairs on young leaves, later
subglabrous, except lower leaves, sessile, 6-20 mm long and 4-12 mm
wide, pinnately or palmately parted, lower lobes smaller and undivided;
upper lobes larger, ternate or digitately divided into lanceolate, scarcely
acute secondary lobes 1.5—3.0 mm long and 0.5—1.5 mm wide; upper
and bracteal leaves less divided, with undivided lobes. Capitula globose,
4—6 mm in dia, crowded, partly remote, spreading or almost drooping,
in simple raceme or in dense racemose panicle. Involucre hairy; outer
involucral bracts oblong or sublanceolate, as long as elliptical inner
bracts, scarious on margin. Receptacle hairy. Peripheral florets pistillate,
15, their corollas narrowly tubular, enlarged at base, punctate-glandular;
stigma lobes exserted from [corolla] tube, linear, obtuse, erect, or
curved; disk florets bisexual, numerous (to 50), their corollas conical,
reddish in upper part, punctate-glandular, glabrous or weakly hairy;
anthers oblong-lanceolate, apical appendages of anthers attenuate, acute,
basal appendages shorter, subobtuse; stigma lobes linear, apically
enlarged, obtuse, ciliate, after anthesis exserted from tube, arcuate.
Achenes 1.2—1.5 mm long, oblong-pyriform or ovoid, glabrous, with
filiform longitudinal ribs, with a flat round areola at apex. Flowering
July to August.

Rubbly desert-steppe slopes of mountains, sometimes on ancient
moraines, on clayey, alkali soils of plains —Western Siberia: Altai
(southeast). General distribution: Western Mongolia (Mongolian Altai).
Described from Altai. Type in Leningrad.

Series 9. Frigidae Rydb. North. Am. FI. 34, part 3 (1916) 258.—
Annuals, biennials or perennials, herbaceous, more or less densely
white-hairy, less often subglabrous; capitula usually hemispherical or
globose, sometimes broadly campanulate; receptacle hairy or glabrous.

74. A. absinthium L. Sp. pl. (1753) 848; MB. FI. taur.-cauc. I,
297; Ldb. Fl. Alt. IV, 62; DC. Prodr. VI, 125; Ldb. Fl. Ross. II, 598;
Hook. FI. Br. Ind. III, 328; Boiss. Fl. or. III, 373; Kryl. Fl. Alt. III,
655; Fedtsch. Perech. Rast. Turk. IV, 202; Rydb. op. cit. 258; Hall
and Clements. Artem. 106; Pampan. in Nouv. Giorn. Bot. Ital. n. s.
XXXIV, 635; Hegi, Ill. VI, 2, 651; Grossh. Fl. Kavk. IV, 139; Kraschen.
in Fl. Yugo-Vost. Evrop. Ch. SSSR, VI, 363; Kraschen. in Kryl. Fl.
Zap. Sib. XI, 2817; Poljak. in Majevski, Fl. 586.—Absinthium vulgare
Lam. Fl. franc. II (1778) 45; Bess. in Bull. Soc. Nat. Mosc. VIII

516

489

(1829) 259.—A. bipetala Gilib. Fl. lithuan. II (1782) 174.—Exs.: GRF
No. 3170.

Perennial. Whole plant grayish from short approximate hairs,
sericeous-tomentose. Root vertical, thick, perennial. Stem 60-100 cm
high, herbaceous, erect, with short branches, leafy, sometimes basally
with short nonflowering branches bearing long-petiolate leaves. Leaves
with 6-9 cm long and 3-7 cm wide lamina broadly ovate, almost
thrice pinnately dissected, terminal lobe lanceolate, short-acuminate;
middle cauline leaves short petiolate, lacking lobes at base of petiole,
twice pinnately dissected; upper leaves subsessile, simple pinnate or
twice ternate; bracteal leaves ternate or simple, undivided, narrowly
lanceolate; lobes of all leaves linear-oblong, scarcely acuminate,
undivided or with few teeth, usually 3-20 mm long and 1-4 mm wide.
Capitula globose, 2.5-3.5 mm in dia, drooping or spreading into
narrowly paniculate inflorescence. Involucral bracts oblong-ovate,
[outer] almost as long as inner bracts, hairy on spine, with wide scarious
margin. Receptacle convex, hairy. Peripheral florets pistillate, usually
25, their corollas filiform-tubular; stigma lobes exserted from tube,
linear, arcuate; disk florets numerous (usually 60), their corollas conical,
glabrous, sometimes weakly hairy; anthers linear, apical appendages
of anthers obtusely angular, with round apex, basal appendages short,
obtuse; stigma lobes broadly linear, after flowering somewhat curved.
Achenes about 1 mm long, oblong-cuneate, rather flat, finely sulcate,
with a round, slightly convex areold at apex. Flowering July to August.

Weed, on recent fallow lands, near roads, dwellings, in kitchen
gardens and gardens, less often in crop fields, meadows, pastures and
forest edge; in forest, forest-steppe and steppe zones, as well as in
mountains.—European part: Karelia-Lapland, Dvina-Pechora, Ladoga-
Ilmen, Baltic Region, Upper Dnieper, Upper Volga, Volga-Kama,
Trans-Volga, Volga-Don, Upper Dnieper, Middle Dnieper, Upper
Dniester, Bessarabia, Black Sea Region, Crimea, Lower Don, Lower
Volga; Caucasus: Ciscaucasia, Dagestan, Eastern, Western, and
Southern Transcaucasia, Talysh; Western Siberia: Upper Tobol, Irtysh,
Ob River Area, Altai; Eastern Siberia: Angara-Sayans; Soviet Central
Asia: Aralo-Caspian Region, Lake Balkhash Region, Dzhungaria-
Tarbagatai, Tien Shan, Syr-Darya, Pamiro-Alai Region, mountainous
Turkmenia. General distribution: Scandinavia (except the north), Cen-
tral Europe, Atlantic Europe, western Mediterranean Region (exclud-
ing Italy), eastern Mediterranean Region, Balkans-Asia Minor, Arme-
nia and Kurdistan, Iran-Afghanistan, Indo-Himalayas, northern Africa,
North America. Described from Western Europe. Type in London.

Note. A. absinthium L. is mostly propagated through seeds, but it
also has a highly developed capability for vegetative reproduction. In

5! /

490

the first year of the plant’s life, adventitious buds develop on the root
collar, which in the second year produce shoots that bear fruits already
in the third year (age of the mother plant). When separated from the
mother plant these are capable of producing normally developed plants
identical with the mother plant, producing a large quantity of seeds
every year and, in turn, developing buds. Common wormwood is not
a troublesome weed, as it is very rare in standing crops; in the case of
a significant presence, it can easily be eradicated by simply uprooting
the plants, because its adventitious buds, as mentioned above, are found
on the root collar.

Sometimes this wormwood occurs extensively in the hay meadows
and pastures, significantly lowering the fodder value of the holdings,
since, eaten by the cattle even in small quantity, it imparts a bitter
taste and wormwood smell to the milk and milk products (I.P. Petrov,
Selskoe Khozyaistvo i Lessovodstvo [Agriculture and Forestry], (1902).

Economic Importance. According to different published reports
(M.I. Goryaev, 1953), essential oil of American origin contains 11.6—
24.5% thujyl alcohol, thujone, phellandrene, and pinene; 24—1-36.2%
thujylacetate and cadinene; 0.25% acetic, isovaleric, and palmitic ac-
ids; Italian oil contains 10% a- and B-thujone, thujyl alcohol, and its
esters of acetic, isovaleric, and palmitic acids. According to the data
of Kazakevich and Sobolevskaya, the oil contains a significant quantity
of paraffin in addition to phallandrene and cadinene. The essential oil
of A. absinthium L. is used in the liqueur-vodka industry for making
vodkas and liqueurs, absinthe and vermouth and, in some European
countries, for making beer (Hall and Clements, 1923). According to
the investigations conducted by Troitsky (1950), the aerial part of the
plant contains a rather significant quantity of carotene; according to
Lazurevsky and Sadykov (1945), alkaloids were absent or found only
in traces (Bankovsky et al.). Common wormwood contains a significant
amount of protein (15.6%) dnd, at the same time, also a rather consid-
erable quantity of cellulose (34.1%). Domestic animals do not graze it;
beginning in spring or during a shortage of fodder only the leaves and
inflorescence are eaten. When cows graze on wormwood, the milk, as
stated before, becomes bitter (Parashchuk, 1932). According to the
observations of Zvonnikov (1936), wormwood hay is greatly relished
by rabbits.

Its decoctions and infusions have been used in folk medicine since
ancient times. Even now, they are very often used in the treatment of
rheumatism, stomach disorders, malaria, for improving the appetite,
etc. (Belyaeva, 1946).

518

491

75. A. sieversiana Willd. Sp. pl. II, 3 (1804) 1845; Ldb. FI. Alt.
IV, 61; DC. Prodr. VI, 126; Ldb. Fl. Ross. I, 599; Maxim. in Bull.
Acad. Sc. Pétersb. VIII, 537; Hooker. Fl. Br. Ind. III, 329; Kryl. Fl.
Alt. III, 656, excl. var. B.; Kom. Fl. Manchzh. III, 679; Nakai, FI.
Korean. II (1911) 29; Fedtsch. Perech. Rast. Turk. IV, 204; Pampan.
in Nuov. Giron. Bot. Ital. n. s. XXXIV, 700; Kom. and Alis. Opred.
Rast. Dalnevost. Kr. II, 1032; Krascheninn. in Fl. Yugo-Vost. Evrop.
Ch. SSSR, VI, 362; Pampan. in Lavori Inst. bot. Univ. Cagliari, XL,
570; Krascheninn. in Kryl. Fl. Zap. Sib. XI, 2818; Poljak. in Majevski,
Fl. 586—Absinthium sieversianum Bess. in Bull. Soc. Nat. Mosc. VIII
(1829) 259.—Exs.: GRF No. 3169a, b, c.

Annuals, biennials. Whole plant grayish or whitish from densely
appressed hairs. Root vertical, slender. Stem 30-100 cm high, erect,
strongly ribbed, branched. Leaves punctate-glandular; lower and middle
cauline leaves long-petiolate, 1.5-12.0 cm and even longer; lamina
broadly triangular, thrice or twice pinnately dissected into oblong or
linear-oblong, obtuse or round, 2-10 mm long and 0.5—12.0 mm wide,
secondary lobes upper and bracteal leaves sessile, less divided,
uppermost undivided, linear. Capitula numerous, hemispherical 4-6
mm in dia, drooping, in rather wide, paniculate inflorescence. Outer
involucral bracts hairy, linear-oblong, inner bracts elliptical or almost
round, with wide scarious margin. Receptacle convex, hemispherical,
hairy. Peripheral florets pistillate, 18, their corollas narrowly tubular,
gradually enlarged toward base, punctate-glandular; stigma lobes
exserted from tube, narrowly linear, obtuse, straight, weakly spreading;
disk florets bisexual, numerous (about 100), their corollas conical,
glabrous, anthers oblong-lanceolate, apical appendages of anthers long,
almost subulate, basal appendages subacute; stigma lobes exserted from
tube after anthesis, ciliate at apex, straight, later curved. Achenes ovoid,
almost flat, slender, sulcate, with a round flat areola at apex. Flowering
August.

Steppe and southern part of forest zone; steppe, alkaline and, less
often, dry-valley meadows, open groves, sometimes along coastal slopes,
often as weed near dwellings, roadsides, fields and fallow lands.—
European part: Trans-Volga (introduced), Volga-Kama; Western Sibe-
ria: Upper Tobol, Ob River Area, Irtysh, Altai; Eastern Siberia: Yenisei,
Angara-Sayans, Dauria, Lena-Kolyma (Vilyui District); Far East: Zeya-
Bureya, Uda River Area, Ussuri; Soviet Central Asia: Aralo-Caspian
Region, Lake Balkhash Region, Dzhungaria-Tarbagatai, Tien Shan,
Pamiro-Alai Region. General distribution: Himalayas (Kashmir),
Western Tibet, China, Mongolia, Japan. Described from Siberia. Type
in Berlin.

492

Note. Siever’s wormwood is widely distributed in the steppe and
forest-steppe zones of Siberia and in the southern regions of the forest
zone. It is a common plant of fallow lands of the chernozem steppes,
where it does not avoid even alkaline soils. At the same time, it is a
common weedy plant of crops, where it grows especially luxuriantly in
dry years; it is also ruderal, constantly found by roadsides, near
dwellings, at the edges of fields, etc. Under fallow conditions, it often
dominates the plant cover in the second and even third year, when it
appears to form almost a pure herb cover. Later, as the arable layer of
the soil becomes compact, this wormwood is replaced by rhizomatous
grasses. It is rarely found on old fallow lands. Siever’s wormwood,
normally an annual or a biennial weed, can be fully eradicated by
usual agronomic measures (fall plowing, early fallow). In the opinion
of animal husbandry specialists (Tr. Omsk. Inst. Moloch. Khoz., 1, II
1931-1933), this wormwood can be used also as a highly nutritive
fodder, particularly suitable for making silage. However, because of
the bitterness inherent in wormwoods, it is recommended that, as a
feed for milk cows, it be mixed with other fodders (I.V. Larin, 1956).

Economic Importance. Siever’s wormwood contains 0.02 to 0.4%
essential oil. According to G.V. Pigulevsky and G.P. Berezovsky (1949),
the constants of the oil are: D,,”°—0.9072; D,’?—0.9056, n,”°—1.4845;
acid number—1.04; ester number—38.23; alcohols—24%; ketones—
12%; phenols—12%. The oil contains o-pinene (highly recemated*),
cineole—13.8%, myrcene—10.89%; alcohols, in all probability, are
tertiary, the total quantity of them is 24%; of phenols 12%, of ketones
of unknown nature 12%, and of azulene-traces; sesquiterpenic carbo-
hydrates and alcohols have not been investigated in detail.

76. A. macrocephala Jacq. Cat. (1833) No. 2000; Bess. in Bull.
Soc. Nat. Mosc. IX (1836) 28; DC. Prodr. VI, 125; Hook. FI. Br. Ind.
III, 329; O. Fedtsch. Fl. Pam. 117; Fedtsch. Perech. Rast. Turk. IV,
204; Pampan. in Lavori Inst. bot. Univ. Cagliari, XL, 570; Krascheninn.
in Kryl. Fl. Zap. Sib. XI 2820; Grub. Konsp. Fl. Mong. 266.—A.
griffithiana Boiss. FI. or. III (1875) 376.—A. sieversiana var. pygmaea
Kryl. Fl. Alt. III (1904) 656.—A. akbaitalensis O. Fedtsch. Second
Addendum to Fl. Pam. (1905) No. 88.—A. krytoviana Steinb. (nom.)
in Prokhor. and Lebed. Dush. Rast. Alt. (1932) 15.—Pyrethrum
pamiricum O. Fedtsch. in Tr. Peterb. Bot. Sada, XXI (1903) 345.—
Exs.: GRF No. 3296.

*Optically highly inactive—General Editor.

493

Annual. Whole plant more or less covered with dense, white,
519 appressed hairs; root slender, vertical bearing solitary or numerous,
simple or sometimes branched (at base), 6-20(30) cm high stems. Lower
and middle cauline leaves petiolate, 1.5—4.0 cm long and 1-2 cm wide;
lamina broadly ovate to oblong-ovate, twice pinnately cut, with 2 pairs
of lobes, lobes ternately divided or pinnately cut, secondary lobes
oblong-linear or linear-spatulate, obtuse or round, 1-4 mm long; upper
leaves smaller, subsessile, simple pinnate, bracteal leaves entire, linear.
Capitula hemispherical, 4-10 mm in dia, on more or less long peduncles
in lax racemes in upper half of stem and branches. Outer involucral
bracts herbaceous, linear, densely hairy, inner bracts oblong-ovate or
elliptical, with narrow, herbaceous spine and wide whitish scarious
margin, subglabrous. Receptacle convex, hemispherical, hairy.
Peripheral florets many-rowed, pistillate, numerous (to 70), their
corollas narrowly tubular, enlarged toward base, punctate-glandular;
stigma lobes scarcely exserted from tube, linear; disk florets bisexual,
numerous (about 90), their corollas narrowly cup-shaped-conical,
glabrous, punctate-glandular; anthers on short filaments, oblong-
lanceolate, apical appendages of anthers long, acute, basal appendages
short, round; stigma lobes broadly linear, ciliate, after flowering
divergent. Achenes to 1.2 mm long, ovate-oblong, finely sulcate,
apically with flat, weakly bordered areola. Flowering August.

Gravelly, clayey and rubbly places at high-altitude or alpine zone
of mountains, sometimes descending considerably lower along
streams.—Western Siberia: Altai (Chuya “belki”*); Soviet Central Asia:
Tien-Shan, Pamiro-Alai Region. General distribution: Mongolia, Iran
Region (Afghanistan), Tibet, Himalayas. Described from Himalayas.
Type in Paris.

Economic Importance. According to Prokhorova and Lebedev
(1932), the recovery of essential oil is 0.4-0.5%. The oil contains
(M.A. Favorsky, 1935) l-a-pinene—9%, cineole—12.1%, camphors—
16.3%, azulene—6.5%; and 28.3%, apparently, of tertiary alcohol
(possibly borneol).

According to the data of M.I. Goryaev, K.K. Kruglykhina and
E.N. Sazhdarova (1959), the presence of alkaloids in the plant has
been established.

77. A. kulbadica Boiss. and Buhse in Nouv. Mém. Soc. Nat.
Mosc. XII (1860) 120; Boiss. Fl. or. III, 373.

Annual, biennial. Plant grayish from dense, appressed, short hairs;
root vertical. Stem 80-120 cm high, herbaceous, erect, ribbed, leafy,

*Snow-covered flattened mountain summits in Siberia—General Editor.

520

494

branched. Leaves, especially beneath, with distinct veins; lower cauline
leaves withering before flowering, petiolate, to 4 cm long, lamina
broadly ovate, twice pinnately dissected, with 2-3 pairs of lobes, lobules
3-8 mm long and 1-2 mm wide, undivided or with 1-3 short teeth,
oblong-linear or lanceolate, obtuse or short-acuminate; middle and upper
cauline leaves sessile or subsessile, pinnatisect or ternate, bracteal
leaves simple, oblong-linear. Capitula globose, 3-4 mm in dia,
pedunculate, drooping, usually remote, on long divergent branches, in

_ wide, pyramidal, paniculate inflorescence. Involucral bracts punctate-

glandular, brownish, scarious along margin, irregularly incised; outer
bracts broadly cuneate, densely hairy, inner bracts larger, oblong-ovate
or oblong, subglabrous. Receptacle with hairs. Peripheral florets
pistillate (7-8), their corollas narrowly tubular, punctate-glandular,
weakly hairy or glabrous; stigma lobes scarcely exserted from tube,
linear, but most often disk florets bisexual, 32-40, their corollas conical,
glabrous; anthers lanceolate-elliptical, apical appendages of anthers
large, subacute, basal appendages short, obtuse; style shorter than tube;
stigma lobes short, linear, short-ciliate, after anthesis divergent.
Achenes to 2 mm long, oblong-ovoid or almost pyriform, somewhat
flat, sulcate, apically with a plano-convex areola.

Mountain deserts and foothills along banks of streams, in ravines
on slopes.—Soviet Central Asia: mountainous Turkmenia (western
Kopetdag). General distribution: Northern Iran. Described from Caspian
Coast. Type in Geneva; cotype in Leningrad.

78. A. samoiedorum Pampan. in Lavori Inst. bot. Univ. Cagliari,
XXXV (1938) 2.—A. anethifolia Fisch. var. macrocephala F. Schmidt.
in Mém. Acad. Sc. Pétersb. VII, XVIII, 1 (1872) 107.—Exs.: GRF
No. 3297.

Annual. Plant herbaceous, more or less densely hairy; root slender,
vertical. Stem 6—12—18(28) cm high, brownish-violet, branched from
base, with more or less numerous, upright, arcuate branches. All leaves
petiolate, simple pinnate or in part ternate, terminal lobe narrowly
oblanceolate, round or short acuminate. Capitula at apices of stem and
branches, solitary or many, in simple recemes, sessile or long
pedunculate, divergent, somewhat plano-convex, round in cross section,
6-10 mm in dia. Involucral bracts oval, outer bracts densely hairy
along narrow, linear, raised, herbaceous spine and with wide, irregularly
incised, scarious, margin; inner bracts brownish completely scarious,
weakly hairy. Receptacle convex, hemispherical, finely tuberculate,
glabrous or hairy. Peripheral florets pistillate, 14-15; their corollas
narrowly tubular, slightly enlarged toward base, glabrous, punctate-
glandular; stigma lobes exserted from tube, narrowly linear, obtuse,

—

495

straight; disk florets bisexual, rather numerous (to 170); central florets
sometimes not fully mature or undeveloped, their corollas conical,
glabrous, punctate-glandular; anthers oblong-linear, apical appendages
of anthers long, acute, basal appendages not developed; stigma lobes
exserted from tube at maturity of achenes, linear, cilia at apex, curved.
Achenes to 1.2 mm long, oblong-prismatic or fusiform, angular, sulcate.
Flowering August.

Riverbanks, sands and gravel beds in tundra zone, partly in forest-
tundra.—Arctic: Arctic Siberia (low-lying areas of Yenisei,
Turukhansk), Anadyr, Chukotka (upper reaches of Anadyr River).
Endemic. Described from Arctic Siberia. Type in Florence.

79. A. succulenta Ldb. Fl. Alt. IV (1838) 81; Bess. in Bull. Soc.
Nat. Mosc. IX, 71; DC. Prodr. VI, 120; Ldb. FI. ross. II, 593; Fedtsch.
Perech. Rast. Turk. IV, 201; Krascheninn. in Fl. Zap. Sib. XI, 2822.—
Ic.: Ldb. Ic. pl. Fl. Ross. V, tab. 474.

Annual or biennial. Plant covered with variously dense, bifid and
simple hairs, as well as capitate glandular hairs and simple glands.
Root slender, fusiform, vertical. Stems single or few, 8 to 50-60 cm
high, slender or thicker, erect or ascending, ribbed, simple or strongly
branched. Leaves succulent, initially hairy, later glabrescent, strongly
variable in size and degree of cutting; basal and lower cauline leaves
withering early, 7-15 cm long, twice or thrice pinnately incised, lobes
fewer, obliquely antrorse, lobules oblong, obtuse; middle and upper
cauline leaves less large (2-4 cm long), twice pinnately incised, with
narrower secondary lobes; upper bracteal leaves simple, linear,
sometimes with auricles. Inflorescence in small specimens simple,
racemose, with fewer capitula, paniculate in larger plants and with
many capitula. Capitula hemispherical, 4-5 mm in dia, pedunculate,
somewhat remote, drooping. Involucral bracts densely hairy and
punctate-glandular; outer bracts small, lanceolate, raised along spine,
with narrowly scarious margin; inner bracts larger, oval, widely scarious.
Receptacle convex, tuberculate, glabrous. Peripheral florets pistillate,
16; their corollas narrowly tubular, punctate-glandular; stigma lobes
exserted from corolla tube, linear, obtuse, divergent or recurved; disk
florets numerous (to 75); their corollas conical, glabrous; anthers linear,
apical appendages of anthers long, acute, basal appendages short, round;
stigma lobes at maturity of achenes exserted from tube, linear, obtuse,
arcuate. Achenes to 1.2 mm long, oblong, sulcate, apically with round,
areola with border. Flowering August.

Saline [marshy] soils, [marshy] saline meadows, sometimes a weed
in irrigated areas or trash dumps, in steppe and desert zones.— Western
Siberia: Irtysh (southern Semipalatinsk); Soviet Central Asia:

522

496

Aralo-Caspian Region (Kyzyl-Orda, Aral Region), Kara-Kum (lower
reaches of Amu-Darya). Endemic. Described from Kazakhstan. Type —
in Leningrad.

Economic Importance. According to the data of M.I. Goryaev,
R.N. Sazonova and N.I. Bekanina (1951), the yield of essential oil was
0.52% from plants collected in the Kzyl-Orda District. The liquid was
viscous, bright yellow, with a strong bitter taste. The constants:
Dedl=- OPaliy D* —1.4772; acid number—4.6; ester number—37.56;
saponification number—42.16. It does not dissolve in 70% alcohol,
but in 80% alcohol it dissolves in the ratio of 1:1 and completely in
90% alcohol.

80. A. jacutica Drob. in Tr. Bot. Muz. Akad. Nauk SSSR, XII
(1914) 108.—A. sieversiana var. septentrionalis Pamp. in Lavori Inst.
bot. Univ. Cagliari, XXXV (1938) 3.—Ic.: in Tr. Bot. Muz. Akad.
Nauk SSSR, XII, 109, Fig. 2 and Plate XV.—Exs.: GRF No. 3298.

Annual or biennial. Whole plant grayish from dense white,
appressed hairs. Root slender, vertical. Stems single or few, 25-40 cm
high, erect or basally ascending, almost circular, sometimes strongly
branched almost from base. Lower leaves long-petiolate, 2.0-4.5 cm
long, lamina round or broadly ovate, twice or thrice pinnately incised,
lobes 2 pairs, pinnati-, twice pinnately incised or ternate, terminal
lobule narrowly linear, 0.5-1.0 mm wide, middle cauline leaves
subsessile, upper ones much less divided, palmatipartite or ternate.
Capitula hemispherical, 6-8 mm in dia, pedunculate, inclined or
drooping, in broad, lax, paniculate inflorescence. Involucral bracts
prolate, with narrow green spine and wide, whitish, scarious margin;
outer bracts densely covered with white hairs, inner ones glabrous or
subglabrous. Receptacle convex, glabrous or hairy. Peripheral florets
pistillate (to 21); their corollas narrowly tubular, punctate-glandular;
stigma lobes linear, exserted from tube, linear, obtuse, divergent; disk
florets bisexual numerous (to 100), of which central florets often
undeveloped; their corollas conical, punctate-glandular, anthers oblong-
linear, on short filaments, apical appendages of anthers long, almost
subulate, basal appendages very short, obtuse, stigma lobes linear,
short-ciliate, exserted from corolla at maturity of achenes, arcuate.
Achenes | mm long, oblong-ovoid or pyriform, somewhat flat, sulcate,
with a round areola at apex. Flowering August.

Peat bogs, lake shores, old fields and weedy places.—Eastern
Siberia: Angara-Sayans, Dauria, Lena-Kolyma. Endemic. Described
from Yakutia. Type in Leningrad.

523

497

Series 10. Anethifoliae Poljak.—Plant usually densely hairy in
beginning, later glabrous; leaves twice or thrice pinnately incised into
filiform secondary lobes; capitula 2-4 mm in dia; receptacle with floral
hairs or glabrous.

81. A. anethifolia Web. in Stechm. Artem. (1775) XXIX, No.
XXV; Willd. Sp. pl. III, 1828; DC. Prodr. VI, 126; Ldb. Fl. Ross. II,
600; Turcz. Fl. baic.-dahur. II, 2, 207; Maxim. in Bull. Acad. Sc.
Pétersb. VIII, 537; Kryl. Fl. Alt. III, 657; Kom. Fl. Manchzh. III, 678;
Pampan. in Nouv. Giron. Bot. Ital. n. s. XXXIV, 646.—A. anethifolia
stelleriana, var. erectiflora and var. multicaulis DC. Prodr. VI (1837)
126.—A. multicaulis Ldb. Fl. Alt. [V (1833) 60; Krascheninn. in Kryl.
Fl. Zap. Sib. XI, 2821.—Absinthium divaricatum Fisch. ex Bess. in
Bull. Soc. Nat. Mosc. VIII (1829) 263.—A. 6. multicaulis Bess. in
Bull. Soc. Nat. Mosc. IX (1836) 31.—A. korotkyi Krasch. in Spisok
Rast. Gerb. Fl. SSSR, X (1936) 73.—Ic.: Gmel. Fl. Sib. II, tab. LIV.—
Exs.: GRF Nos. 3171a, b, 3151.

Annual or biennial. Root vertical, sometimes thicker. Stems solitary
or numerous, 20-35 (45) cm or 3-10 cm high (var. korotkyi (Krasch.)
Poljak.), erect or decumbent, ribbed, glabrous, brownish-violet, strongly
branched, sometimes almost from base. Leaves whitish-woolly, later
glabrous; basal leaves withering early, long-petiolate, to 3.0-4.5 cm
long, twice or thrice pinnately incised, terminal lobes linear-filiform,
subobtuse, 3-8 mm long; middle cauline leaves short-petiolate or sessile,
less divided; uppermost leaves bracteal, simple, linear-filiform. Capitula
broadly campanulate, 2-4 mm in dia, more or less on long, filifom
peduncles, drooping, somewhat remote, in lax paniculate inflorescence.
Involucral bracts whitish-hairy or subglabrous; outer bracts small,
lanceolate, herbaceous, inner bracts oval, with wide scarious margin.
Receptacle convex, hemispherical, with floral hairs or glabrous (var.
korotkyi (Krasch.) Poljak.). Peripheral florets pistillate few (3-6), their
corollas narrowly tubular, punctate glandular; stigma lobes narrowly
linear, obtuse, straight or slightly divergent; disk florets bisexual (to
18), their corollas conical, punctate-glandular, usually purple-pink,
glabrous, less often yellow (f. multicaulis DC.); anthers linear, apical
appendages of anthers long, subulate, basal appendages of anthers short,
subobtuse; stigma lobes linear, dilated, ciliate, weakly divergent.
Achenes to 1.5 mm long, oblong-conical, somewhat flat, brown, apically
with a round areola, strongly bordered. Flowering August. (Plate XX VII,
Fig. 2).

Valleys of rivers and lakes in salt marshes, alkali soils, sands in
steppe or desert-steppe zones and in mountains.—Western Siberia:
Altai (Chuya Steppe); Eastern Siberia: Angara-Sayans, Dauria. General

524

498

distribution: Mongolia. Described from Steller’s specimen from the
salt marshes near Lake Baikal. Type in Leningrad.

Note. A. multicaulis Ldb. is a variety of A. anethifolia Web.,
which is distinguished by the color of the corolla of the disk florets.
Likewise, recognition of A. korotkyi Krasch. is not justified, since it is
a form distinguished only by its short height and the absence of hairs
on the receptacle, i.e., by characters of no taxonomic significance.

Section 5. Artanacetum (Rzazade) Poljak. emend. comb. nov.—
Artanacetum Rzazade in Izv. Akad. Nauk. Azerb. SSR (1957) 3, pro
gen.—Capitula in fasciculate corymb at apices of branches; involucre
many-rowed, imbricate, outer involucral bracts small, round, inner bracts
considerably larger, oval; receptacle narrow, cylindrical-conical,
glabrous; achenes oblong-prismatic.

82. A. incana (L.) Druce in Rept. Bot. exch. Cl. Brit. Isles, III
(1914) 414.—Tanacetum incanum L. Sp. pl. (1753) 844.—T. orientale
Willd. Sp. pl. II (1804) 1812.—T. canescens DC. Prodr. VI (1835)
129.—Artemisia orientalis Willd. loc. cit.; Grossh. Fl. Kavk. III, 143.—
A. fasciculata MV. FI. taur.-cauc. II (1808) 293; Bess. in Nouv. Mém.
Soc. Nat. Mosc. III, 62; DC. Prodr. VI, 116; Ldb. Fl. Ross. II, 587;
Boiss. Fl. or. III, 368; Grossh. op. cit. III, 142.—A. fasciculata var.
iberica and var. armeniaca Bess. in Bull. Soc. Nat. Mosc. VIII (1835)
63, 64.—Artanacetum fasciculatum (MB) Rzazade in Izv. Akad. Nauk.
Azerb. SSR (1957) 3.—Exs.: GRF No. 3168a, b; Herb. Fl. Cauc. No.
497.

Perennial. Whole plant whitish from dense, silky appressed hairs.
Root thick, woody, with perennial, woody, more or less branched,
sterile shoots covered with brownish gray bark. Flowering shoots many,
erect, (20)25-40 cm high, finely branched in upper part, with long,
straight, divergent branches. Lower and middle leaves petiolate, 1.0—
2.5 cm long and 0.8—1.7 cm wide, their laminas broadly round, pinnately
or ternately palmate lobes petiolulate, pinnate, palmate, or ternate,
terminal lobules lanceolate or linear-lanceolate, scarcely acuminate,
sometimes round, 2-6 mm long; upper leaves sessile, more often
palmately cut or ternate, bracteal leaves mostly simple, lanceolate-
linear. Capitula ovate-conical, to 4 mm long, pedunculate, upright,
partly drooping, in dense fasciculate corymbs, at apices of more or
less divergent branches forming racemose-paniculate inflorescence.
Involucre conical, many-rowed imbricate; outer involucral bracts small,
round; inner considerably larger, oval, hairy along spine, glabrous in
remaining part, smooth, with wide, unevenly scarious margin.
Receptacle convex, narrow, cylindrical-conical, glabrous. Peripheral

525

499

florets pistillate, few; their corollas narrowly tubular, enlarged at base,
with 2 teeth, punctate-glandular and with sparse hairs; stigma lobes
projecting, linear, truncate, divergent; disk florets 9-12, bisexual; their
corollas narrowly conical, with 5 teeth, with white, long, erect hairs on
teeth, glabrous and punctate-glandular in remaining part; anthers
exceeding tube, linear-lanceolate, apical and basal appendages of anthers
acute; stigma lobes linear, arcuate after flowering, with dense straight
cilia at apex. Achenes up to 1.5 mm long and 0.4—0.5 mm thick, dark
brown, oblong-prismatic, angular, with flat, round apex, bordered on
one side. Flowering August.

Stony slopes of mountains.—Caucasus: Eastern and Southern
Transcaucasia, Dagestan. Endemic. Described from Georgia (Kura
River). Type in Leningrad.

Subgenus 2. Dracunculus Bess. sect. in Bull. Soc. Nat. Mosc.
VIII (1835) 16.—Oligosporus Cass. in Bull. philom (1817) 33 and
Dict. Nat. XXXVI (1826) 25.—Capitula smaller, ovate or globose,
sessile or pedunculate, in spicate, racemose or paniculate, less often in
dense capitate inflorescence. Involucral bracts in 2-3 rows, oval, round,
or oblong-lanceolate, herbaceous along spine, glabrous or hairy, with
more or less wide scarious margin, usually not incised-toothed,
uniformly long but outer involucral bracts smaller than inner bracts.
Receptacle flat or somewhat convex. Peripheral florets pistillate, always
fertile; their corollas tubular or narrowly conical, often slightly enlarged
at base, with 2-3 teeth, glabrous, stigma lobes 2, less often 3, narrowly
linear, subacute or obtuse; pollen-receptive area as continuous strip
bordering margin of lobe. Achenes small, pyriform, ovoid or oblong,
somewhat flat, narrowed at base, very finely ribbed brown; pappus
absent; disk florets staminate but with rudiment of pistil, few or rather
numerous; their corollas usually narrowly campanulate, with 5 acute,
straight, yellow or pinkish teeth, glabrous or hairy in upper part; anthers
linear, apical appendages of anthers obtuse or acute, basal appendages
smaller, subobtuse, weakly developed, “antheropodia” well-developed,
usually convex, their pistils abortive as style (lacking ovary), with
undivided funnel- or goblet-shaped, densely ciliate stigma, less often
stigma with 2 short, erect, basally connate lobes.

Annual and shrub-like or herbaceous plants with simple, less often
branched (stellate) hairs or glabrous; leaves simple, linear and narrowly
lanceolate or pinnately cut, twice pinnately cut or pinnately divided.
The subgenus includes about 80 species distributed in the extratropical
region of Asia to the extreme north, as well as in Europe, northern
Africa, and North America.

526

500

D+

+

10.

All or almost all leaves simple, linear-lanceolate or narrowly
lanceolate, less often some leaves with 2—5 lobes or 3 lobed at

APEX! wish. sioniel) Haile ae ned hh eae cepetinl borer aioe 2:
Lower and middle cauline leaves simple to twice pinnately in-
cised on pinnatifidi: iJ: isha dean bench aie 8.

Leaves gray from more or less dense pubescence of bifid hairs ..

vais AA Ss Wl teeth ed 9 apt eaten, tema eh sold 87. A. glauca Pall.
Leaves mainly yellowish-green, glabrous, less often pubescent with
simple;on bifidchairs; but notyeray) ted j.ctesscccisdeel octets... eee 3.

Shrub-like perennial, 60-90 cm high, strongly woody at base;

flowering shoots strongly branched. Plants of sandy deserts of
Soviet: CentialbAsia.:a1J.,\oiextien test ath aeons eee 4.

Perennials with herbaceous stems, sometimes weakly woody only
at base, more weakly branched. Mainly steppe, meadow, or
halophylictplants::...ic28...22041..oeheieend..f. ass 5;

All leaves simple, lanceolate-linear, with short appressed hairs;
involucres denselyshainyess ety a 88. A. dimoana M. Pop.

Leaves mainly glabrous, narrowly linear, sometimes with 2—5 lobes
in upper part, involucres glabrous ............ 89. A. kelleri Krasch.
All or most leaves more or less deeply 3-lobed at apex ............
dateitheeied Jes eb eey.. Dieta ct 85. A. dracunculiformis Krasch.
All leaves entire, very rarely some with short teeth ................04 6.

Leaves usually rather densely hairy beneath with simple appressed
hairs;; Plants of Caucasusiwonihte Rahahkkestiane. Bed eee
heh Baas, ale a 86. A. daghestanica Krasch. and Por.
Leaves glabrous or more weakly hairy ..........::csscsesereeeeeseonaeeenes 7.

Stem brownish or violet-brownish; inflorescence more or less
narrow, idense panicle. <st.:.2ce8.c8s be: 84. A. pamirica Winkl.
Stem stramineous; inflorescence more or less spreading or narrow
pyramidalupanicles .tacasteags alse 83. A. dracunculus L.
Annuals or biennials, mostly with solitary stem and more slender
vertical root..s.2:..msaiitidastand inavken enone are eee 9,
Perennials\or:semishnubs, ivsicarwceisccheiien nhts epee sae ae 12.
Plants 5-20 cm high; leaves small, mostly 0.6-1.8 cm long;
peripheral florets pistillate, 15-18, central florets 5—6 .................
snvoegiieghs tue. <tgeeeenenl, binary! salihihgat | deans 116. A. demissa Krasch.

shulibia eolesildlshvg Ghdveicbnice « Raowaiceahs erates ob plese est paed, dinh eta a Seapine 10.
Lobes of middle and upper cauline leaves narrowly linear; stem
reddish-browmisstiatz,ts.n asada his 115. A. scoparia W. and K.
All leaves except lowermost divided into filiform-linear lobes;
stem stramineous;}or brownish: s::...cesssseivgzlss shoes ¥oocteaee-saeeese LA.

We
15(12).
at
16.
L
17.

-Ts

18.

19.

20.

al.

501

Plants glabrous; lobes of middle cauline leaves 8-12 mm long. ...
WRU NIG. UCD NAAR, Bias EI a 104. A. capillaris Thunb.
Plants appressed-hairy; lobes of leaves 3-5 mm long .........
CWA alannah 115. A. scoparia var. scoparioides Poljak.
Leaves of sterile shoots or all, or at least some lower cauline,
leaves oblong-cuneate, toothed or lobed ......... cee eeseeeseeeees 13.
Lower leaves pinnately cut, twice or almost thrice pinnately incised
oe pinnatelydividedsy.c.ytheon. 20 s20Gel gehen. cAingi..2... Ad 5.
Inflorescence spreading panicle, with straight, long more or less
strongly divergent branches ................0 100. A. japonica Thunb.
Panicle narrow, dense, with short branches appressed to
inflorescence axis or branches weakly divergent .................... 14.
Leaves of sterile shoots and partly lowermost cauline leaves
obcuneate, toothed, but middle and partly upper cauline leaves
pinnately incised; plant glabrous ..... 102. A. desertorum Spreng.
Leaves mostly obcuneate, toothed or toothed-lobate; plant more

or less pubescent in upper part. ............ 101. A. littoricola Kitam.
Plant densely caespitose, pulvinate; stem 1-3 cm high; capitula in
dense capitate inflorescence. ............ 112. A. henriettae Krasch.
Plants taller; capitula in other type of inflorescence ............. 16.
Semishrubs (shrub-like perennials) ............:cccccssesseesseeseeseeeeee LT:
Herbaccous.perenniiaks: d sinsssg tat. LO dened NAG Aiectens 23.
Involucres always, but leaves and stem mostly densely appressed
ramen... Ceaser aaa. de Ria wwesins 18.
Plants glabrous or more weakly hairy only in beginning of
vegetative growth; involucres glabrous ............scscsseeressseeereseees 19,

Lower cauline leaves 5-8 cm long, divided into narrowly linear,
0.7-2.7 cm long and 1-2 mm wide lobes; capitula ovate, sessile,
crowded on branches .............:.000 90. A. trautvetteriana Bess.
Lower leaves 3—5 cm long, divided into narrowly linear, 5—10(15)
mm long and 1 mm wide lobes; capitula subglobose, pedunculate,
Pemiote aa satan anid 93. A. quinqueloba Trautv.
Capitula subglobose, 1.5—-2.0 mm wide and as long, on almost
horizontal divergent branches in broad, subglobose, lax
inflorescence; plants more or less ephemeral, drying out at
Serena OF July. sic. .ccsereseascecccesnsesss 97. A. songarica Schrenk.
Capitula larger, ovoid; inflorescence of other types ............... 20.
Inflorescence narrow, dense panicle or raceme ...............00004 ai.
Capitula in more or less lax racemose-paniculate inflorescence

Capitula 2.5—3.5 mm long, in racemose inflorescence; lower leaves
2.0-3.5 cm long, pinnately cut; plant glabrous or sometimes with
occasional stellate hairs .............:seeee 91. A. salsoloides Willd.

502

528 22.

23(16).

28.

29.

30.

31!

32(23).

+
33:

“b

Capitula 4-5 mm long, in paniculate inflorescence; lower leaves
3-6 cm long, once or twice pinnately cut; plant with simple hairs

or glabrous), ..2svas!..20..22 92. A. halodendron Turcz. ex Bess.
Lower cauline leaves 2—5(6) cm long, twice pinnately cut, their
lobes: 5-215, mmulongys..tin.10. annie alies 95. A. arenaria DC.
Leaves 5-8 cm long, pinnately cut, their lobes to 3-4 cm long ...
miauretien ceabenlt.sacacen tg me earth tie. il ass 95. A. albicerata Krasch.
Capitula subglobose or broadly Ovate ...........ceeeesseeeseceeseeeeeees 24.
Capitula ovate or narrowly OVALE ........ cee eeeecesseeeeneeeeereeeeneeeees B2}
eaves usually glabrous ‘c:. 2odamaatl inet tiga ke 25.
Leavesimore or lessidensely, hairy!.®:...2ccawh.eklagee.. 27.
Lobes narrowly linear or filiform-linear, to 7-16 mm long ...........
sluea.ceoniubek bltene, shee 103. A. macilenta (Maxim.) Krasch.
Lobes linear or lanceolate-linear, mostly to 7 mm long .......... 26.
Capitula 4—5 mm long ....... cee eeeeeeeeeeeeee 113. A. limosa Koidz.
Capitula!2=3mn long@=ciue:..2teen ios. clive eee. Be.
popes: eck by. i 105. A. commutata Bess. var. dolosa Krasch.
Capitula in narrow, racemose, partly racemose-paniculate
inflorescénde if.4.hi...c0is..2, oie, Bie ee. ee 28.
Capitulaanipaniculateanflorescenct: cadet eet .. .cbe 30.
Capitula small, 2.5-3.0 mm in dia ........ 114. A. pyenorhiza Ldb.
Capitula.3=4immnin. dia:..c20.cldeneeet eek 29.

Leaves hairy, twice or almost thrice pinnately incised; involucral
bracts and central florets usually hairy ...... 111. A. borealis Pall.
Leaves glabrous or hairy, pinnately cut or almost twice pinnately
incised; florets and involucre usually glabrous .................00008
MATEUOES, AWS 134. A. pycnorhiza Ldb. var. depauperata Krasch.
Plant green, moderately hairy; capitula small, to 2 mm in dia
deedientspank hs 8 105. A. commutata Bess. var. pubescens Ldb.
Plant whitish from dense semidivergent hairs; capitula larger
Qala igen el aeeodae Labeler hae, wae, Sa 31.
Lower cauline leaves twice pinnately incised; capitula 3-4 mm in
diagtah ne tune cnbbertern. 00.08, Laos 109. A. ledebouriana Bess.
Leaves spatulate-pinnately divided; lobes undivided or partly
pinnately divided; capitula 2-3 mm in dia ........cccsssssssessesseeseeeeeeeeeees

Leaves palmately divided into 4—7 parts .........:cssseseeseeseeeeeeees 33.
Leaves. more:compotndlac. naieehs. aceite eevee tes aeen se 34.
Stems 34-40 cm high, branched; lower cauline leaves 3-5 cm
long; inflorescence narrow panicle ........... 98. A. lipskyi Poljak.
Stems 5—12(15) cm high, usually simple or with short branches;
leaves 1-3 cm long; inflorescence narrow, racemose-paniculate.
DNR, Bask Qs ssivvniiananbboysecsannin MRE 99. A. kuschakewiczi Winkl.

529

34.

36.

37(35).

a
38.

39%

cot

503

Capitula in simple racemes in upper part of stem, on short branches
STOW SAMI ci Rog heid. BINT, 96. A. saposhnikovii Krasch. ex Poljak.
inflorimienee-pamculnte, LY. ..26 ae, fk Wate li RUL.. 35.
Capitula 3.0-4.5 mm long, in dense narrow panicle; plants glabrous
OP Sabplabrass Gat ninan..dcnnanle. LANs 3.0... ial Ve 36.
Capitula 2.0—2.5(3.0) mm long, in more or less spreading, less
often somewhat narrow panicle; plants mostly pubescent,
SOM@pIiee Nn alMMSs~>.05.. O58 23.5.4 Be Ae. Ah dle 37.
Capitula 4-5 mm long and 3-4 mm in dia, with flat broad base,
upright, with numerous florets (to 40 and more) ...........ccceeseeeees
DULY, ORB OLR LON, LW LLY 108. A. bargusinensis Spreng.
Capitula 3.0-3.5 mm long and 1.5-3.0 mm in dia, with broadly
cuneate base, divergent or drooping; florets 23-30 ......0....

PONE RHE RARE Le... PRI ADR 105. A. commutata Bess.
ea eS SommraOIS A... AW, ess. GL Ld, Deh ae... 38.
Leaves glabrous or more weakly hairy .............cceseesseeeseeeneees 39.

Lower cauline leaves once or almost twice pinnately cut, lobes
linear-lanceolate, 0.3—3.0 cm long; capitula on primary branches
in short racemes or glomerules ........... 107. A. tomentella Trautv.
Leaves pinnately cut, lobes narrowly linear, often 7-15 mm long;
capitula in spikes on terminal branches .......... cies eeeseeeeeeeees
pte home Mat ce 106. A. campestris L. var. sericophylla Rupr.
Lower cauline leaves twice, less often thrice pinnately incised,
their lobes 3—10(20) mm long, linear; capitula ovoid, 1.5—3.0 mm
in dia; outer involucral bracts glabrous or weakly hairy ......
TOW. AERO SALA LD al 106. A. campestris L.
Lower cauline leaves twice pinnately incised, their lobes narrowly
linear, 7-15 mm long; capitula narrowly ovoid, up to 1.5 mm in
dia; outer involucral bracts glandular or hairy ...........:csceseeeee
ORI ARG. 106. A. campestris L. var. araratica Novopokr.
Lower cauline leaves twice pinnately incised, their lobes to 10
mm long, linear; capitula ovoid or globose, 2-3 mm in dia; outer
involucral bracts glandular or hairy .............ccssssssssseessseereseseeeees
See 8) RS Se 106. A. campestris L. var. sosnovskyi Novopokr.

Series 1. Dracunculi Poljak.—Perennials with simple or branched
hairs or glabrous; leaves simple, linear or lanceolate-linear.

83. A. dracunculus L. Sp. pl. (1753) 849; Bess. in Bull. Soc. Nat.
Mosc. VIII, 54; DC. Prodr. VI, 97; Ldb. Fl. Ross. I, 563; Turcz. Fl.
baic.-dahur. II, 2, 49; Maxim. in Bull. Acad. Sc. Pétersb. VIII, 524;
Kryl. Fl. Alt. III, 633; Kom. Fl. Manchzh. III, 658; Fedtsch. Perech.
Rast. Turk. IV, 190; Rydb. North Am. FI. 34, part 3, 251; Pampan. in

530

504

Nuov. Giorn. Bot. Ital. n. s. XXXIV (1927) 653; Hegi, Ill. Fl. VI, 2,
635; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1035; Krascheninn.
in Fl. Yugo-Vost. Evrop. Ch. SSSR, VI, 350; Krascheninn. in Kry]. FI.
Zap. Sib., XI, 2768; Poljak. in Majevski, Fl. 583.—A. inodora Willd.
Enum. horti berol. II (1809) 864.—A. redowskyi Ldb. in Mém. Acad.
Sc. Pétersb. V (1815) 574, non MB.—A. desertorum var. macrocephala
Franch. Pl. Turk. (1883) 88, non Spr.—Oligosporus condimentarius
Cass. in Dict. Sc. Nat. XXXVI (1826) 26.—Exs.: GRF Nos. 3178a, b,
3179.

Perennial. Rhizome woody, 0.5—1.5 cm thick, sparsely covered
with rootlets and sometimes with well developed stolons; whole plant
glabrous, smooth, green, less often hairy when young. Stems erect,
solitary or few, 20-150 cm high, ribbed, usually branched, lower
branches sterile. Leaves undivided, linear-lanceolate or almost linear,
1.5-8.0 cm long and 1-10(14) mm wide; lowermost cauline leaves
sometimes with 3 lobes at apex. Capitula numerous, globose, drooping,
(2)2.5-4.0 mm in dia, at apices of stem and branches in racemes,
forming paniculate inflorescence. Involucral bracts smooth, outer bracts
oblong or sublanceolate, inner bracts round-oval, with wide scarious
margin. Peripheral florets pistillate, usually 7; their corollas tubular,
expanded toward base; stigma lobes narrowly linear, subacute, exserted
from corolla tube, divergent; disk florets staminate, 11-14, their corollas
conical, with 5 teeth; anthers linear, apical appendages of anthers
obtuse-angled, subacute, basal appendages shorter, subobtuse; stigma
of abortive pistil not lobed, funnel-shaped at apex. Achenes small, 0.6
mm long, somewhat flat, ovoid, finely sulcate, brown. Flowering July
to August.

Alkaline meadows, near birch groves, river terraces of meadows,
steep valley slopes and old fallow lands, forest-steppe and steppe zones,
as well as meadow slopes of low mountains and mountains.—Euro-
pean part: Baltic Region, Upper Dnieper, Upper-Volga, Volga-Kama,
Middle Dnieper, Volga-Don, Trans-Volga, Black Sea Region, Crimea,

~ Lower Don, Lower Volga; Soviet Central Asia: Aralo-Caspian Region

533

(northern part), Lake Balkhash Region, Dzhungaria-Tarbagatai, Tien
Shan, Pamiro-Alai Region; Western Siberia: Upper Tobol, Irtysh, Altai,
Angara-Sayans, Dauria, Zeya-Bureya, Lena-Kolyma. General distribu-
tion: Central Europe, Balkans-Asia Minor, Mongolia, northern China,
North America. Described from Siberia. Type in London.

Note. A. dracunculus L. varies within its range, which provides a
basis to distinguish:

1) Var. pratorum Krasch. (Spisok Rast. Gerb. Fl. SSSR, X, No.
3179)—distinguished by a stem up to 2 m high, long and lax inflores-
cence, and leaves that are partly divided into 2 or 3 lobes; found in
Kazakhstan steppes.

505

:

4

Plate XXV.
1—Artemisia maximovicziana Krasch. ex Poljak. habit; 2—A. medioxima

531

Krasch. ex Poljak., habit.

506

2) Var. turkestanica Krasch.—distinguished by rather tall stems,
large lanceolate leaves, and capitula up to 3-4 mm [wide]; distributed
in Tien Shan, Pamiro-Alai.

3) Var. pilosa Krasch. (Fl. Yugo-Vost. Evrop. Ch. SSSR, VI. No.
1956)—distinguished by more or less densely hairy leaves and stems
at the beginning of the vegetative growth; distributed in the Trans-
Volga steppes.

4) Var. humilis Kryl. (Fl. Alt. III, p. 633)—distinguished by short
plants (20-30 cm high) and a narrowly paniculate or almost racemose
inflorescence; distributed in the West Siberian steppes.

5) Var. redovskyi Ldb.—distinguished by stramineous stems,
globose, 2-3 mm wide, drooping capitula on long peduncles, forming
a rather lax paniculate inflorescence; distributed in eastern Siberia and
Far East.

Economic Importance. According to the data of M.I. Goryaev
and R.N. Sazonov (1959), the recovery of essential oil is 0.5%. The
oil contains aldehydes and a small quantity of ketones; bound alcohols—
38.8%.

The above-ground parts of the plant contain the essential oil; the
yield from plants collected near Saratov reaches 0.54% (of dry weight)
and 0.17% (of fresh herb). According to the work of L. Kazakevich
and O. Sobolevskaya (1928), the oil has the following constants: D°—
0.9408; a + 29.35°; n,?°—1.4712; acid number 0.37; ester number 8.39;
ester number after acetylation 29.96. It is insoluble in 70% alcohol,
but dissolves in 80% alcohol in the ratio 1:0.76, and completely in
90% alcohol.

Tadzhik oil, according to the data of N.P. Kiryalov and D.P.
Snegirev (1936), contains 84-85% d-sabinene, 7—8% resin, precipitat-
ing crystals at melting point of 48°, and 7% of a high melting part
consisting of esters and, possibly, carbohydrates. According to the
data of I.P. Tsukervanik and N. Mirkina (1935), the yield of oil was
0.06-0.3%; the oil contained up to 65% sabinene and 10% myrcene.
According to foreign sources (cited in Goryaev, 1956), the oil contains
up to 60-70% methylchavicol (estragol), 15—20% myrcene and ocimene,
phellandrene, methoxycinnamic, anisic, and acetic aldehydes. The data
of M.I. Goryaev, (cf. Efirnye masla [Essential Oils], Fl. SSSR, 1952,
pp. 40, 41) erroneously attributed to this species, in fact, relate to
Artemisia schrenkiana Ldb. Probably, the data of V.A. Vyshensky
(1935) also relate to some other plant, as A. dracunculus does not
grow in Turkmenia.

Traces of alkaloides have also been found in the leaves, stems and
roots (Bankovsky, Zarubina, and Sergeeva, 1947).

534

507

The leaves and stems are used for pickling cucumbers and cab-
bage, and are used in folk medicine as an antiscorbutic and diuretic.
Often it is a noxious weed. I.V. Larin (1956) suggests the following
control measures: two mowings or pruning with a spade at a depth of
10 cm. The mowed plants can be used for making silage or, in extreme
cases, as fuel. The fodder quality of the plant is not high.

84. A. pamirica Winkl. in Tr. Peterb. Bot. Sada, XI, 10 (1890)
329; O. Fedtsch. Fl. Pam. 114; Fedtsch. Perech. Rast. Turk. IV, 191;
Pampan. in Nuov. Giron. Bot. Ital. n. s. XXXIV, 684.

Perennial. Plant more or less pubescent with forked hairs, later
only in upper part or throughout glabrescent. Rhizome woody, branched,
developing erect, sterile, leafy shoots; fertile shoots (20)35-45 cm
high, straight or slightly bent, ascending at base, brownish or brownish-
violet, branched (var. aschurbajevii Winkl.) or almost simple, leafy.
All leaves simple, sessile, 1.5—3.0(4.0) cm long, lanceolate; lower leaves
usually subobtuse, upper leaves acuminate. Capitula on short peduncles,
divergent or drooping, globose, 2.0—3.5 mm in dia, crowded on branches
or main stem, forming narrow, more or less long, racemose-paniculate
inflorescence. Outer involucral bracts broadly lanceolate or oval, inner
bracts larger, broadly oval, with wide scarious margin. Peripheral florets
pistillate, 12-13, their corollas narrowly tubular; stigma lobes exserted
from [corolla] tube, 2-3, linear, obtuse, divergent; disk florets 12-13,
staminate, their corollas conical, with purple teeth; anthers linear; stigma
of abortive pistil narrow funnel-shaped, not lobed. Flowering August.

Clayey and rubbly-clayey meadow-steppe slopes of mountains, in
alpine zone and lower.—Soviet Central Asia: Dzhungaria-Tarbagatai,
Tien Shan, Pamiro-Alai Region. General distribution: Dzhungaria-
Kashgaria. Described from Lake Kara-Kul in Pamir. Type in Leningrad.

Note. The reported occurrence of A. dracunculus in western Tibet
(Hook. Fl. Brit. 1882, 122), most probably relates to A. pamirica
Winkl.

Economic Importance. According to the data of M.I. Goryaev,
G.K. Kruglykina, and E.I. Satdarova (1959), the plant contains alkaloids.
According to Goryaev and Sazonova, the yield of essential oil is 0.06—
0.2%; the oil has an unpleasant smell and contains aldehydes and
ketones. The distillate is very turbid.

85. A. dracunculiformis Krasch. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk. SSSR, IX (1946) 172.

Perennial. Root woody, vertical. Stem herbaceous, 40—80 cm high,
thick, erect, greenish-yellow, glabrous, sulcate-ribbed, almost woody
at base, weakly leafy. Leaves yellowish-green or green, somewhat thick,

535

508

glabrous, 3-6 cm long and 4—6 cm wide, almost simple, more or less
with 3 deep lobes at apex, lobes 1—2 cm long, lanceolate, subacute or
subobtuse, contorted, or leaves simple, long-acuminate, gradually
narrowed toward base, upper bracteal leaves simple, lanceolate, 5—10
mm long. Capitula on short peduncles, drooping, subglobose, 4-5 mm
in dia, in long, narrow-pyramidal panicle. Involucral bracts glabrous,
lustrous, white scarious on margin; outer involucral bracts narrowly
lanceolate, inner bracts ovate, subobtuse. Peripheral florets pistillate,
their corollas narrowly tubular, enlarged at base; stigma lobes narrowly
linear, acute, exserted from corolla tube; disk florets staminate, their
corollas reddish-violet in upper part, weakly hairy; anthers linear; stigma
of abortive pistil narrow funnel-shaped, not lobed. Flowering July to
August.

River valleys and slopes of terraces, along northern edge of forest-
tundra and in southern part of arctic zone.—Arctic: Arctic Siberia
(lower reaches of Lena, Olenek, Indigirka and Kolyma rivers). Endemic.
Described from lower reaches of Lena River. Type in Leningrad.

86. A. daghestanica Krasch. and Por. in Izv. Bot. Sada Akad.
Nauk SSSR, XXX (1932) 709-710; Grossh. Fl. Kavk. IV, 138.

Perennial. Plant 40-50 cm high, densely hairy with appressed hairs,
later subglabrous; rhizome slender, ascending, developing sterile leafy
shoots and fertile, erect, more slender, brown or violet-brown, straight
or slightly flexuous branched stems. Cauline leaves simple, 1.0—3.5 cm
long and 1-3 mm wide, narrowly linear, short-acuminate, strongly
deflexed, somewhat stiff, uppermost small, narrowly paniculate [sic.;
recte narrowly lanceolate], often 3-6 mm long. Capitula pedunculate,
1.5-4.0 mm long, drooping, remote, subglobose, 1-2 mm in dia, in
depauperate pyramidal paniculate inflorescence. Involucral bracts
glabrous, lustrous, with wide scarious border; outer involucral bracts
ovate, inner bracts large, obovate. Peripheral florets 10, pistillate, their
corollas filiform-tubular; stigma lobes linear-filiform, exserted from
corolla tube, divergent; disk florets 5—6, staminate, their anthers linear
(apical appendages of anthers obtuse), large, basal appendages smaller,
subobtuse; stigma of abortive pistil almost funnel-shaped, densely [sic.;
recte deeply] lobed. Achenes about 0.8 mm long, ovoid, somewhat
flat, finely ribbed, brown. Flowering August.

Southern steppe slopes of mountains, in steppe zone, at 1,000-
1,900 m.—Caucasus: Dagestan. Endemic. Described from Dagestan.
Type in Leningrad.

87. A. glauca Pall. in Willd. Sp. pl. III (1800) 1831; Ldb. Fl. Alt.
IV, 87; Bess. in Bull. Soc. Nat. Mosc. VIII, 57; DC. Prodr. VI, 97;

536

509

Ldb. Fl. Ross. II, 563; Turcez. Fl. baic.-dahur. II, 2, 50; Kryl. Fl. Alt.
III, 633; Fedtsch. Perech. Rast. Turk. IV, 191; Rydb. North Am. FI.
34, part 3, 251; Pampan. in Nuov. Giron. Bot. Ital. n. s. XXXIV, 657;
Fl. Yugo-Vost. Evrop. Ch. SSSR, VI, 350; Kryl. Fl. Zap. Sib. XI,
2770; Poljak. in Majevski, Fl. 4583.—Exs.: GRF Nos. 67, 3180a,
3181a, b, c, d.

Perennial. Rhizome horizontal or obliquely ascending, densely
covered with rootlets, sometimes branched in upper part, sometimes
nodulose, thickened at point of origin of stem; whole plant grayish-
green or grayish (var. incana Bess.) from more or less dense tomentum
of short, stellate hairs. Stems few (2-12) or single, erect, less often
somewhat divergent, 15—20 cm (var. humilis Kryl.) or usually 25-70
cm high, green or reddish, often ribbed, branched, lower branches
sterile. Leaves undivided, linear or linear-lanceolate, 1-7 cm long and
1-7 mm wide; less often some leaves, mainly on sterile shoots, twice
or thrice divided. Capitula globose, 1.5—2.0 mm in dia, divergent or
drooping, in dense, more or less broadly paniculate inflorescence or
narrow (var. humilis Kryl.) paniculate inflorescence. Involucral bracts
smooth, glabrous; outer involucral bracts lanceolate; inner round, with
wide scarious border. Peripheral florets pistillate, 6-7, their corollas
filiform-tubular, stigma lobes narrowly linear, acute, exserted from
corolla tube; disk florets staminate, 6—7, their corollas narrowly conical,
with 5 teeth, anthers linear, apical appendages of anthers acute, basal
appendage not conspicuous, subacute, stigma of abortive pistil bilobed.
Achenes small about 0.5 mm long, ovate, fusiform, brown. Flowering
August. (Plate XXVIII, Fig. 3).

Alkaline meadows, near birch groves, steep riverbanks, old fallow
lands, less often stony slopes, forest-steppe and steppe zones.—Euro-
pean part: Volga-Kama, Trans-Volga; Western Siberia: Upper Tobol,
Irtysh, Altai; Eastern Siberia: Angara-Sayans, Dauria. General distri-
bution: Mongolia, North America. Described from Siberia. Type in
Berlin?

Note. Hooker (FI. Br. Ind., 1882) reports this species for the
western Himalayas, which is doubtful.

Series 2. Simplicifoliae Krasch.—Semishrubs, glabrous or weakly
hairy (hairs simple); leaves simple, linear-lanceolate, partly with 2-5
lobes in upper part.

88. A. dimoana M. Pop. in Pochv. Eksp. v Bass. rr. Syr-Dari i
Amu-Dari, II (1916) 75, 88.—Ic.: Ibid., Plates 22a, 22b, 22c.

Perennial. Semishrub, 70-90 cm high. Root thick, woody, cordlike,
with thick, woody, strongly reduced, branched stem covered with dull,

537

510

grayish-brown bark. Fertile stems (branches) erect, woody almost to
apex, strongly branched, smooth, lustrous, stramineous. All leaves
undivided, linear or linear-narrowly lanceolate, succulent, with short
appressed hairs, short-acute; middle and partly upper cauline leaves 4—
6 cm long, uppermost bracteal leaves linear, 3-5 mm long. Capitula
sessile or on short peduncle, somewhat remote, divergent or drooping,
ovoid, 3.5—4.0 mm long and 2 mm in dia, in lax racemes forming lax
panicle or branches. Involucral bracts densely hairy; outer involucral
bracts small, almost round, or oval strongly convex, brownish on outer
side; inner bracts larger, oblong-ovate, scarious along margin. Peripheral
florets pistillate, 3-4, their corollas small, tubular, narrowed in upper
part; stigma lobes linear, divergent; disk florets staminate, 3-5, their
corollas conical; anthers linear, apical appendages of anthers acute,
basal appendages short, subobtuse; stigma of abortive pistil funnel-
shaped at apex, not lobed. Achenes to 3 mm long, pyriform-ovoid,
fusiform, ribbed. Flowering July to August.

Sands of desert zone.—Soviet Central Asia: Kara-Kum. Described
from vicinity of Khiva. Type in Tashkent.

89. A. kelleri Krasch. in Otch. o Rab. Pochv.-Bot. Otr. Kazakhsk.
Eksp. Akad. Nauk SSSR, IV, 2(1930) 274; Krascheninn. in Spisok
Rast. Gerb. Fl. SSSR, X, 98.—Exs.: GRF No. 3188.

Semishrub, 60-80 cm high, densely hairy with appressed hairs,
later glabrous; root woody, vertical, thick, developing short, rather
thick, woody, perennial, prostrate or erect sterile shoots covered with
gray, peeling bark. Stems rather numerous, strong, 3-4 mm thick,
smooth, straw-yellow or brownish, woody in lower part, branched above.
Leaves succulent, yellowish-green, 3-5(9) cm long and 1-3 mm wide,
simple, narrowly linear or in upper part with 2-5 lobes, lobes linear,
acute; upper cauline leaves and leaves on branches small. Capitula
usually on long peduncle, ovoid, 3-4 mm long, drooping, in broad,
lax, paniculate inflorescence. Involucre 3-rowed, glabrous; outer
involucral bracts succulent, small, round, middle bracts ovate, inner
ovate-oblong with wide scarious border. Peripheral florets pistillate,
fertile, 5, their corollas, narrowly tubular, punctate-glandular; stigma
lobes linear, short-acuminate, divergent; disk florets staminate, 9, their
corollas narrowly conical, anthers on short filaments, linear, apical
appendages of anthers subulate, basal appendages short, round; stigma
of abortive pistil funnel-shaped, not lobed. Achenes to 1.8 mm long,
oblong-ovoid, fusiform, finely ribbed, brown. Flowering August.

Sands of desert zone.—Soviet Central Asia: Aralo-Caspian Region,
Lake Balkhash Region (Muyun-Kumy), Kara-Kum. Endemic. Described
from Akhcha-Kuim in Turkmenia. Type in Leningrad.

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511

Series 3. Psammophilae Poljak.—Semishrubs; lower cauline leaves
long-petiolate, divided into 3—7 lobes or pinnately cut, lobes rather
long.

90. A. trautvatteriana Bess. in Mém. Acad. Sc. Pétersb. IV (1845)
464.—A. pauciflora MB. FI. taur.-cauc. II (1808) 290 p. p. exl. syn.—
A. pauciflora MB. B. cernua Bess. in Bull. Soc. Nat. Mosc. VIII (1835)
19.—Ic.: Bess. op. cit. tab. V.

Perennial. Semishrub; root woody, branched in upper part, with
short, woody, leafy sterile shoots. Stems 50-65 cm high, rather thick,
strong, woody in lower part, covered with grayish-brown, peeling bark,
brownish above, initially pubescent, later subglabrous, leafy, branched.
Leaves densely hairy with appressed hairs, later subglabrous; lower
cauline leaves and leaves of nonflowering shoots in fascicles, long-
petiolate, 5—8 cm long, pinnatisect in 3-7 palmate lobes, lobes narrowly
linear, subacute, (0.5)0.7—2.0(2.7) cm long; middle and upper leaves
shorter, sessile, usually divided into 3—7 lobes; uppermost leaves short,
simple, linear. Capitula ovate, 3 mm long and 1.5-1.8 mm in dia,
sessile, densely spicate, crowded on branches, forming conical panicle.
Outer involucral bracts succulent, broadly elliptical, short-acuminate,
densely hairy, inner bracts larger, oval, keeled, glabrous or weakly
hairy, with wide scarious margin. Peripheral florets pistillate, 5-7,
their corollas in narrow in upper part, expanded below like bottle,
stigma lobes linear, subobtuse, divergent; disk florets staminate, 7-9,
their corollas narrowly conical, anthers linear; stigma of abortive pistil
funnel-shaped, short-ciliate. Flowering August.

Coastal sandy areas.—European part: Black Sea Region, Lower
Volga. Endemic. Described from southern Ukraine (near Odessa and
mouth of Dnieper River) and from vicinity of Astrakhan (var. B.).
Type in Kiev.

91. A. salsoloides Willd. Sp. pl. III (1800) 1832; Bess. in Bull.
Soc. Nat. Mosc. VIII, 17; DC. Prodr. VI, 94; Bess. in Mém. Acad. Sc.
Pétersb. IV, 455; Ldb. Fl. Ross. II, 560; Boiss. Fl. or. III, 362; Fedtsch.
Perech. Rast. Turk. IV, 190; Grossh. Fl. Kavk. IV, 141; Krascheninn.
in Fl. Yugo-Vost. Evrop. Ch. SSSR, VI, 351; Kryl. Fl. Zap. Sib. XI,
2766; Poljak. in Majevski, Fl. 585.—A. tanaitica Klok. in Bot. Mat.
Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954) 362, 364.—Ic.: Bess.
op. cit. (1845) tab. II.—Exs.: GRF No. 3184a, b.

Perennial. Short semishrub, 20—30(45) cm high, with thick, woody
root developing short, woody, strongly branched, perennial sterile
shoots. Flowering stems numerous, erect, in lower part woody, brownish,
glabrous or sometimes sparsely pubescent. Leaves glaucescent, glabrous

539

512

or sometimes covered with inconspicuous hairs; lower cauline leaves
petiolate, 2.0-3.5 cm long, pinnately cut; middle cauline leaves
palmately cut, upper simple, {—2 cm long, linear, subacute. Inflorescence
narrow, short, more or less dense raceme; capitula ovate, 2.5—3.0(3.5)
mm long. Involucral bracts oval, glabrous, inner bracts wider, scarious
along margin. Peripheral florets pistillate, usually 4—6, their corollas
narrowly tubular; stigma lobes narrowly linear, exserted; disk florets
staminate, 8—9, their corollas tubular-conical; anthers on short filaments,
linear, apical appendages, of anthers narrowly angular, basal appendages
short, subobtuse; stigma of abortive pistil funnel-shaped, with 2 short
divergent lobes. Achenes to 1.2 mm long, ovate, somewhat flat, with
slightly attenuate apex, dark brown. Flowering August.

Slopes of limestone outcrops, chalks, sometimes on outcrops of
igneous rocks.—European part: Volga-Don, Trans- Volga, Volga-Kama,
Black Sea Region, Lower Don, Lower Volga; Caucasus: Dagestan;
Western Siberia: Upper Tobol (southwest). Endemic. Described from
Western Siberia. Type in Berlin.

Note. Hooker’s report (Hook. Fl. Br. Ind., 1882, 321) about the
occurrence of A. salsoloides Willd. in western Tibet is erroneous.
According to the determination of I.M. Krascheninnikov, this plant
belongs to Artemisia welbyi Hemsl. and Pears. The reported distribution
of this species in the Altai is not supported by specimens.

92. A. halodendron Turcz. ex Bess. in Bull. Soc. Nat. Mosc. VIII
(1835) 17; DC. Prodr. VI, 94; Bess. in Mém. Acad. Sc. Pétersb. IV,
459; Ldb. FI. Ross. Ii, 560; Turcz. Fl. baic.-dahur. II, 2, 48; Maxim.
in Bull. Acad. Sc. Pétersb. VIII, 527; Kom. Fl. Manchzh. III, 658.—
Ic.: Bess. op. cit. (1845) tab. III.

Perennial. Semishrub to 50 cm high; root woody with thick, woody,
perennial, sterile, leafy shoots. Stem ascending or erect, woody in
lower part, brown, strongly branched with somewhat divergent branches.
Leaves green, moderately hairy with long appressed hairs, later
subglabrous; leaves of sterile shoots and lower cauline leaves petiolate,
3-5(6) cm long, usually once or almost twice pinnately incised into
narrowly linear, often 1.0—1.5(2.0) cm long and 0.5—1.0 mm wide short-
acuminate lobes; middle and upper leaves sessile, pinnate to palmately
cut, at base with 3-5 narrow, incised auricles; uppermost bracteal leaves
simple, linear, sometimes with 3-5 lobes. Capitula on short peduncles
or sessile, ovoid, 4-5 mm long and 2.0-2.5 mm in dia, straight, in
sparse racemes on weakly divergent branches, forming narrow panicle.
Involucral bracts oval, glabrous, smooth, with wide scarious border,
peripheral florets 5, pistillate, fertile, their corollas narrowly tubular,
slightly enlarged at base; stigma lobes linear, obtuse, divergent; disk

540

513

florets staminate, 10-11, their corollas narrowly conical; anthers linear,
apical appendages of anthers acute, basal appendages smaller, obtuse,
roundish; stigma of abortive pistil funnel-shaped, not lobed. Achenes
1.5 mm long, ovate, with somewhat attenuate apex, finely ribbed, dark
brown. Flowering August. (Plate XXVIII, Fig. 1).

Saline lands, saline sands, sometimes on slopes of [mud] volcanic
cones.—Eastern Siberia: Dauria. General distribution: Mongolia,
northern China (west). Described from saline lands of Dauria and
Mongolia. Type in Leningrad.

93. A. quinqueloba Trautv. in Bull. Soc. Nat. Mosc. XXXIX, I
(1866) 348; Fedtsch. Perech. Rast. Turk. IV, 190.

Perennial. Semishrub, (40)50-—75(85) cm high; root vertical, woody,
thick, thickened above, with strongly reduced, branched, woody trunk
covered with brownish-gray fibrously cracking bark. Stems rather
numerous, straight or slightly flexuous, more slender, virgate, branched,
stramineous, usually densely hairy or partly glabrous (particularly in
fall). Leaves more or less hairy on both sides with dense appressed
hairs; lower leaves long-petiolate, usually 3—4(5) cm long, pinnatisect
into 7 linear, 5—10(15) cm long and 1 mm wide, subacute lobes; middle
and upper leaves with 3-S incised, sessile, lobes at base with palmately
cut or ternate auricles; uppermost leaves bracteal, simple, linear.
Capitula pedunculate, drooping, broadly ovate or subglobose, to 2.5
mm in dia and 3 mm long, remote on divergent branches in lax,
spreading, racemose-paniculate inflorescence. Involucral bracts hairy,
with short appressed hairs; outer involucral bracts oblong-ovate, inner
bracts wider, oval, with scarious border. Peripheral florets pistillate,
3-5, their corollas narrowly tubular, punctate-glandular, stigma lobes
linear, acute, divergent; disk florets staminate, 5-9, their corollas
narrowly conical, with straight, hairy teeth, anthers linear; stigma of
abortive pistil funnel-shaped, not lobed. Flowering August.

Sands of desert zone.—Soviet Central Asia: Aralo-Caspian Region,
Lake Balkhash Region. Endemic. Described from sands along Sarysu
River. Type in Leningrad.

Note. It is distinguished from A. arenaria DC. by less divided
leaves, broadly ovoid or subglobose capitula, a scarcely hairy involucre,
and squarrose, stiff hairy corolla teeth of the disk florets.

94. A. arenaria DC. Prodr. VI (1837) 94; Bess. in Mém. Acad.
Sc. Pétersb. IV, 474; Ldb. FI. ross. II, 2, 561; Boiss. Fl. or. III, 362;
Fedtsch. Perech. Rast. Turk. IV, 190; Grossh. Fi. Kavk. IV, 141;
Krascheninn. in Fl. Yugo-Vost. Evrop. Ch. SSSR, IV, 352; Krascheninn.
in Kryl. Fl. Zap. Sib. XI, 2767; Poljak. in Majevski, Fl. 585.—A.

54

—

514

pauciflora MB. FI. taur.-cauc. II (1808) 290 p. p.; Bess. in Bull. Soc.
Nat. Mosc. VII, 18.—A. sabulosa Willd. in herb. and A. fruticosa
Willd. ex adnot. Stev.—A. inodora steveniana Bess. in Bull. Soc. Nat.
Mosc. VIII (1835) 44.—A. inodora Ldb. Fl. Ross. II (1844-1846)
566, pro parte (non Web.).—A. tschernieviana Bess. in Bull. Soc. Nat.
Mosc. VIII (1835) 31; DC. Prodr. VI, 95; Bess. in Mém. Acad. Sc.
Pétersb. IV, 480; Ldb. Fl. Ross. II, 562; Krascheninn. in Spisok Rast.
Gerb. Fl. SSSR, XI, 47.—Ic.: Bess. in Mém. Acad. Sc. Pétersb. IV,
tab. XII.—Exs.: GRF Nos. 3187, 3288a, b, sub. nom. A. tschernieviana
Bess.

Perennial. Semishrub (35)50—75(100) cm high; root thick, woody,
with woody, perennial, strongly reduced, branched trunk covered with
brownish-gray cracking fibrous bark. Flowering stems rather numerous,
straight or weakly flexuous, curved, more slender, virgate, straw-yellow,
more or less hairy, later glabrous or subglabrous, branched above,
branches more or less divergent. Leaves hairy, later glabrous; lower
cauline leaves withering early petiolate, 2—-5(6) cm long, twice pinnately
incised, terminal lobe often 5-15 mm long, narrowly linear, short-
acuminate; middle cauline leaves usually sessile, simple pinnate or
with 3—5 lobes; uppermost leaves bracteal, simple, narrowly linear or
sometimes with 2 lateral lobes at base. Capitula on short peduncle,
ovate or broadly ovoid, 2.5—3.0-4.0 mm long and 1.6—2.5(3.0) mm in
dia, in lax or rather dense racemes on branches, forming spreading
paniculate inflorescence. Involucral bracts elliptical, with scarious
border; outer involucral bracts herbaceous, inner bracts membranous,
smooth. Peripheral florets pistillate, fertile, 3-5, their corollas obconical,
enlarged at base; stigma lobes narrowly linear, divergent; disk florets
staminate, 6-9, their corollas narrowly conical; anthers linear, apical
appendages of anthers linear, acute, basal appendages shorter, subacute;
stigma of abortive pistil with 2 short lobes. Achenes 1 mm long, oblong-
ovate, somewhat flat, finely ribbed, blackish-brown. Flowering August.

Sands in steppe zone, sometimes near pine forests, along
seacoasts.—European part: Volga-Don (southern part), Black Sea
Region, Bessarabia, Lower Don, Lower Volga; Caucasus: Eastern
Transcaucasia, Dagestan; Soviet Central Asia: Aralo-Caspian Region,
Kara-Kum, Kzyl-Kum, Lake Balkhash Region. General distribution:
Balkans-Asia Minor. Described from area between Odessa and mouth
of Dniester River. Type in Geneva.

Note. Var. tschernieviana (Bess.) Grossh. (FI. Kavk IV, 141),
distributed in the western Caspian Region, is distinguished by long
branchlets and remote capitula. However, separation of this variety as
an independent species, as proposed by Besser, is not adequately
justified.

542

515

Economic Importance. From a half-dried plant doubtfully be-
longing to A. arenaria DC., V.I. Isaev (1932) obtained by distillation
0.9-1.1% of light yellow, highly odorous essential oil with the follow-
ing constants: D'*—0.869; «wD—0.06°; n,—1.4820; acid number—0.3,
ester number—33; ester number after acetylation—27.8. The presence
in the oil of 2% phenols and 2% ketones was established. A plant
collected 6 km from Irgiz, according to N.I. Bakanina (Goryaev, 1952),
yielded 0.14% essential oil from absolute dry matter. The oil is yellow
in color with a pleasant smell, and a pleasant but slightly pungent
taste. Constants of the oil are: D,?°—0.8692; n,?°—1.4183; acid num-
ber—2.05; ester number—28.4; saponification number—30.45. Its
reaction to aldehydes was positive.

It is a good stabilizer of shifting sands and ought to be widely
used for such purposes. The data on its fodder qualities are inadequate,
and it is necessary to conduct additional studies for a definitive
evaluation of the nutritive value of this psammophilous wormwood for
pasturing animals (I.V. Larin, 1956).

95. A. albicerata Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, IX (1946) 173.—Exs.: GRF No. 3289.

Perennial. Semishrub, (35)45—65(70) cm high, densely appressed
hairy, later glabrous; root vertical, woody, with short, perennial, woody,
sterile shoots covered with gray peeling bark. Fertile shoots straight,
smooth, lustrous, whitish-waxy in lower part, brownish-yellow above,
with divergent branches. Leaves somewhat thick, green; lower and
middle cauline leaves 5-8 cm long, mostly pinnately cut or ternate,
with fewer lobes, lobes narrowly linear, semicylindrical, 3-4 cm long
and 1 mm wide, with stiff somewhat obtuse apices; upper leaves simple,
undivided or sometimes some with 5-7 lobes, almost pinnately cut,
lobes strongly elongated, flat; uppermost bracteal leaves small. Capitula
numerous, ovoid, 2.5-3.0, mm long and 2 mm in dia, on short peduncle,
drooping, in broad, pyramidal-paniculate inflorescence. Involucre
glabrous; outer involucral bracts small, almost round, fleshy, gibbous
on outer side; inner bracts oval, convex, with white scarious border.
Peripheral florets pistillate, few (3-4), their corollas narrowly tubular;
stigma lobes linear, obtuse, straight; disk florets staminate (7-9), their
corollas tubular-conical, anthers on short filaments, lanceolate-linear,
apical appendage of anthers acute, basal appendage shorter, acute,
stigma of abortive pistil with 2 short lobes. Achenes up to 1 mm long,
ovoid, flat, finely ribbed, dark brown. Flowering August.

Sands of desert zone.—Soviet Central Asia: Lake Balkhash Region.
Endemic. Described from Ili River valley. Type in Leningrad.

54

WwW

516

Note. This species is very close to A. arenaria DC., but differs
from it by usually less divided leaves and rather long, narrowly linear
leaf lobes. In my opinion, these characters are not adequate to treat it
as. a species.

96. A. saposhnikovii Krasch. ex Poljak. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XVII (1955) 412.

Perennial. Whole plant glabrous, 25-35 cm high; root vertical,
rather thick, woody. Stems several or numerous, straight, slender, pale
straw-yellow, woody in lower part, sometimes red, simple or almost
simple. Lower cauline, leaves petiolate, 2.0—-2.5 cm long, pinnatisect
or more or less twice pinnately incised, lobes fewer, linear, 3-7 mm
long, subacute; middle and partly upper cauline leaves sessile (usually
with linear auricles at base), with 5 lobes; uppermost leaves bracteal,
simple, linear. Capitula on short peduncles, approximate, divergent or
drooping, broadly ovoid, 3-4 mm long and 2.0—2.5 mm in dia, at stem
apices in simple raceme, below on short, weakly divergent branches in
narrow racemose-paniculate inflorescence. Involucral bracts glabrous,
with wide scarious border; outer involucral bracts oval, small, inner
bracts considerably larger, oblong or elliptical, acute. Peripheral florets
pistillate, 4, their corollas tubular, enlarged at base; stigma lobes 2-3,
linear, obtuse, divergent; disk florets staminate, 8; anthers linear; stigma
of abortive pistil with 2 short, straight lobes. Achenes to 1.5 mm long,
ovate, terete, finely ribbed, brown. Flowering August. (Plate XXX,
Fig. 3).

Stony mountain slopes, in valleys on gravel beds.—Soviet Central
Asia: Tien Shan. Described from Inylchek River valley. Endemic. Type
in Leningrad.

97. A. songarica Schrenk, Enum. pl. nov. I (1841) 49; Ldb. FI.
Ross. II, 561; Boiss. Fl. or. III, 362; Krasn. in Spisok Rast. Sbor. in
V. Tyan-Shan. 67; Fedtsch. Perech. Rast. Turk. IV, 190; Pampan. in
Nuov. Giorn. Bot. Ital. n. s. XXXIV, 03.

Perennial. Semishrub, 30—40 cm high, weakly hairy when young,
later glabrous; root woody, vertical, usually with numerous, strongly
reduced, woody, perennial, ascending shoots covered with light
yellowish-gray, peeling bark. Fertile shoots numerous, virgate, somewhat
flexuous, smooth, stramineous brownish, strongly branched, with almost
horizontally divergent branches. Lower and middle cauline leaves
pinnately cut, with 5-7 oblong, simple lobes or palmately cut: lobes
linear, 4-12 mm long, short-acuminate; upper bracteal leaves small,
sessile, narrowly linear. Capitula ovate-globose or subglobose, sessile
or on short peduncle, 1.5—2.0 mm in dia, remote, divergent, in lax,

544

517

spreading panicle. Involucral bracts roundish-ovate, with wide scarious
border. Peripheral florets pistilate, 2—4, their corollas narrowly tubular;
stigma lobes linear, straight; disk florets staminate, 6-8, their corollas
conical, pinkish; anthers linear; stigma of rudimentary pistil funnel-
shaped. Achenes 1.5 mm long, ovate, indistinctly finely ribbed, brown.
Flowering July.

Hummocky sands and sandstone outcrops, in desert zone.—Soviet
Central Asia: Lake Balkhash Region (basin of Ili River). General
distribution: Dzhungaria-Kashgaria. Described from Dzhungaria. Type
in Leningrad.

Series 4. Palmatisectae Poljak.—Leaves grayish from thin, silky,
appressed hairs; lower cauline leaves round or oval, palmately cut into
4-7 lobes.

98. A. lipskyi Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVII (1955) 411.

Perennial. Root thick, woody, with strongly reduced, erect, woody,
peripheral shoots covered with light brown, peeling bark. Fertile shoots
few, 35-40 cm high, arcuate or almost straight, weakly flexuous,
brownish, almost round, finely tomentose, weakly leafy, branched.
Leaves whitish-lanate from thin, silky, appressed hairs on both sides,
lower cauline leaves long-petiolate, 3-5 cm long, petioles strongly
thickened at base, amplexicaul, lamina round or broadly oval, palmately
parted into 4—7 lobes, lobes with (2)3—5 secondary lobes, such lobes
lanceolate-linear, short-acuminate, 3-7 mm long; middle leaves short-
petiolate or sessile; bracteal leaves mostly simple, linear. Capitula ovoid,
to 2 mm wide, on short peduncles or subsessile, crowded on branches
in narrowly paniculate inflorescence. Involucral bracts whitish-lanate,
with scarious margin; outer bracts oval, inner bracts somewhat larger,
oblong-ovate. Receptacle convex, hemispherical, glabrous. Peripheral
florets pistillate, 6, their corollas narrowly tubular, enlarged at base;
stigma lobes linear, acute, straight, divergent; disk florets staminate,
12, their corollas narrowly conical, anthers linear, apical appendages
of anthers acute, basal appendages short, obtuse; stigma of abortive
pistil funnel-shaped. Flowering August.

Sandy places in mountain river valleys.—Soviet Central Asia:
Pamiro-Alai Region (Karategin). Described from Surkhob. Type in
Leningrad.

Note. Close to A. kuschakeviczi Winkl. but is distinguished by
greater height, narrow capitula, and a hairy punctate-glandular corolla.

545

518

99. A. kuschakeviczi. Winkl. in Tr. Peterb. Bot. Sada, XI (1890)
330; O. Fedtsch. Fl. Pam. 116; Fedtsch. Perech. Rast. Turk. IV, 190.

Perennial. Whole plant whitish-silky from fine appressed hairs;
root thick, woody, vertical, with underground, short, woody, perennial
shoots with above-ground parts having densely leafy apices forming
thickset tussocks. Stems rather numerous, 5—12(15) cm high, erect or
ascending, weakly leafy, simple or with short branches, covered with
remnants of leaf petioles at base. Lower cauline leaves and leaves of
sterile shoots petiolate, 1-3 cm long, petioles thickened at base, lamina
round, divided into 5—7 palmate lobes, with 3-5, less often 2 secondary
lobes, less often simple, such lobes lanceolate-linear, acute, 2-5 mm
long; upper cauline leaves sessile, divided into 3-5 lobes; uppermost
leaves bracteal, simple, linear. Capitula ovoid, 25 mm in dia, on short
peduncles, upper capitula crowded; lower capitula remote, in simple
racemose, less often racemose-paniculate inflorescence. Involucral
bracts oval, with scarious border, outer involucral bracts convex,
densely hairy, inner subglabrous. Receptacle convex, conical, glabrous.
Peripheral florets pistillate, fertile, 5-6, their corollas narrowly tubular,
somewhat enlarged at base; stigma lobes linear, acute; disk florets
staminate, 9, their corollas tubular-conical, glabrous; anthers linear,
apical appendages acute, basal appendages short, obtuse; stigma of
abortive pistil funnel-shaped. Achenes to 2 mm long, narrowly linear,
with 4-angled, blackish-brown. Flowering August.

Rubbly-gravelly deposits along mountain streams, near lakes in
high altitude-desert zone, at 3,500-4,000 m.—Soviet Central Asia:
Pamiro-Alai Region. Endemic. Described from Chonsu River and Lake
Karakul in Pamir. Type in Leningrad.

Note. It is quite likely that this species is found in the high-alti-
tude areas of western China and the Himalayas bordering eastern Pamir.

Series 5. Japonicae Krasch. ex Poljak.—Leaves of sterile shoots
and lower cauline leaves divided, with acute teeth, middle cauline
leaves pinnatifid.

100. A. japonica Thunb. Fl. Japon. (1784) 308; Willd. Sp. pl. III,
1847; Bess. in Bull. Soc. Nat. Mosc. VIII, 50; Kom. Fl. Manchzh. III,
654, excl. synon.—A. japonica Thunb. a. japonica Maxim. in Bull.
Acad. Sc. Pétersb. VIII (1872) 526, excl. synon.; Nakai, Fl. Korean.
33; Pampan. in Nuov. Giron. Bot. Ital. n.s. XXXV, 660, includ. var.
japonica Maxim. and f. typica Nakai; Sugaw. Ill. Fl. Saghal. IV,
1807.—Ic.: Sugaw. ibid. tab. 828.

Perennial. Plant glabrous, 65-90 cm high; root thick with long
(sometimes almost as long as stem). Sterile shoots and densely leafy

546

519

and solitary or few, erect, glabrous, branched stems. Leaves of sterile
shoots and lower cauline leaves 5—8 cm long, almost round-cuneate or
oblong-cuneate, incised-acute-toothed, entire below, at base with 1-2
pairs of narrow, linear, acute lobes; middle cauline leaves somewhat
shorter, narrowly cuneate, incised-acute-toothed or almost pinnately
divided; upper leaves smaller, simple, lanceolate or lanceolate-linear,
entire or with few, acute, linear-lanceolate teeth; upper-most leaves
bracteal, very small, usually not exserted from inflorescence, simple,
narrowly lanceolate. Capitula numerous, pedunculate, broadly ovoid,
to 2 mm long, more or less approximate on branches, forming broad
paniculate inflorescence. Involucral bracts oval, smooth, with more or
less wide scarious margin; outer bracts smaller than inner. Peripheral
florets pistillate, 7, their corollas obconical, strongly expanded at base;
stigma lobes exserted from corolla tube, narrowly linear, subacute,
recurved; disk florets staminate, 7, their corollas conical, glabrous,
anthers linear, apical appendages acute, basal appendages shorter,
subobtuse, stigma of abortive pistil funnel-shaped, not lobed. Achenes
1 mm long, fusiform, ovoid, finely sulcate, dark-brown. Flowering
August.

Riverbanks and gravelly deposits, dry meadows, slopes of mountain
ridges, among scrub thickets or forests.—Far East: Ussuri. General
distribution: Northeastern China, Japan. Described from the vicinity
of Nagasaki. Type in Uppsala.

101. A. littoricola Kitam. in Acta Phyt. et Geobot. V (1936) 94.—
A. desertorum Takeda in Bot. Mag. Tokyo, XXIV (1911) 21 non
Sprengel; Nakai in Bot. Mag. Tokyo, XXVI, 99.—A. japonica var.
macrocephala f. sachalinensis Pamp. in Nuov. Giron. Bot. Ital. n.s.
XXXIV (1927) 668.—A. japonica var. rotundifolia f. vestita Pamp.
ibid. p. 669, pro parte.

Perennial. Rhizome creeping, woody, developing long, sterile, leafy
shoots. Stems erect or weakly divergent, 30-100 cm high, grayish-
lanate, later glabrous, branched, branches arcuate or straight, divergent.
Leaves of sterile shoots oblong-cuneate, including petiole, 5-10 cm
long, palmately or pinnately divided or pinnately lobed, lobes attenuate,
pinnate, secondary lobes obtuse, somewhat succulent, on both sides
grayish-woolly; lower cauline leaves withering early; middle cauline
leaves oblong-cuneate or elliptical, with auricles, pinnately divided,
palmate or ternate, lobes remote, long, linear-lanceolate, with distinct
venation beneath, grayish-tomentose, later subglabrous; upper cauline
leaves simple, linear-lanceolate, obtuse. Capitula numerous, crowded,
pedunculate, subglobose, to 2.5—-3.0 mm long and up to 2 mm in dia,
on branches, with dense drooping racemes forming long pyramidal

520

Plate XXVI.
Artemisia lagocephala (Bess.) DC. Plant habit. involucral bracts; peripheral

547

pistillate floret; bisexual disk floret.

54

Vo)

521

panicle. Involucral bracts rectangular, oval, with scarious border, outer
bracts small, on spine gibbose; inner involucral bracts larger, obtuse.
Peripheral florets 7-8, pistillate, their corollas tubular, enlarged; stigma
lobes exserted from corolla tube, linear, obtuse, divergent; disk florets
6, pistillate, their corollas narrowly campanulate, apical appendages of
anthers acute, basal appendages smaller, subobtuse; anthers linear;
stigma of abortive pistil funnel-shaped, undivided. Achenes 1 mm long,
oblong-ovoid, somewhat flat, smooth, brown. Flowering August to
September.

Seacoasts.—Far East: Sakhalin (and Kuril Islands). General
distribution: Japan. Described from Hokkaido Island. Type in Tokyo.

102. A. desertorum Spreng. Syst. veg. III (1826) 490; Bess. in
Bull. Soc. Nat. Mosc. VIII, 64; Turcz. Fl. baic.-dahur. II, 2, 51; Maxim.
Pr..Fl. Maur. 157; Krascheninn. in Spisok Rast. Gerb. Fl. SSSR, XI,
3284.—A. desertorum var. sprengeliana Bess. in Bull. Soc. Nat. Mosc.
VIII (1835) 65; DC. Prodr. VI, 98; Ldb. Fl. Ross. II, 564; Pampan. in
Nuov. Giron. Bot. Ital. n. s. XXXIV, 650; Ling Jong in Contrib. Inst.
bot. Acad. Peiping. II, No. 10 (1934) 512.—A. japonica Thunb. var.
desertorum Maxim. in Bull. Acad. Sc. Pétersb. VIII (1872) 527.—A.
japonica Thunb. B. manchurica Kom. Fl. Manchzh. III (1907) 654;
Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1035.—Ic.: Kom. and
Alis. op. cit. Plate 306.—Exs.: GRF No. 3284.

Perennial. Plant glabrous, with short woody rhizome thickened in
upper part. Stems solitary, less often few, (50)65—90 cm high, straight,
smooth, light brown, sometimes reddish-violet, leafy, branched above;
sterile shoots short, leafy. Leaves strongly variable in size and degree
of cutting; lower leaves long-petiolate, most often 4-10 mm long,
pinnatifid, with broadly linear or linear-lanceolate lobes often, in turn,
divided into 3 sharp teeth or oblong-obcuneate, strongly narrowed
toward base, more or less incised-toothed with acute teeth in upper part,
middle cauline and upper leaves smaller, short-petiolate or sessile, at base
with auricles divided into 3-5 linear lobes, lamina often divided into
3-5(7) lobes, less often pinnate, lobes linear, short-acuminate, undivided,
sometimes with short, subacute teeth; uppermost bracteal leaves simple,
small, linear. Capitula numerous, on short peduncles, divergent or drooping,
broadly ovoid, 2.0—-2.5 mm long, in dense racemes, clustered in narrow
compact panicle. Involucral bracts glabrous, with wide scarious border;
outer bracts oval; inner bracts slightly larger. Peripheral florets pistillate,
their corollas narrowly tubular; stigma lobes exserted from corolla
tube, linear, subobtuse, divergent; disk florets fewer, staminate, their
corollas narrowly conical, anthers linear, stigma of abortive pistil with
2 short divergent lobes. Achenes 0.5 mm long, ovate, convex, finely
ribbed, black. Flowering August. (Plate XXX, Fig. 4).

550

522

River valleys in dry meadows, mountain slopes, in scrub thickets
or in deciduous forests.—Eastern Siberia: Dauria; Far East: Zeya-
Bureya, Uda River Area, Ussuri. General distribution: Northeastern
China. Described from Siberia. Type in Berlin.

Note. Erroneously cited for Soviet Central Asia and western Tibet.

103. A. macilenta (Maxim.) Krasch. in Mat. po Istor. Fl. i Rast.
SSSR, II (1946) 156.—A. campestria var. macilenta Maxim. in Prim.
Fl. Amur. (1859) 158.—A. capillaris var. simplex Maxim. in Bull.
Acad. Sc. Pétersb. VIII (1872) 524.—A. commutata Bess. var. helmiana
Bess. in Bull. Soc. Nat. Mosc. VIII (1835) 69.—A. desertorum Spr.
var. macilenta Pampan. in Nuov. Giron. Bot. Ital. n. s. XXXIV (1927)
651.—Exs.: GRF No. 3183.

Perennial. Plant green, with occasional, very fine, appressed hairs.
Rhizome woody, short, nodulose above, developing long, sterile, leafy
shoots; stems straight, slender, glabrous, light brownish, sometimes
violet-brown in lower part, 40-70 cm high, almost simple, with short
branches above. Leaves glabrous; leaves of sterile shoots (clustered at
their apices) and lower cauline leaves 4-5 cm long including petiole,
twice pinnately incised, lobes narrowly linear, 7—-12(15) mm long, 0.3-
0.6(1.0) mm wide, scarcely acuminate; middle and upper cauline leaves
sessile or subsessile, palmately cut; uppermost bracteal leaves small,
simple, filiform-linear. Capitula numerous, approximate, pedunculate,
divergent or drooping, subglobose or broadly ovate, 2.0-2.5 mm long
and 1.0-1.5 mm in dia, in lax racemes on short branches usually
appressed to stem and forming narrow, dense panicle. Involucral bracts
glabrous, with wide scarious border; outer bracts oval; inner bracts
larger, broadly lanceolate, subacute. Peripheral florets pistillate, their
corollas small, obconical, at base strongly enlarged; stigma lobes
filiform-linear, straight; disk florets staminate, their corollas conical;
anthers on short filaments, linear, apical appendages of anthers acute,
basal appendages short, subobtuse; stigma of abortive pistil with 2
short, linear straight lobes. Achenes about 0.7 mm long, ovate, finely
ribbed, brown. Flowering August.

River valleys in sandy places, among scrubs, on slopes of
[mud]volcanic cones in forest zone.—Far East: Zeya-Bureya, Uda River
Area, Ussuri. General distribution: Northern China. Described from
Bureya Mountain. Type in Leningrad.

104. A. capillaris Thunb. Fl. Japon. (1784) 309; Willd. Sp. pl.
III, 1818; Spreng. Syst. veg. III, 487; Maxim. in Bull. Acad. Sc. Pétersb.
VIII, 524; Kom. Fl. Manchzh. III, 653 p. p.—A. capillaris var.
sachalinensis Pamp. in Nuov. Giron. Bot. Ital. n. s. XXXIV (1927)

5323

646; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1035; Ling Jong
in Contr. Inst. bot. Acad. Peiping. II, No. 10, 509 p. p.—A.
sachalinensis Tiles. ex Bess. in Bull. Soc. Nat. Mosc. VIII (1835) 48;
DC. Prodr. VI, 96; Ldb. Fl. Ross. Il, 562.—A. scoparia Maxim. op.
cit. 523; Nakai, Fl. Korean. (1911) 32.

Annual. Root vertical, slender. Stem solitary, slender, straight, 30-
50 cm high, brown or reddish-brown. Leaves of sterile shoots and
basal leaves sericeous pubescent, 2-4 cm long, simple pinnate or almost
twice pinnately incised, lobes 8-12 mm long and 0.5-1.0 mm wide,
simple, linear or narrowly lanceolate-linear, flat, short-acuminate, or
with occasional more or less long teeth; cauline leaves glabrous, short-
petiolate and sessile, once or twice pinnately incised, lobes long,
narrowly filiform, obtuse; uppermost bracteal leaves small, simple,
filiform. Capitula numerous, approximate, drooping, pedunculate,
broadly ovoid, 1.3-2.0 mm long and 1.0-1.5 mm in dia, on branches,
in secund, dense, drooping racemes forming more or less lax panicle.
Involucral bracts glabrous, smooth, with wide scarious border; outer
bracts oval, acute; inner bracts larger, elliptical. Peripheral florets
pistillate, 3-5, their corollas narrowly tubular, with 2 teeth; stigma
lobes narrowly linear, straight; disk florets staminate, 5—7, their corollas
narrowly conical; anthers on short filaments, linear, apical appendages
of anthers acute, basal appendages short, round, stigma of abortive
pistil with 2 erect lobes. Achenes 0.7 mm long, ovoid, somewhat flat,
with attenuate apex, dark brown, finely ribbed. Flowering August. (Plate
XXX, Fig. 2).

Riverbanks and marine coastal areas in forest zone.—Far East:
Uda River Area, Ussuri, Sakhalin. General distribution: Japan, north-
ern China. Described from Pescadores Islands. Type in Uppsala.

Series 6. Commutatae Poljak.—Perennials; leaves divided into
lanceolate-linear, narrowly linear, or linear-filiform lobes; inflorescence
paniculate.

105. A. commutata Bess. in Bull. Soc. Nat. Mosc. VIII (1835)
70; DC. Prodr. VI, 98; Ldb. Fl. Ross. II, 567 p. p.; Turcz. Fl. baic.-
dahur. II, 2, 51; Kryl. Fl. Alt. III, 635 p. p.; Krascheninn. in Fl. Yugo-
Vost. Evrop. Ch. SSSR, VI, 352; Kryl. Fl. Zap. Sib. XI, 2774.—A.
commutata var. helmiana Bess. op. cit. 695; Kom. and Alis. Opred.
Rast. Dalnevost. Kr. II, 1036.—A. commutata var. gebleriana Bess.
op. cit. 70; DC. Prodr. VI, 98; Ldb. Fl. Ross. II, 567; Kryl. Fl. Alt. III,
635.—A. campestris Maxim. Pr. Fl. Amur. (1859) 158.—A. desertorum
var. sprengeliana Bess. f. helmiana (Bess.) Pamp. and f. gebleriana
(Bess.) Pamp. in Nuov. Giron. Bot. Ital. n. s. XXXIV (1927) 650.

352

524

—A. pubescens Ldb. in Mém. Acad. Sc. Pétersb. V (1812) 568; Bess.
op. cit. 67; Ldb. op. cit. 565:—A. dolosa Krasch. in Sistemat. Zam.
Gerb. Tomsk. Univ. 1, 2 (1949); Kryl. Fl. Zap. Sib. XI, 2775.—A.
borealis var. mertensii Bess. and A. borealis var. ammaniana Bess. in
herb. quoad pl. altaic.—A. monostachya Bge. nom. in herb.—
Oligosporus affinis Less. in Linnaea, IX (1835) 191.—Exs.: GRF Nos.
3182, 3285 (pro var. gebleriana), 3183, 3286 (pro var. helmiana),
4048 (pro var. pubescens).—Ic.: Fl. Yugo-Vost. Evrop. Ch. SSSR,
Fig. 684.

Perennial. Tap root, woody, to 1.5 cm thick. Stems few or solitary,
35-60(70) cm high, herbaceous, erect, slender, reddish-brownish,
glabrous or hairy, leafy. Leaves glabrous or hairy beneath when young;
leaves of sterile shoots, basal and lower cauline leaves long-petiolate,
often 4—8(12) cm long and 1.5-—2.0 cm wide, twice pinnately incised,
with linear or linear-lanceolate, acute, 1-3 cm long and 0.5—1.5(2.0)
mm wide secondary lobes; middle cauline leaves simple pinnate, sessile;
upper bracteal leaves simple, narrowly linear. Capitula on short or
elongated peduncles, oblong or broadly ovoid, 3.0—3.5 mm long and
(1.5)2.0-3.0 mm in dia, divergent or drooping, on branches in lax
racemes (forming a narrow panicle) or sometimes in more or less
broadly paniculate inflorescence. Involucral bracts glabrous, outer
involucral bracts broadly oval, subacute, inner bracts elliptical, with
wide scarious margin. Receptacle convex, glabrous. Peripheral florets
pistillate, fertile, 10-15, their corollas narrowly tubular, to 15 mm
long, with 2 teeth; stigma lobes filiform-linear, exserted from tube,
divergent; disk florets 13-15, staminate, their corollas tubular-conical,
glabrous; anthers on short filaments, linear, apical appendages of anthers
subulate, basal appendages short, subobtuse; stigma of abortive pistil
with 2 short, narrowly linear, divergent lobes. Achenes 1 mm long,
oblong-ovoid, indistinctly finely ribbed, blackish-brown, with flat round
edge. Flowering August. (Plate XXVIII, Fig. 4).

Alkaline steppe meadows, clayey herb-steppe slopes, low mountains
and [mud]volcanic cones, sometimes in open birch groves and pine
forests, in forest-steppe zone and in adjoining regions of forest and
steppe zones.—European part: Volga-Kama; Western Siberia: Upper
Tobol, Irtysh, Altai, Ob River Area; Eastern Siberia: Angara-Sayans,
Yenisei, Dauria, Lena-Kolyma; Far East: Zeya-Bureya. General dis-
tribution: Mongolia. Described from southern Urals? Type in Kiev.

Note. The following varieties can be distinguished:

1. Var. helmiana Bess.—Stem solitary, slender, 1-2 mm thick;
panicle narrow, with small, drooping, globose, capitula 1.5—2.0 mm in
dia.

553

525

2. Var. gebleriana Bess.—Stems thicker, 2-4 mm thick; plant
glabrous or weakly pubescent, with large (2.5-4.0 mm long) ovate,
upright capitula.

3. Var. dolosa (Krasch.) Poljak.—Similar to var. helmiana Bess.,
with capitula up to 3 mm wide, in narrow, racemose-paniculate
inflorescence, with stems 8-30 cm high, distributed in southeastern
Altai.

4. Var. pubescens (Ldb.) Poljak.—With rather densely hairy leaves
and capitula to 2 mm in dia; eastern Siberia.

106. A. campestris L. Sp. pl. (1753) 846 and (1763) 1185; Willd.
Sp. pl. II, 1827; Bess. in Bull. Soc. Nat. Mosc. VIII, 40; Ldb. FI.
Ross. II, 565; Boiss. Fl. or. III, 363; Kryl. Fl. Alt. III, 634 p. p.;
Fedtsch. Perech. Rast. Turk. IV, 191 p. p.; Rydb. North Am. FI. 34,
254; Hegi, Ill. Fl. IV, 2, 666 p. p.; Grossh. Fl. Kavk. IV, 142; Vilda
vaxter i Nord. ut Lagerberg, IV, 1641; Poljak. in Majevski, Fl. 585.
—A. campestris var. bottnica Hartm. Handbook Scand. FI. XI (1897)
6.—A. commutata var. bottnica Krasch. in Mat. po Istor. Fl. i Rast.
SSSR, II (1946) 152.—A. borealis ssp. bottnica (Lundstr.) Hult. Atlas
over vaxter utbr. (1950) tab. 1721.—A. inodora auct. non Mill.; MB.
Fl. taur.-cauc. II, 245; Fedtsch. Perech. Rast. Turk. IV, 192;
Krascheninn. in Fl. Yugo-Vost. Evrop. Ch. SSSR, VI, 351.—A.
marschalliana Spreng. Syst. veg. III (1826) 496; Grossh. Fl. Kavk.
IV, 142; Kraschennin. in Spisok Rast. Gerb. Fl. SSSR, XI, 46;
Krascheninn. in Kryl. Fl. Zap. Sib. XI, 2772; Grossh. Opred. Rast.
Kavk. 465.—A. sericophylla Rupr. in Beitr. Pflanzenk. Russ. Reich. II
(1845) 41; Perf. Fl. Sev. Kraya, III.—A. sosnovskyi Krasch. in Grossh.
Opred. Rast. Kavk. (1949) 465 nom.—A. sosnovskyi Krasch. ex
Novopokr. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XI (1949)
178.—A. eldarica Rzazade in Izv. Akad. Nauk Azerb. SSR, 3 (1955)
22.—A. araratica Krasch. ex Novopokr. op. cit. 179.—Oligosporus
campestris (L.) Cass. in Dict. Sc. Nat. XXXVI (1826) 25.—Exs.: GRF
No. 1025 (sub A. inodora), 3287a, b, c, (sub A. marschalliana Spreng.).

Perennial. Whole plant sparsely covered with short, semiappressed
hairs, sometimes almost glabrous; root vertical, woody, with sterile,
leafy shoots. Flowering shoots strong, (20)50—65 cm high, usually erect,
ribbed, brown or slightly reddish, branched. Leaves of sterile shoots
and lower cauline leaves long-petiolate, (3)4—8(10) cm long, twice or,
less often, thrice pinnately incised, lobes narrowly linear, most often
3-10(20) mm long, subacuminate; middle and upper cauline leaves
sessile, more simply divided, usually pinnately cut or divided into
3-7 lobes, uppermost leaves bracteal, small, narrowly linear, sometimes
with 2-4 lobules at base. Capitula numerous, erect, sessile, ovate,

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526

1.5-—3.0(4.0) mm long and 1.5—3.0 mm in dia, crowded on branches in
more or less dense or lax spikes forming elongated, rather narrow,
conical-paniculate inflorescence. Outer involucral bracts oval, almost
round, convex, green on outer side, glabrous or hairy; inner bracts
slightly larger, oblanceolate-oblong, smooth, glabrous, with wide
scarious border. Peripheral florets pistillate, fertile, 7-9, their corollas
narrowly tubular, with 2 teeth; stigma lobes exserted from corolla tube,
narrowly linear, erect or weakly divergent; disk florets staminate, 7—
9, their corollas conical, glabrous; anthers on short filaments, lanceolate-
linear, apical appendages of anthers obtuse, basal appendages short,
subacute, stigma of abortive pistil with 2 short, erect lobes. Achenes
to 1 mm long, ovoid, terete, dark brown. Flowering August.

Steppe meadows, sandy steppe, pine forests, weed in old fields,
along river valleys on terraces, forest, steppe and forest-steppe zones
and rubbly mountain slopes of.—European part: Ladoga-Ilmen, Dvina
Pechora, Upper Dnieper, Upper Volga, Volga-Kama, Upper Dniester,
Middle Dnieper, Volga-Don, Trans-Volga, Lower Volga, Lower Don,
Black Sea Region, Bessarabia, Crimea; Caucasus: Eastern and Southern
Transcaucasia, Dagestan, Talysh; Western Siberia: Upper Tobol, Irtysh,
Altai; Soviet Central Asia: Aralo-Caspian Region, Lake Balkhash Region.
General distribution: Atlantic Europe, Central Europe, Balkans-Asia Mi-
nor, North America. Described from Western Europe. Type in London.

Note. Based on a study of the variation, the following varieties
can be distinguished:

1. Var. marschalliana (Spreng.) Poljak.—With highly crowded
spicate capitula on branches forming a rather narrow, spicate-paniculate
inflorescence; distributed in Kazakhstan steppe.

2. Var. sericophylla (Rupr.) Poljak.—Distinguished by dense
sericeous pubescence; distributed on sand dunes at the mouth of the
Northern Dvina River.

3. Var. sosnovskyi (Novopokr.) Poljak.—Distinguished by smaller,
subglobose capitula and somewhat elongate branches; distributed in
western Caspian Region (Dagestan, eastern Transcaucasia).

4. Var. araratica (Novopokr.) Poljak.—With small, narrowly ovate
capitula, glandular or hairy involucral bracts, and narrowly linear leaf
lobes; found in Armenia.

Economic Importance. According to the data of L. Kazakevich
and O. Sobolevskaya (1928), the yield of essential oil from the air-
dried plant matter is 0.028—0.07%. Oil constants: D?°—0.8696; acid
number—S.59; ester number—16.77; ester number after acetylation—
57.49.

According to I.V. Larin (1956), var. marschalliana is a plant of
average quality in regard to its fodder value.

555

527

107. A. tomentella Trautv. in Bull. Soc. Nat. Mosc. XXXIX, 1
(1866) 351; Fedtsch. Perech. Rast. Turk. IV, 192; Krascheninn. in
Kryl. Fl. Zap. Sib. XI, 2772.—A. inodora MB. ssp. tomentella (Trautv.)
Krasch. in herb.—Exs.: GRF No. 3186.

Perennial. Tap root sometimes branched, many-headed in old
specimens, to 3 cm thick. Stems few or numerous, 35—60(70) cm high,
at base woody, densely covered with fine white hairs. Leaves sericeous-
gray or pubescence absent (var. subglabra Krasch.), lower leaves
petiolate, middle and upper leaves sessile; all leaves oblong, once or
almost twice pinnately incised, their lobes broadly linear or linear-
lanceolate, 0.5—3.0 cm long and 0.3-1.5 mm wide, their terminal lobe
3-parted; uppermost leaves simpler 3—5 lobed, with 1-3 linear auricles
on each side; bracteal leaves linear, sometimes with 1-2 secondary
lobes at base. Inflorescence oblong, rather narrow panicle. Capitula
short-pedunculate, oblong-ovoid, 2.5—3.0 mm long and 1.5 mm in dia,
on main branches in short racemes or head. Involucral bracts oval,
with scarious margin; outer bracts hairy; inner bracts glabrous.
Peripheral florets pistillate, 7, their corollas narrowly tubular, punctate-
glandular; stigma lobes linear, erect, or divergent; disk florets staminate,
5, their corollas conical; anthers lanceolate-linear, their apical!
appendages round, long-acuminate, basal appendages short, subobtuse,
stigma of abortive pistil not lobed, funnel-shaped. Flowering August.

Sands at northern edge of desert zone.—Soviet Central Asia: Aralo-
Caspian Region, Lake Balkhash Region. Endemic. Described from Chu
and Sarysu rivers. Type in Leningrad.

108. A. bargusinesis Spreng. Syst. veg. III (1826) 493; Kryl. Fl.
Zap. Sib. XI (1949) 2776.—A. brealis wildenovii Bess. in Linnaea,
XV (1841) 96.—A. pallasii Willd. in herb. No. 15342.

Perennial. Root woody, distinctly thickened in upper part, with
fascicles of basal leaves. Stem solitary, erect, glabrous, weakly ribbed,
yellowish-green, stem 40-50 cm high. Leaves usually glabrous or
grayish-pubescent from appressed hairs, long-petiolate; lower cauline
leaves 10-15 cm long and 2-3 cm wide, twice pinnately incised,
secondary lobes 10-15 mm long and 1-2 mm wide, linear-lanceolate,
short acuminate, spinescent or falcate; upper cauline leaves simpler,
pinnately cut; bracteal leaves pinnate, ternate, or undivided, often
distinctly exceeding inflorescence. Capitula broadly ovate, 4-5 mm
long and 3—4 mm in dia, with flat broad base, erect, on short peduncles
on lower appressed branches, but upper capitula borne along entire
main axis of inflorescence on long peduncles in narrowly oblong-
paniculate inflorescence. Involucral bracts glabrous, lustrous, with white
scarious border; outer bracts smaller, oval, acuminate, inner bracts

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528

elliptical or lanceolate, subobtuse. Peripheral florets pistillate, fertile,
15, their corollas narrowly tubular, with 2 teeth; stigma lobes narrowly
lanceolate, subacuminate, divergent; disk florets staminate, numerous
(25-27), their corollas narrowly conical; anthers on short filaments,
linear, apical appendages of anthers oblong, acute, basal appendages
short, obtuse; stigma of abortive pistil not lobed. Achenes 1.5 mm
long, narrow, ovate, somewhat flat, apically attenuate, finely ribbed,
brown. Flowering August.

On mountain and steppe slopes, gravel beds in forest and alpine
tundra zones.—European part: Volga-Kama (southern Urals); Western
Siberia: Altai; Eastern Siberia: Dauria. Endemic. Described from
Barguzin District. Type in Berlin.

109. A. ledebouriana Bess. in Bull. Soc. Nat. Mosc. VIII (1835)
39; DC. Prodr. VI, 99; Ldb. FI. Ross. II, 562.—A. borealis o. ledebouri
Ldb. ibid. 567.

Whole plant with dense, long, semierect, white, hairs, later
sometimes subglabrous. Root woody, vertical, rather thick, with
perennial, woody, approximate, erect shoots covered with remnants of
leaf petioles and with rosettes of leaves. Stems 2-3 or solitary, 20-
35(40) cm high and up to 3 mm thick, strong, straight, brownish,
simple, less often branched, leafy. Basal leaves and leaves of sterile
shoots petiolate, 4-6 cm long, twice pinnately incised, terminal lobes
lanceolate-linear, most often about 1.5 mm wide, acute; middle cauline
leaves short-petiolate or sessile, with pinnately cut auricles at base;
lamina pinnately cut, upper bracteal leaves simple, short, narrowly
linear. Capitula pedunculate, subglobose, 3—4 mm long, in lax racemes
on short branches forming short pyramidal panicle. Involucral bracts
oval, with wide scarious margin; outer bracts densely hairy; inner bracts
glabrous. Peripheral florets pistillate, 9, their corollas narrowly tubular,
with 2 teeth, broadened at base, in upper part purple-violet; stigma
lobes exserted from corolla tube, narrowly linear, obtuse, recurved;
disk florets staminate, 16-18, their corollas narrowly conical; anthers
on shorter filaments, linear-lanceolate, apical appendages of anthers
acute, basal appendages considerably shorter, subacute; stigma of
abortive pistils funnel-shaped, not lobed. Achenes to 2 mm long, oblong-
ovoid, attenuate at apex, finely ribbed, blackish-brown. Flowering
August. (Plate XXIX, Fig. 2).

Sandy coasts.—Eastern Siberia: Dauria (Lake Baikal) Endemic.
Described from eastern Siberia. Type in Leningrad.

110. A. pannosa Krasch. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser.
1, No. 3 (1936) 347.

529

Perennial. Plant sericeous-shaggy from long appressed hairs. Stems
almost simple, 15-28 cm high. Basal and lower cauline leaves 4-7 cm
long, long-petiolate, petioles winged at base, lamina broadly oblong or
broadly ovate, with 1-2 lobes on each side, lobes undivided, lanceolate
or broadly lanceolate or pinnately divided, broadly cuneate at base;
secondary lobes 3-7 mm long, linear or broadly lanceolate, short-
acuminate, undivided or toothed; sometimes basal leaves also undivided,
lobate or toothed; upper cauline leaves sessile, pinnately divided, with
linear lobes, apical leaves undivided, linear. Capitula broadly ovate or
subglobose, 2-3 mm in dia, on short branches, in narrow, long (10-15
cm long) panicle. Involucral bracts bulged, with scarious border; outer
bracts oval or elliptical, densely hairy; inner bracts oblong-lanceolate
or narrowly elliptical, weakly hairy, sometimes glabrous. Peripheral
florets pistillate, 10, obconical, their stigma lobes linear, divergent;
disk florets staminate, 8-9, their corollas conical; anthers lanceolate-
linear, apical appendages of anthers acute, basal appendages short,
subobtuse; stigma of abortive pistils funnel-shaped, not lobed. Achenes
to 2 mm long, flat, ovate, attenuate at apex, brownish-black. Flowering
August to September. (Plate XXVIII, Fig. 2).

Seacoasts and stony slopes.—Far East: Ussuri. Endemic. Described
from coast of Sea of Japan between Vladivostok and mouth of Amur
River. Type in Leningrad.

Series 7. Boreales Krasch. ex Poljak.—Tussocky perennials;
inflorescence simple, spicate or narrow, racemose-paniculate.

111. A. borealis Pall. Reise, III, Anhanf. (1776) 735, tab. H, h,
f, 1; Willd. Sp. pl. III, 1839; Bess. in Bull. Soc. Nat. Mosc. VIII, 80;
DC. Prodr. VI, 98; Ldb. Fl. Ross. II, 567 p. p.; Maxim. Pr. Fl. Amur.
158; Maxim. in Bull. Acad. Sc. Pétersb. VIII, 524; Kryl. Fl. Alt. III,
636; Kom. Fl. Manchzh. III, 659; Nakai, Fl. Korean. 33; Rydb. North.
Am. FI. 34, part 3, 256 p. p.; Pampan. in Nuov. Giorn. Bot. Ital. n. s.
XXXIV, 641; Kom. Fl. Kamch. III, 146; Kom. and Alis. Opred. Rast.
Dalnevost. Kr. II, 1035; Krascheninn. in Kryl. Fl. Zap. Sib. XI, 2777.—
A. violacea Ldb. in Mém. Acad. Sc. Pétersb. V (1812) 567; Ldb. FI.
Alt. IV, 78.—A. borealis var. purshii Bess. in Hook. Fl. Bor. am. I
(1833) 326; Bess. op. cit. 80.—A. borealis Pall. var. spithamaea (Pursh)
Torrey and Gray, Fl. North Am. FI. 2 (1843) 417; Krascheninn. in
Spisok Rast. Gerb. Fl. SSSR, X, 96.—A. spithamaea Pursh, Fl. Amer.
Sept. II (1814) 522; Rydb. op. cit. 255.—A. campestris ssp. spithamaea
Hall and Clements. The Phylog. Method (1923) 123.—A. campestris
ssp. borealis Hegi, Ill. Fl. VI, 2 (1929) 669.—A. remosa Sugaw. PI.
of Saghal. (1934) 364.—A. camtschatica Schlechtd. in Herb. Berol.—
Exs.: GRF No. 3185a, b.

558

530

Perennial. Root thick (to 1.5 cm ), many-headed, with very short,
erect, approximate perennial shoots, covered with thickened bases of
leaf petioles and with rosettes of leaves forming rather compact, small
tussocks. Stems few or solitary, 8—25(35) cm high, erect or somewhat
bent, simple or branched, more or less hairy or subglabrous, sometimes
violet. Leaves entire, densely hairy at base, on both sides or only
beneath, later glabrescent; lower cauline leaves and leaves of sterile
shoots long-petiolate, 2—-6—10 cm long, twice or almost thrice pinnately
incised, lobes lanceolate-linear or linear, 2-10 mm long and 0.5-1.5
mm wide, short-acuminate; middle cauline leaves less divided, often
simple pinnate or palmate or sometimes even undivided, linear; apical
leaves bracteal, usually undivided, linear, considerably exceeding
inflorescence and short lateral branches. Capitula subglobose, 3-4 mm
in dia, borne directly on main stem or on peduncles, gradually longer
downward forming simple raceme or even compact and strongly
shortened racemes on lateral branches of narrow racemose or partly
racemose-paniculate inflorescence. Involucral bracts densely hairy or
glabrous; outer bracts oval, usually brownish-purple, inner bracts
somewhat larger, broadly oval, with white scarious margin. Peripheral
florets pistillate, 15-17, their corollas small, tubular; stigma lobes linear,
obtuse, divergent; disk florets staminate, 17-20, their corollas narrowly
conical, glabrous or in upper part hairy, often purple, anthers linear,
apical appendages of anthers acute, rather large, basal appendages
very short, subobtuse; stigma of abortive pistil funnel-shaped, not lobed.
Achenes about 2 mm long, oblong-ovate, somewhat flat, blackish-brown.
Flowering August. (Plate XXIX, Fig. 3).

Dry sandy banks, well-drained river terraces, slopes of uplands in
arctic zone, as well as stony and rubbly slopes of mountains in alpine
zone, descending to limits of forest zone.—Arctic: Arctic Europe,
Novaya Zemlya, Arctic Siberia, Chukotka, Anadyr; Western Siberia:
Altai; Eastern Siberia: Angara-Sayans, Yenisei, Dauria, Lena-Kolyma;
Far East: Sakhalin, Kamchatka. General distribution: Scandinavia,
North America. Described from lower reaches of Ob River. Type in
London.

Note. This species is characterized by great variation in the pu-
bescence, whereby young plants are often whitish from an abundance
of hairs, but older plants are often subglabrous; the inflorescence is
sometimes paniculate or spicate, when the capitula are more or less
crowded on very short peduncles. The degree of leaf cutting also varies;
the apical leaves are either undivided or partly 3-lobed or even
pinnatisect. It is possible to distinguish:

559

531

1. Var. ledebouri Bess.—distinguished by pinnately cut basal leaves
and simple racemose inflorescence;

2. Var. mertensii Bess.—with twice or thrice pinnately incised
basal leaves and capitula that are crowded in lax panicle with longer
branches.

112. A. henriettae Krasch. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, IX (1946) 175.

Perennial. Root woody, rather thick, with strongly shortened
perennial shoots bearing rosettes of leaves forming compact velutinous
tussocks scarcely 5—8 cm in dia with short (1-3(7) cm high), erect
stems. Leaves small, covered with long, usually dense hairs; petioles
thickened at base; lamina pinnatifid, lobes 2—3 on each side, small, 2—
3 mm long, fleshy, linear or lanceolate, subacute. Capitula globose, 4—
5 mm in dia, in dense, terminal head. Involucral bracts oval, pilose,
with broadly scarious margin. Receptacle convex, glabrous. Peripheral
florets pistillate, fertile, 17-32, their corollas narrowly tubular or
narrowly goblet-shaped-tubular with 3-5 teeth, in upper part purple-
violet, glabrous or weakly hairy; stigma lobes exserted from corolla
tube, linear-up-right; disk florets (12-19) staminate, their corollas
narrowly goblet-shaped-conical with 5 teeth, purple-violet, anthers
lanceolate-linear, apical appendages round, acuminate, basal
appendages short, round; stigma of abortive pistil funnel-shaped,
indistinctly lobed. Flowering August.

Well-drained, clayey slopes in arctic zone.—Arctic: Novaya
Zemlya, Arctic Siberia (islands of Kara Sea, De Long islands). En-
demic. Described from Henriette Island. Type in Leningrad.

113. A. limosa Koidz. in Sugaw. Pl. Saghal. (1937) 303; Honda
Nom. pl. Japon. 342; Sugaw. Ill. Fl. Saghal. IV (1940) 1825.—Ie.:
Sugaw. ibid. tab. 824.

Perennial. Root slender, vertical, with solitary, perennial, very short
shoots covered with leaf petioles and bearing rosette of leaves. Stem
solitary, 20-28 cm high, erect, glabrous, brown or violet-brown, sparsely
leafy, simple or branched, with more or less short, upright, flowering
branches. Leaves glabrous, smooth; lower cauline leaves and leaves of
sterile shoots 2-3 cm long, long-petiolate with petioles strongly
thickened at base, lamina oblate or almost round, twice pinnately cut,
lobes linear or narrowly lanceolate-linear, often 3-7 mm long, acute;
middle cauline leaves less compound, uppermost leaves bracteal, short,
simple, narrowly linear. Capitula broadly ovoid, 4-5 mm long, upright,
crowded, on more or less long peduncles, borne on main stem or in
depauperate racemes on branches, forming narrow, linear, racemose-

560

Sa2

paniculate inflorescence. Involucral bracts glabrous, with wide scari-
ous margin; outer bracts oval or elliptical, inner bracts considerably
larger, narrowly elliptical or narrowly oblanceolate.

Mud volcanos.—Far East: Sakhalin? Endemic. Described from
southern Sakhalin. Type in Japan.

Note. The available material is insufficient and precludes the
possibility of describing the flowers. The original description by the
author of the species is incomplete.

114. A. pycnorhiza Ldb. Fl. Alt. [IV (1833) 79; Bess. in Bull.
Soc. Nat. Mosc. VIII, 77; DC. Prodr. VI, 98; Ldb. Fl. Ross. II, 567;
Kryl. Fl. Alt. II, 636; Kryl. Fl. Zap. Sib. XI, 2775.—A. borealis var.
ledebourii Fedtsch. Perech Rast. Turk. IV (1911) 192.—A. depauperata
Krasch. in Sistemat. Zam. Gerb. Tomsk. Univ. 1-2 (1949); Kryl. FI.
Zap. Sib. 2777.—Ic.: Ldb. Ic. pl. Fl. Ross. V, t. 473.

Perennial. Root 1.0-1.5 cm thick, woody, many-headed, with few
strongly reduced, thick, perennial shoots (with rosette of leaves),
forming rather dense tussocks. Fertile stems few, (5)10-18(25) cm
high, erect, simple, hairy, later subglabrous, stramineous. Leaves densely
covered with semiappressed long hairs; basal and lower cauline leaves
petiolate, 2-6 cm long and 1-2 cm wide, ovate or lanceolate, once or
twice pinnately incised, lobes 3-6 mm long and 1.0-1.5 mm wide,
lanceolate, subulately spinescent; middle and upper leaves sessile,
smaller and more or less divided to simple pinnate; uppermost leaves
undivided, usually not rising above lateral branches of inflorescence.
Capitula broadly ovoid, 2.5—4.0 mm in dia, upright, in simple racemose-
spicate or racemose-paniculate inflorescence; lower capitula often
remote on rather long peduncles, but upper ones sessile. Involucral
bracts sparsely hairy or glabrous, smooth; outer bracts oval, acute;
inner bracts broadly elliptical or almost round, obtuse or somewhat
acuminate with scarious border, and brownish longitudinal stripe in
upper part. Peripheral florets pistillate usually 15, their corollas narrowly
tubular, often enlarged at base; stigma lobes narrowly linear,
subacuminate, upright; disk florets staminate, 10-15, their corollas
conical, punctate-glandular, glabrous, sometimes with sparse hairs;
anthers on short filaments, linear, apical appendages acute, basal
appendages short, subacute; stigma of abortive pistil short, linear, not
lobed. Achenes 1.0-1.2 mm long, ovoid, somewhat flat, angular-ribbed,
dark brown. Flowering July to August.

Stony steppe slopes and valleys, upper part of mountains or in
alpine zone.—Western Siberia: Altai (south-east); Soviet Central Asia:
Dzhungaria-Tarbagatai. General distribution: Mongolia. Described from
Altai. Type in Leningrad.

56

ot

533

Note. In Tarbagatai and part of the Altai, we find var. depauperata
(Krasch.) Poljak. with larger capitula, partly in a spicate arrangement,
and bracteal leaves that project from the inflorescence.

Series 8. Scopariae Krasch.—Annuals or biennials; leaves divided
into narrowly filiform or filiform lobes; capitula ovoid.

115. A. scoparia W. and K. PI. rar. Hung. I (1802) 66, t. 65; Ldb.
Fl. Alt. IV, 87; Bess. in Bull. Soc. Nat. Mosc. VIII, 92; DC. Prodr. VI,
99; Ldb. FI. Ross. II, 569; Boiss. Fl. or. III, 364; Hook. Fl. Br. Ind.
322; Kryl. Fl. Alt. III, 637; Fedtsch. Perech. Rast. Turk. IV, 192;
Hegi, Ill. Fl. VI, 2, 670; Grossh. Fl. Kavk. IV, 139; Krascheninn. in
Fl. Yugo-Vost. Evrop. Ch. SSSR, VI, 353; Kryl. Fl. Zap. Sib. XI,
2778; Poljak. in Majevski, Fl. 584.—A. scopariaeformis M. Pop. in
Pochv. Eksp. Bass. rr. Syr-Dari i Amu-Dari, 1 (1915) 50.—A.
scoparioides Grossh. in Tr. po Geob. Obsl. Pastbishch Azerb. 2 (1929)
69; Grossh. Fl. Kavk. IV, 139; Grossh. Opred. Rast. Kavk. 463.—A.
paniculata Bess. in Mém. Acad. Sc. Pétersb. V (1945) 471, non Lam.—
Oligosporus scoparius (W. and K.) Less. in Linnaea, IX (1834) 191.—
Exs.: GRF Nos. 623, 3177a, b.

Annual or biennial. Root slender, straight, vertical. Stem 30-70
cm high, solitary, less often 2-3, erect, pubescent, later glabrous,
reddish-violet or brown, strongly branched in middle and upper parts,
with divergent branches. Young leaves pubescent, older ones glabrous;
lower leaves petiolate, twice or thrice pinnately incised into linear-
lanceolate-acute lobes, withering before anthesis; middle cauline leaves
sessile, 1-4 cm long, with narrowly linear or filiform-linear lobes.
Capitula small, 1.5-2.0 mm long, ovate or broadly ovate, on short
peduncles, divergent or drooping, on branches in secund racemes
forming pyramidal panicle. Involucral bracts glabrous, with scarious
margin; upper bracts oval, inner larger, green, oblong or broadly
lanceolate, glabrous, glaucous. Peripheral florets pistillate, 6, their
corollas narrowly tubular, stigma lobes narrowly linear, acuminate,
divergent; disk florets 6, staminate, their corollas conical, anthers linear,
stigma of abortive pistil not lobed. Achenes 0.6 mm long, ovoid,
somewhat flat, finely ribbed, brown. Flowering August.

Alkaline steppe meadows, light sandy loamy and sandy soils, as
weed in fields, pastures, old fields, open pine groves, and also in
irrigated fields, along river valleys on rubbly slopes, in forest-steppe
and desert zones and also in lower mountain zone in steppes.—
European part: Karelia-Lapland, Dvina-Pechora, Ladoga-Ilmen,
Upper Dnieper Baltic Region, Upper Volga, Volga-Kama, Volga-Don,
Upper Dniester, Middle Dnieper, Bessarabia, Black Sea Region, Lower

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534

Don, Lower Volga, Trans-Volga, Crimea; Caucasus: Ciscaucasia;
Eastern, Western and Southern Transcaucasia, Dagestan, Talysh; West-
ern Siberia: Upper Tobol, Irtysh, Altai, Angara-Sayans, Lena-Kolyma,
Dauria; Soviet Central Asia: Aralo-Caspian Region, Lake Balkhash
Region, Kyzyl-Kum, Kara-Kum, Syr-Darya, Amu-Darya, Tien Shan,
Pamiro-Alai Region. General distribution: Atlantic and Central Europe,
Balkans-Asia Minor, Armenia and Kurdistan, Iran. Described from
Hungary. Type in Vienna.

Note. One can distinguish var. scopariiformis (M. Pop.) Poljak.
with long, strongly divergent branches and a small number of flowers
(pistillate florets 1-2, staminate disk florets—2—4); found in the deserts
of Uzbekistan and Turkmenia.

Economic Importance. The above-ground parts of the plant con-
tain 0.09-1.1% essential oil. According to the data of L. Kazakevich
and O. Sobolevskaya (1928), the oil from a plant collected in the
Astrakhan Region had the following constants: D—0.913; n,”°—1.49;
acid number —3.32; ester number—14.50; ester number after acetyla-
tion—43-45. The oil contained up to 4% a—pinene, up to 30% B—
pinene, about 10% myrcene. According to the data of A. Katz (1938),
the oil contains up to 3% aldehydes and ketones and 1.5% phenols.

According to the data of V.A. Vyshensky (1930), the oil from a
plant collected in Turkmenia (probably var. scopariformis (M. Pop.)
Poljak.) had the following constants: D—0.8695; a—1.3°; n—1.4894;
acid number—0.16; ester number—32.3; ester number after
acetylation—53.17. The oil contained apiol. In Vyshensky’s opinion,
the oil is of little industrial interest, but, according to M.I. Goryaev
(1956), it can be used in perfumery.

116. A. demissa Krasch. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser.
1, c, (1936) 348.

Annual or biennial. Root slender, vertical. Stem 5-20 cm high,
violet, sometimes greenish, more or less densely hairy, basally branched,
lower branches prostrate in lower part, ascending above, usually
arcuately bent, fertile. Leaves covered with long appressed, white hairs;
basal and lower cauline leaves petiolate, with expanded semiamplexicaul
petioles, 1.5-2.0 cm long and 1.0—1.5 cm wide, lamina broadly oblong
or broadly ovate, twice pinnately incised, lobes 2-4 mm long, linear-
lanceolate, with short, stiff, acuminate apices; upper cauline and apical
leaves sessile, 5-10 mm long, simple pinnate, with linear lobes. Capitula
numerous, erect, sessile or on short peduncles, ovoid-campanulate, 2
mm long and 1.5 mm in dia, remote, in lax racemose-paniculate
inflorescence. Involucral bracts glabrous, glaucous, convex dorsally,
greenish, with more or less wide scarious border, pale, sometimes

564

535

violet, outer bracts oval-deltoid, inner bracts broadly lanceolate.
Peripheral florets pistillate, rather numerous (19), their corollas narrowly
tubular, with one very short tooth, stigma lobes filiform-linear,
acuminate, divergent; disk florets staminate, usually 5, their corollas
narrowly conical, anthers linear; stigma of abortive pistils not lobed,
funnel-shaped. Achenes 0.6—0.7 mm long, flat, ovoid, finely ribbed,
brown. Flowering August. (Plate XXIX, Fig. 1).

Sandy coasts and alpine deserts, sometimes near irrigated areas.—
Soviet Central Asia: Pamiro-Alai Region (western Pamir). General
distribution: Tibet. Described from banks of Kuku-Nor River. Type in
Leningrad.

Subgenus 3. Seriphidium (Bess.) Rouy. Fl. France. VIII (1903)
298.—Seriphidium Bess., Sect. in Bull. Soc. Nat. Mosc. I (1828) 222.—
Capitula with homogamous florets; involucre small, ovate or narrowly
cup-shaped, 3—6-whorled. Involucral bracts imbricate, with scarious
margin, innermost bracts considerably larger than outer ones, partly
covering each other by their apices, hence involucre appearing like
closed receptacle. Florets 2-10, bisexual, cleistogamous, their corollas
tubular, with 5 teeth, yellow, less often pinkish; anthers lanceolate-
linear, as long as filaments. Apical appendages of anthers subulate or
narrowly linear, basal appendages very short, roundish; “antheropodia”
distinct, round; style shorter than or as long as stamens, style lobes
short, linear, upright, truncate, with short, erect ciliate hairs throughout
upper margin. Achenes flat-ovoid, sulcate; pappus absent; common
receptacle very small, conical, glabrous.

Perennial shrubs or semishrubs with woody perennial shoots at
base, or very rarely annuals, at beginning of vegetative period densely
covered with simple or bifid arachnoid hairs, later glabrescent or
glabrous; lower cauline leaves often twice or thrice pinnately incised
into narrowly linear or filiform lobes, upper leaves more simple.
Inflorescence paniculate or less often spicate-paniculate. There are
more than 100 species in nature, distributed in the Mediterranean
countries and extending up to the eastern parts of Soviet Central Asia.
In the flora of the Soviet Union, 58 species belong to this subgenus.

1. All leaves simple, entire or pinnate-short lobate, or even with 3—

UME UM AL GEEMOK ress oun ass nantMGie ibiensas cnauate ge <ssrnnghigh gh rnnessssaaMsecaves 2.
+ Lower cauline leaves ternate or twice to four times pinnately incised
00), ANAS ADIL CIM CREP URE COMER, “OER 4.
2. Lower cauline leaves with 3—5 broadly linear teeth at apex, others
DA Ng ook ent Xi eheasitenslt x> diis<-nnennsesnasé 172. A. deserti Krasch.

+ All leaves pinnately short-lobed or subentire ............:cceeeeeee 3:

536

563 Plate XXVII.

1 —Trichanthemis aulieatensis (B. Fedtsch.) Krasch.,
anethifolia Web., habit.

habit. 2 — Artemisia

4(1).

565

N +

7(4).

So +

10.

11.

537

Lateral segments of leaves simple, round, sometimes slightly elon-
gated or even divided into 2—3 small, round lobes; sometimes
leaves even entire or with broad, obtuse teeth; uppermost leaves
bracteal, small, linear. Capitula ovate, up to 5 mm long, with 3-
A floretarreol Ha. Wis ol. fat. ies 170. A. santolina Schrenk.
Lateral segments deltoid-oval or oval, with very short decurrent
rachis, simple or twice or thrice pinnately incised lobes; upper
bracteal leaves scaly, broadly linear; capitula oblong, up to 4 mm
long, with 5—6 florets ...........ccsseeeeeeee 171. A. lobulifolia Boiss.
Middle and partly lower cauline leaves ternate; pubescence of
hairs short, stiff, appressed or long, and erect ...........::cseeeeeeees 5.
Lower leaves twice or thrice pinnately cut; hairs arachnoid, flexuous

Annual or biennial, with slender, readily peeling bark; hairs dea
long, MoStly SQUAITOSE ...........ssseeeeeeeeee 174. A. leucodes Schrenk.
Perennials with thick woody root; hairs short, appressed .......... 6.
Leaves of sterile shoots and lower cauline leaves 1.5—2.0 cm long,
terminal lobes 4-6 mm long; capitula 5-6 mm long, densely
qpabecarit’ Sie hn ieee nak. PAL IA cedabensee
ere reee ere 173. A. juncea Kar. and Kir. var. macrosciadia Poljak.
Leaves larger; capitula usually less hairy, up to 4.0-4.5 mm long
jcsetaES SPOTS oe RET tase endless 173. A. juncea Kar. and Kir.
Stems at flowering glabrous, blackish-purple or deep dark brown,

indtwous:!. 23.2) Seed ete a, 154. A. turanica Krasch.
Stems lighter in color; pubescent to different degrees, less often
pOUUAST TOT RA ys BP cncccsensennssnnarrannctarswesacossescncnsers 8.

Short, up to 35 cm high, strongly branched shrub forming compact,
stiff pulvinate crown; flowering shoots almost horizontally
divergent, almost prickly at end of vegetative period ................
5» aah. ee A: Se elisha asekt ss 166. A. sieberi Bess.

Plantiwith chinracters otherwise cake. 2100.8 28. 4 EU aeadees nee evens 9.
Capitula few, clustered in usually compact, oblong or subglobose,
PST ON ATO MEOIALS SPOS) aici sestosenonponanccensncetidddroneese 10.
Capitula approximate or remote on branches but not forming
Cove expanse saree Pk eda eae ease eee ited oo onde 14.
Lower leaves round, thrice- or almost thrice pinnately cut;
SeMiabribyetoonel yoRtanChed iyi. ts. dieses. Sakis scds onan bbidaes onthe i
Lower leaves ovate, twice pinnately cut; herbaceous perennials
SUPER steht Tee ee ele, hae ont HER Lethe ened EZ.
Plant 35-45 cm high, initially densely pubescent, later partly
glabrous or subglabrous .................. 167. A. lehmanniana Bge.

Plant 12-30 cm high, whitish-tomentose up to end of vegetative
ERSRMIGIOIE, HOPE NTE WEDINTICt A Mets tecceiker secured ctstsovectecvevecsctoceesnscdenseseneaveeceses

538

18.

++

19(14).

-+-
20.

+

IAN

High-mountain (alpine) plants with strongly reduced underground,
partly aerial, woody shoots forming rather compact tussock; fertile
stems 4—10(15) cm high, with simple spicate inflorescence ......
Rial SENT. 09. Gk BNO, ROR A. 128. A. rhodantha Rupr.
Mountain plants, partly in foothills, with lax tussock and branched,
muchitaller stems J.44:..20. Nn RI aR. ARs, Le ee 13.
Fertile stems 15-35 cm high; lower cauline leaves short-petiolate,
2=3 semilonge witae2 TRO LIRR Ee 127. A. compacta Fisch.
Stems 50-60 cm high; leaves 4-5 cm long
eet. AEE Rea. BRAS, 129. A. prolixa Krasch. ex Poljak.
Inflorescence simple, spicate or spicate-racemose ................+. td:
Inflorescencepaniculatevt.2g, 220281 DOS, Se 19,
Lower leaves simple or partly almost twice pinnately cut; plant
usually S=i5:em highvs:..x2hiate Ce... Rs ae ee, 10.
Lower leaves always pinnate; plant 20-35 cm high ............... 18.
Plants 15(25) cm high, with lax, weakly developed tussock
FA occiisd. OB, SSE, PO, DAR MOONS. 132. A. korovinii Poljak.
Plants 5-15 cm high, forming compact tussock ................006 Ly.
Lower leaves once pinnately cut; capitula in simple spicate
inflorescence. 184..1541.008. 18 Bee 131. A. knorringiana Krasch.
Lower leaves once or twice pinnately cut; capitula in spicate or
FACEMIOSEVIMMIOLESCENCE -. «0c. esseirensiniuncecanbengeveneesnte codeemieeoeecennine Mace
190.262 123. A. nitrosa Web. ex Stechm. var. gobica Krasch.
Entire plant whitish-tomentose; lower leaves 2.5-4.0 mm long;
capitula up to 2.5 mm long
ot cadtndalaue eee cieten A ccepicd 130. A. dubjanskyana Krasch. ex Poljak.
Plant glaucescent-green, subglabrous; lower leaves up to 1.0—1.5
cm long; capitula usually smaller, up to 2 mm long
ve SLL. AIO, 1 168. A. vachanica Krasch. ex Poljak.
Plant tomentose, later subglabrous; lower leaves 1-3 cm long;
capitula up to-3.mmilong 9 Ret. ee 158. A. sogdiana Bge.
Lower cauline leaves once pinnately cut or in part twice pinnately
GUE Brevvectesbavedinsvisaiiasseseidavuen Quan « ame aI, et MERE cate ca 20.
Lower cauline leaves twice to four times pinnately cut .......... 29:
Inflorescence; usually narrow, branches mostly short, 1—2(3) cm
Love) €6. Se, AOEOLG EON SL RA eee 21.
Inflorescence more or less spreading, branches longer, obliquely
upright.or/almost horizontal enw. a8 Re. ess cents 24.
Plant glabrous or subglabrous; leaves divided into filiform-linear
lobes, up to 5-10 mm long .................. 136. A. lessingiana Bess.
Plant grayish, more or less densely pubescent; terminal lobes of
leaves linear or narrowly lanceolate-linear ...............:ccssccecceseees Pagde

Peewee roses eres eoeeseeeesesssoee

Pewee ere sees se rer ee esse ee esse eS EH SESE SEES SEES ES OE SEES

567

22.

24(20).

20.

28.

+

++

29(19).

ee
30.

539

Branches of inflorescence horizontally divergent or even droop-
ingsnalantgledsanthy aromatic, 55. pi2isc2..cosisehi6).-16i0h GEGEN G..0sseerereee
siihicaiseabinteas 135. A. terrae-albae Krasch. var. suaveolens Poljak.
Branches adpressed to stem or obliquely upright .................06 yen
Terminal lobes of leaves mostly up to 1 mm wide and 4-12 mm
long; plant whitish-tomentose up to end of vegetative period,
endemic-tov Altai jecccesiss.cic.stvtvcsesbes vesves.0ss 143. A. amoena Poljak.
Terminal lobes shorter and narrower; plant grayish-green, partly
SiR earn aa IN et cnc gciononnsicennconhanny es enciddenanppsosoeeiand
139. A. sublessingiana Krasch. ex Poljak. var. polistichum Poljak.
Stems branched from base or lower part, with long (lower branches
EpstonlO—25. crilong)virgate branche 5. i0.....:sssisaconeccerecnesneavensens
papchantcertd hes eavhss th aanderadhe whtaads ohashsvtas 142. A. kaschgarica Krasch.

Stems always branched in upper part ............ccccesseseseesseeeeeeeees 25.
Plants grayish-green, not densely pubescent, later subglabrous or
weakly pubescent, green or glaucescent-green .........:..:scsceeeeees 26.
Plants whitish-tomentose, later glabrescent ............::csceeeeeees 1

Fertile stems 30-50(65) cm high; inflorescence rather wide,
paniculate with long, rather strongly divergent branches; lower
cauline leaves 3-5 cm long, their terminal lobes up to 4.5 mm
Weinchos~ainsssi-hastanntiGit 123. A. nitrosa Web. ex Stechm.
Fertile stems 7—18(20) cm high; inflorescence oblong, paniculate,
with obliquely upward-spreading branches; lower cauline leaves
1.0-2.5 cm long; their terminal lobes 24 mm long .................
Geeks 145. A. cina Berg. ex Poljak. var. mogoltavica Poljak.
Lower leaves pinnately cut, lateral segments entire or more or less
deeply divided into 2-4 lanceolate, acute lobes; inflorescence
narrow, lax, pyramidal panicle; plants of clayey salt marshes and
allsalisndiloadss ..2a¥. 2200re get hnndseel3i.i.. 124. A. scopaeformis Ldb.
Lower cauline leaves once or twice pinnately cut, terminal lobes
narrowly linear; panicle rather wide, usually with strongly divergent
or upward-spreading branches; plants of brown-desert soils and
dete sands tierce coves. .7eoi.i0.exiol lanienes.....8 28.
Stems 12-25 cm high; capitula 3.0-4.5 cm long; involucral bracts
convex, punctate-glandular .................. 156. A. kemrudica Krasch.
Stems 15-30 cm high; capitula 2-3 mm long; involucral bracts
not strongly curved, eglandular ....... 135. A. terrae-albae Krasch.
Stems 30-40 cm high; capitula 2-3 mm long, eglandular ........
ier ae 135. A. terrae-albae Krasch. var. massagetovii Krasch.
Lower cauline leaves thrice or four times pinnately cut ........
Dish ABAUSENGE, BMG, cade ie 169. A. baldshuanica Krasch. and Zapr.
Lower leaves twice or thrice pinnately Cut ..........cccseeseesseeereees 30.
Lower leaves mainly thrice pinnately Cut ..........:ssscceseesseeereees 31.

540

jd.

32:

568 33(30).

Lower leaves partly thrice pinnately cut i.e., some primary lobes
pinnate but others twice pinnately cut or leaves twice pinnately
CUE ek BA ollie lide ARR RRA ds adebeteee ine. a.
Plant with aerial, woody, strongly reduced, perennial, sterile shoots;
inflorescence narrowly compact panicle with adpressed branches
WG SAO, . LOR AOI SRL hRLL,. MOREE TENE + 141. A. tenuisecta Nevski.
Plant with weakly developed or almost undeveloped, perennial,
woody shoots; panicle with obliquely upward-directed branches
wisactzanaasvilecaiabvonsosiirebelnas bys dehssmenqent tlt tEAM eee Nee IEE ENS CIM ants 32)
Inflorescence broadly paniculate, with rather long (up to 10-15
cm) lateral branches; capitula remote or approximate ................
sthneeBcesiapis viteses vanes MEM: RULE LOI 126. A. schrenkiana Ldb.
Panicle narrowly pyramidal, with shorter lateral branches; capitula
crowded.,on. branchesis}. 2579.85, Lees, AEA TO onc seen
TACOS. TOME MTR 117. A. maritima L. var. boschniakiana DC.
Inflorescence more or less wide pyramidal panicle with divergent
branches, «iiss: LAOReNI RR. TA Ae AG et oe 34.
Inflorescence mostly compact panicle with branches adpressed to
stem/axis/an obliquely uprights .te...22.0es, oes LS we 59:
Stems mainly 50-60 cm high, sometimes even mote .............. 35.
Stems mainlyil5—45 cm high. sivosscsoeioimcervorearr eee 41.
Lower leaves twice or thrice pinnately cut, i.e., primary lobes
partly twice or once pinnately cut; plants of Caucasus .......... 36.
Lower leaves mainly twice ae cut; plants of Soviet Central
NST ASIOINE 5 IBY. DAC A RE, UR i Nh cctessestviytioen cons Bite
Lower leaves often up to 10 cm long, terminal lobe linear, 5—10
mm longs fates... eee 119. A. szowitziana (Bess.) Grossh.
Lower leaves smaller, their lobes up to 3-5 mm long, linear-
filiformoc2..& ASS. 120. A. fragrans var. phyllostachys Boiss.
Lower leaves up to 1 cm long; capitula narrowly ovoid or narrowly
oblong i:..auw.. ea .2 159. A. stenocephala Krasch. ex Poljak.
Leaves)larger;, capitulalovoid 2c5248. 2228aiie Re. ee 38.
Terminal lobes of lower leaves narrowly linear, usually up to 7—
20: mmi long: and «0:3! mami pwide), ,4.240). ce teeee sh RTRs... oseven sen
rk) SOE 157. A. kochiiformis Krasch. and Lincz. ex Poljak.
Terminal lobes of lower leaves linear, shorter and wider ........ 39.
Capitula spicately borne on secondary branches; lower leaves up
to 3-5 cm long, terminal lobe small, up to 2-4 mm long ............
ISA RR RAAB PE. Ha aL ee EONS A 150. A. serotina Bge.
Capitula more or less remote or approximate, few in each
inflorescence; lower leaves up to 4-8 cm long, terminal lobe longer
avavscnussapgnedbanendus ceebdes dom eeOeagRe: ANSELM AUS, AEG EEN RON SUS, SRA he 40.

40.

+

41(34).

+
42.

45.

46(41).

+

47(46).

+

48(46).

+
49.

541

Terminal lobe of lower leaves up to 1.5 mm wide, all or some
With DAS heath wb POR). BO es ARI eg UT oege esenneh ennnh

SUPRA be Bd HA, AHS 148. A. ferganensis Krasch. ex Poljak.
Terminal lobes of lower leaves 0.5—1.0 mm wide, acuminate, not

tothedawulei.dannauacata 149. A. porrecta Krasch. ex Poljak.
Lower leaves twice or thrice pinnately Cut ............cscceesseeeseeeees 42.
Lawen leavesiiwick pititiately: cut ......2.....:..0sr0isesscsnorcsgaceenseeesees 46.

Capitula mainly (2.5)3.0-4.0 mm long; lower leaves 24 cm long;
bracteal leaves usually exceeding capitula; plants with perennial,
Short: creeping, istertlesShapts, 105.366 iiic isc eceaks ang libttlan baal sp sennen 43.
Capitula 2.0-2.5 mm long; lower leaves up to 1.5—-2.0 cm long,
upper bracteal leaves not exceeding capitula; plants with perennial,
reduced, woody ascending Shoots ...........ccscssecsceseersesneeeneees 44,

Capitula 3-4 mm long, with 7-10 florets; bracteal leaves usually
Sa: fe 117. A. maritima L.

Capitula 2.5 mm long, with 4—5 florets .........ccccsssesseeseeeseeneseeneneees
itachi atleast ~aleatedmtbiiengs 117. A. maritima L. var. palmata L.
Inflorescence branches obliquely upward-spreading; stems slender,
slabréscents bio Win iiss, Bee BE an Legh eden sh chan cclldd daleddh deneny oabldanenee
Lenmdenebibbe deiuin» <peinclb etd 154. A. turanica Krasch. var. pallida Poljak.
Branches almost horizontal; stems partly glabrescent, partly
pubescence till end of vegetative Peri0d ..........sssseeeeeeseeeeees 45.
Stem up to 40 cm high, at base often up to 3 mm thick, pubescent
almost to end of vegetative period; capitula more or less
approximate ON branche ...........csseseseeeeserenseeeeneeseeeesessneseeeteseaeees
rates rete a 154. A. turanica Krasch. var. diffusa (Krasch.) Poljak.
Stems low, up to 25 cm high, more slender, partly glabrescent,
weakly branched; capitula fewer, remote on branches ...............++.
amare Ae d, 154. A. turanica Krasch. var. eremophila Poljak.
Terminal lobes of leaves narrowly linear or almost filiform-linear;

capitula broadly ovate, less Often OVOIC ........sscseesesereeesetseeeees 47.
Terminal lobes linear or narrowly linear; capitula more or less
Ovate OF NALrOWLY OVALE ...........cccsssccercsscssseessseessetsceneeeseeseees 48.
Capitula broadly ovate ...........scseeeeeees 120. A. fragrans Willd
Capitula Ovate ..........ssscssssssssssseessesssssssessssensscssssscecessssnsvssssanasaneneeaeeenes
ARG AE OS 120. A. fragrans Willd. var. gurganica Krasch.
Lower leaves mainly 34 cm long or to 7 cm long ...............+5 49.
Lower leaves up to 1-2-3 cM JONG ........esseecereeereereereetsersceneeees al.

Plants with numerous, ascending, woody, perennial shoots forming
low tussocks; capitula in compact panicle, up to 3 mm long, upright
cide e RR teeksaasimsinidinwinike 151. A. valida Krasch. ex Poljak.
Plants not forming tussock; capitula in spreading or lax, usually
narrow panicle, somewhat approximate or partly remote, 3-4 mm
long, erect OF CrOOPING .............secssesseseesessseresenessseccensseesseesssees 50.

570

542

51(48).

+
Ses

+
53:

54.

55°
56.

57.

58.

59(33).

Stems initially grayish, densely tomentose, later almost glabrous,
brown, smooth; branches of inflorescence almost horizontal;
capitula pedunculate, often drooping, in lax inflorescence ..........
JOBRb. obi theh: B22 RA) eee 118. A. monogyna Waldst. and Kit.
Stems in part glabrescent, light brownish, inflorescence branches
obliquely upright in lower part; capitula sessile, erect ................
Sent aaonclicton dit seisbeen chan cee em 123. A. nitrosa Web. ex Stechm.
Capitula small, usually 2-3 mm long .........eeeeeeseseseeeeeneeeeeee 52:
Capitularlarger,) 3=4omar longe.3.4 Hate aee ee JO ce ccaae 54.
Low shrub, 12—15(30) cm high; terminal lobes of lower leaves 1—
2 mm long, acuminate ...............0 138. A. camelorum Krasch.
Plant 30-45 cm high; terminal lobes longer ........... cece 53.
Stems slender, green above, with short, more or less divergent
branches; mature capitula divergent or drooping ..............:ccsee0
ba: As eM csccsivcuntns Bute dazedea Seema eee 137. A. aralensis Krasch.
Stems virgate, brown, with long, obliquely upright branches;
eapitula’. sessmletnerect: Ail LS cnscsrsanswsncmansstainamnaanaddnsbundaaaquecaea team
eT Ee 153. A. namanganica Poljak. var. uzbekistanica Poljak.
Terminal lobes of lower leaves small, broadly linear; mature
capitula mainly divergent or droOping .......... ee eeseeeseeeneeeeeeeeeees
LPIA GRIS Non disy LASERS RRS cect ERO 125. A. halophila Krasch.
Terminal lobes of lower leaves narrowly linear; Capitula mainly
Greeti da tektites aleel, Ra ap li DUN scant Re 55.
Terminal lobe of lower leaves usually up to 4-6 mm long.. 56.
Stems usually straight; leaf lobes narrowly linear; inflorescence
branches*miostlyiupr'tos4=7memmlongi28 215... 2B RE.
ATR. VL bends 160. A. kopetdaghensis Krasch. ex Poljak.
Stems partly flexuous; leaf lobes linear or oblong-linear,
inflorescence branches longer .................. 156. A. scotina Nevski
Perennial plant with woody, short, perennial, sterile shoots forming
small tussocks; stems rather numerous; capitula with grayish,
arachnoid-hairy involucte .............. 153. A. namanganica Poljak.
Semishrubs with woody, erect or ascending, sterile shoots covered
with grayish bark; capitula with bright glaucescent green, punctate-
glandular, involucte:...ck002 0004.10 CIR Bie See 58.
Inflorescence branches shorter; leaves grayish; upper bracteal
leaves simple, not exceeding capitula; capitula 3-4 mm long
UR acdetvtede des. ELE Renee, 164. A. prasina Krasch. ex Poljak.
Inflorescence branches elongate; leaves grayish-green, upper
bracteal leaves pinnate, some simple, exceeding capitula; capitula
D3 imma LOB KEE. SR ah Reade sda eat ed baabaees
Yoh Geck geen 163. A. badhysi (Krasch.) and Lincz. ex Poljak.
Lower leaves twice or thrice pinnately cut ...........cssseeeseeeees 60.

Srl

60.

él.

62.

63.

64.

65(60).

66.

Lower teavesitwice pinnately Gut... ci2ii. WON Baik 1 vdoeetn 69.
Plant up to 40-75 cm high, glabrous or subglabrous at end of
vegetative growth; terminal lobes of lower leaves usually up to 4—
8 mm long; capitula 3-5 mm Iong ........ elec eeeeeeeeneeeeeeeeees 61.
Plant usually 10-40 cm high, more or less hairy up to the end of
vegetative growth; terminal lobes of lower leaves shorter, less

often longer than 4 mm; capitula 2.0—-3.5 mm long .................. 65.
Mature capitula: 3=Siminivlong) iis c2v.i0. align albeul......2 62.
Capitula not:more: than 3mm longs ii..0....05. tnd. Sead... 64.

Stems stramineous-brownish, with narrowly linear pancile; terminal
lobes of lower leaves filiform-linear, 2-8 mm long ................
A lo tated 146. A. ciniformis Krasch. and M. Pop. ex Poljak.
Stems light brownish; panicle narrowly oblong, with obliquely
upright branches; terminal lobes of lower leaves narrowly linear
or litter, 1 Sisane Tore. 15012. 2c Att taeda ge. Borat... oee 63.
Lower leaves with oblong lamina; their terminal lobes 3-5 mm
long, linear; plants with pleasant lemon fragrance ................00++
NNER Sons. tw, abe tae. 14 161. A. balchanorum Krasch.
Lower leaves with oval or roundish lamina, their terminal lobes
1-3 mm long, almost lobate; plant with camphor fragrance ........
poles 000 Ges SS. RS a 162. A. turcomanica Gndg.
Terminal lobes of leaves linear; upper bracteal leaves usually
exceeding capitula; capitula oblong-ovoid (2 mm in dia and up to
3yummilong)"....c0nie. 147. A. glanduligera Krasch. ex Poljak.
Terminal lobes of leaves narrowly linear; upper bracteal leaves
usually not exceeding capitula; capitula narrowly ovoid ................
.... 141. A. tenuisecta Nevski var. karataviense Krasch. ex Poljak.
Small shrub, 10-25 cm high, with numerous, woody, strongly
reduced, sterile stolons forming rather compact tussocks; stems
numerous, initially arachnoid-hairy, later glabrous, dark brown or
brownish; capitula narrow, 2.0—-2.5 mm long, in compact condensed
PR MMIM i Sec ee cea nssxacninctamenccennsn se nn ssanaswnnsn= RAGS MAcIREMODD MAREIEE isso 66.
Small shrub, 20-40 cm high, with loose, inconspicuous tussocks;
stems fewer, by the end of vegetative growth partly glabrescent,
arachnoid hairy; capitula ovoid or oblong-ovoid, 2.0—3.5 mm long,
imeomparatively lax! pamiche tov... 2.5. 0a). sed. doth). VOW oie. 00nd 67.
General color of plants dark greenish-brown from caducous
pubescence; stems glabrous in second half of summer; dark brown;
terminal lobes of leaves linear lanceolate; involucral bracts glabrous
or subelabrous; yellowish, lustrous. ........... [ci0sdies Seva decnneess dn anss
sie hpe teeth ir. a ie 133. A. pauciflora Web. ex Stechm.
General color of plants grayish from pubescence persisting up to
the fall; stems usually hairy, stramineous-brownish; terminal lobes

544

67.

68.

572 69(59).

71(69).

73.

74.

of leaves narrowly linear; involucral bracts whitish hairy and
punctate-glandular .......... 134. A. gracilescens Krasch. and Iljin
Capitula oblong-ovoid, 3.0-3.5 mm long; lower leaves 1.5—2.5
cm long, their terminal lobes filiform-linear 3-5(7) mm long
121. A. taurica Willd.
Capitula ovoid, 2-3 mm long; lower leaves 2-4 cm long, their
terminal lobes narrowly linear, 2-3 mm long ..............:ceceeeee 68.
Middle cauline leaves always withering and usually falling by the
second half of summer; plants of clayey alkali soils
aT, SEN ROE 122. A. lercheana Web. ex Stechm.
Middle cauline leaves persisting into summer and not withering;
plants of desert-steppes
ky 122. A. lercheana Web. ex Stechm. var. astrachanica Poljak.
Plants mostly 50-80 cm high; lower leaves usually not less than
4-8 cm long; terminal lobes 4-12 mm long ............ eee 70.
Plants shorter, with smaller leaves ............:ccsccesecssseseseeesseeeeees 71.
Lower leaves 4—6 cm long, their terminal lobes 4-8 mm long,
narrowly linear, 0.5—0.7 mm wide, with short stiff tip; capitula 3
mimilonatt hein! Cee Oh ag 140. A. transiliensis Poljak.
Lower leaves 5-8 cm long, their terminal lobes 7-12 mm long
and 1 mm wide, scarcely acuminate; capitula up to 4 mm long
pe be ee i 8 152. A. gypsacea Krasch., Pop. and Lincz. ex Poljak.
Capitula narrowly cylindrical, up to 4-5 mm long, with densely
hairy and tuberculate-glandular involucre; stem slender, glabrous
at the end of vegetative growth, dark brown
we CREED RONG LADS EGE ot cc Re ae 165. A. spicigera C. Koch.

POOP eee oem eee HEHEHE HET TSO SESESE SEES ESSERE SEES ESSE ODE EEEEOES

we eeeeresceneees

PCCP CORE eee eT T EEE HETTETE SOOO HE EEE OSES OHSS SHEE EEE T SEH OES OEES

Capitula ovoid; plant with characters otherwise ..................+ 72.
Capitula 3-4 mm long; plant whitish from dense tomentum almost
up'tocéndofivegetative erowthr).: 2nG.0k A eee 73.

Capitula 2-3 mm; plants grayish-green or green, at the end of
vegetative growth glabrous or relatively weakly hairy, less often
plants’ densely. haiti: 22.22 cc.siacctebcwousduneedoveud tattoo e eee tees WD.
Lower leaves 2-4 cm long, their terminal lobes linear 3 mm long,
subobtuses flowers! 6=Bo 206 Ae a ee OL Went.
rg aa fe ips| feL bbe 120. A. fragrans Willd. var. ericanica Bess.
Lower leaves up to 2.5 cm long, their terminal lobes narrowly
linear and acute or filiform-linear, 2-4 mm long; flowers 3-4
Lgedoas SUNRISE AE RE 74.
Stems relatively slender, 25-40 cm high; panicle more or less
dense, oblong
a LY 135. A. terrae albae Krasch. var. heptapotamica Poljak.
Stems thicker, 20—35 cm high; panicle narrow, condensed, almost
firteaty jean? ot iil ee eR ct So
.. 135. A. terrae albae Krasch. var. semiarida Krasch. and Lavr.

75(72).

78(75).

573 a

81.

545

Plant glaucescent green, with stramineous brownish smooth stems
bh ARO ER Meet Ait erieekl. Rages: 145. A. cina Berg. ex Poljak.

Plants green or grayish-green, more or less hairy .............0 76.
Terminal lobes of leaves linear or oblong-linear .................. 77.
Terminal lobes of lower leaves narrowly linear ............... es 78.

Stems slender, 16—25(30) cm high; panicle very narrow with fewer
capitula; terminal lobes of leaves linear or prolate-linear ................
RAMS AA Orden ARIA G IO 144. A. fedtschenkoana Krasch.
Stems sturdy, virgate, 25-45 cm high; panicle oblong, dense, with
large number of capitula; terminal lobes of leaves linear ..........
pos ARAL ROM Qe weeclhainhn. 151. A. valida Krasch. ex Poljak.
Stems branched almost from base; capitula few, remote or more
or less approximate in comparatively narrow, pyramidal panicle
sedis 144. A. fedtschenkoana Krasch. var. issykkulense Poljak.
Stems branched in upper part; capitula approximate in prolate

PATOL 2H TAIRA ROA hic Hlth ede ate a teoaee 79.
Lower cauline leaves up to 3-4 cm ION ...........c:cccseeeeeeeeeseeeees 80.
Lower 'cauling leaves! 1,:5—310 cnn long 2.26600...) ne selicdedieel 81.

Terminal lobes of lower leaves 5-12 mm long; panicle oblong-
Bpcary plurity 50-245 gents Gi Ch Wid, UA lcs A RAG case
Tole Pate nieve Obata] 139. A. sublessingiana (Keller) Krasch. ex Poljak.
Terminal lobes of leaves much shorter; panicle narrow linear;
plant 25-30 eri Migh (fs, 2G ken .0de ANI oo I ee et edeay cen ad ends
.... 139. A. sublessingiana Krasch. ex Poljak. var. gorjavii Poljak.
Plants 12-20 cm high, grayish-green from dense hairs persisting
till, end) of wemetative, STOWE... ;..c<..1..c0n0..0casenrnnsbblenesgunteaen na ubaaess
cre eaeerey tent ee 141. A. tenuisecta Nevski var. mucronulata Poljak.
Plants 30-40 cm high, weakly hairy in second half of summer,
MGCHUMEK Ri 141. A. tenuisecta Nevski var. glaucina Poljak.

Section 1. Serphidium.—All leaves simple, entire or pinnately
lobed or even partly, at apex, with 3-5 teeth.

Series 1. Maritimae Poljak.—Perennials, more or less densely
hairy, with short, perennial, creeping or ascending nonflowering shoots,
not or partly forming tussock; lower cauline leaves or leaves of
vegetative shoots twice or thrice pinnately incised.

117. A. maritima L. Sp. pl. (1753) 846.—A. maritima Stechm.
Dissert. Artemis. (1775) 17, pro parte. —A. maritima linnaeana Bess.
in Bull. Soc. Nat. Mosc. VII (1834) 33.—A. maritima genuina Ldb.
Fl. Ross. II, 2 (1844-1846) 571, pro parte.—A. maritima L. ssp.
maritima (L.) Hegi, Ill. Fl. II, 662.—A. maritima var. genuina Kryl.

574

546

Fl. Alt. III (1904), non. L.—A. palmata Lam. Encycl. meth. I (1789)
268; Bess. op. cit. 29.—A. gallica Willd. Sp. pl. HII (1800) 1834.—A.
salina Willd. ibid. 1884.—A. maritima boschniakiana Bess. op. cit.
39.—A. boschniakiana (Bess.) DC. Prodr. VI (1837) 104.

Whole plant whitish-woolly tomentose. Root vertical, woody, with
short, perennial, creeping shoots. Fertile stems few, 20-—40(50) cm
high, ascending or erect, branched above, with long, partly short
branches, most often not strongly divergent. Lower cauline leaves
petiolate, 2-3(4) cm long, lamina oval, twice or thrice pinnatisect,
their terminal lobes linear, somewhat flat, subacute or obtuse; middle
cauline leaves short-petiolate or sessile, once pinnatisect or twice
pinnately cut, uppermost leaves bracteal, often exceeding capitula,
sessile, simple, linear, sometimes with 2 short lateral lobes at base.
Capitula sessile, ovoid, usually 3-4 mm or 2.5 mm long, with 3 flowers
(var. palmata Lam.), rather densely spicate on branches in somewhat
appressed or more or less spreading paniculate inflorescence. Involucral
bracts with scarious margin; outer bracts small, oval, densely hairy,
inner bracts larger, oblong-lanceolate, glabrous or subglabrous, smooth.
Florets (6)7—10.

Seacoasts.—European part: Baltic Region, Bessarabia, Black Sea
Region; Caucasus: Transcaucasia. General distribution: Scandinavia,
Atlantic Europe, Mediterranean Region (west, east), Balkans-Asia
Minor. Described from Denmark. Type in London.

Note. The following varieties are distinguished:

1) Var. palmata Lam.—with small (up to 2.5 mm long), few-
flowered capitula.

2) Var. boschaniakiana Bess.—With bi- or tripinnatisect leaves
having narrowly linear, almost filiform lobes.

3) Var. salina (Willd.) Koch.—With dense whitish pubescence
and very strongly divergent branches.

118. A. monogyna Waldst. and Kit. Descr. et Icon. pl. rar. Hung.
1 (1802) 77, tab. 75; Poljak. in Majevski, Fl. 584.—A. santonica
Waldst. and Kit. k. c. p. XXX, non L. —A. nutans Willd. Sp. pl. Ill
(1800) 1832.—A. maritima var. wilhelmsiana Bess. var. blumiana Bess.
var. O. kitaibelliana Bess. in Bull. Soc. Nat. Mosc. VII (1834) 32,
40.—A. maritima var. monogyna Ldb. FI. Ross. II, 2 (1844-1846)
573.—A. maritima salina Kell. V Obl. Polupustyni (1907) 104-109.—
A. salina ssp. monogyna Sagorski in Oest. Bot. Zeitschr. (1907) 16.—
A. salina ssp. monogyna var. kitaibelii Sagorski in Mitt. Thir. Bot.
Ver. Neue Folge, XXIII (1908) 82.—A. santonicum var. monogyna
Fritsch. in Sched. ad. Fl. exs. Austro-Hung. No. 2265.—A. maritima
var. monogyna Fedtsch. in Perech. Rast. Turk. IV (1911) 194, non

575

547

Waldst. and Kit.—A. maritima ssp. monogyna (Waldst. and Kit.) Hegi,
Ill. Fl. If, 2 (1929) 664.—A. salina Kell. in Fl. Yugo-Vost. Evrop. Ch.
SSSR, VI (1930) 354, non Willd. (pro parte).—A. stepposa Kell. and
Kom. Ibid. 355 (pro parte).—A. maritima salina Kell. f. citriodora
Kazakevicz., nom.—A. kobstanica Rzazade and A. paucifloriformis
Rzazade in Izv. Akad. Nauk Azerb. SSR, 3 (1955) 25.—Exs.: FI.
Hung. No. 585.

Perennial. Plant grayish, densely arachnoid-hairy, later weakly hairy
or subglabrous. Root relatively slender, woody, vertical, with short,
ascending, perennial, woody shoots, not forming tussocks. Fertile shoots
more often 12—20, sometimes even more, less often few, erect or weakly
flexuous, slender, stiff, 30-40(50) cm high, branched above, with
obliquely upward-spreading, sometimes drooping branches. Leaves of
vegetative shoots and lower cauline leaves, long-petiolate, 2-4(5) cm
long, lamina oblong-ovate or oblong, twice pinnately cut, their terminal
lobes narrowly linear, subacute; middle cauline leaves sessile, uppermost
bracteal, simple, linear. Capitula usually pedunculate, narrowly
campanulate, with mature, 3-4 mm long, approximate or somewhat
remote, often drooping, in more or less spreading paniculate
inflorescence. Involucral bracts with wide scarious margin; outer bracts
oval, pubescent dorsally, inner bracts larger, oblong, in upper part
weakly hairy or glabrous, smooth. Flowers 3-4, usually purple red.

River terraces, ravines, depressions, salt marshes and alkali soils,
sometimes on outcrops of calcareous rocks.—European part: Black
Sea Region, Bessarabia, Crimea, Lower Don, Trans-Volga, Lower
Volga. General Distribution: Central Europe. Described from Hun-
gary. Type in Vienna.

Note. The form citriodora Kazak., found on the left bank of the
Volga River, is distinguished by a rather strong lemon fragrance.
Morphologically, it is closest to A. monogyna Waldst. and Kit.
According to the data of Spiridonov, the yield of essential oil from f.
citriodora is as high as 0.4%, and the oil contains pinene, about 8%
camphors, about 19% borneole and cadinene (?), 34%—citral and 2%
phenols.

119. A. szowitziana (Bess.) Grossh. Fl. Kavk. IV (1934) 142;
Opred. Rast. Kavk. 465.—A. maritima var. szowitziana Bess. in Bull.
Soc. Nat. Mosc. VII (1834) 34; Ldb. Fl. ross. li, 2, 574.—A. maritima
var. hanseniana Bess. op. cit. 44.—A. hanseniana Grossh. Fl. Kavk.
IV (1934) 143 p. p.—A. prilipkoana Rzazade and A. nachitschevanica
Rzazade (p. p.) and A. iskenderiana Rzazade p. p. in Izv. Akad. Nauk
Azerb. SSR, 3 (1955) 26-28.

576

548

Perennial. Plant grayish, densely arachnoid hairy, later weakly
pubescent, partly glabrous. Root woody, with short nonflowering shoots —
bearing rosettes of leaves in upper part. Fertile stems few, straight,
strong, 60-100 cm high, branched above with long, sulcate, strongly
divergent, sometimes almost horizontal branches. Lower cauline leaves
and leaves of vegetative shoots deciduous, up to 10 cm long and 4 cm
wide, long-petiolate, with pinnately cut auricles; lamina twice or thrice
pinnately cut, lobes remote, terminal lobes flat, linear, 5-10 mm long
and 1 mm wide, subacute; middle and upper cauline leaves shorter,
sessile or short-petiolate, less divided; bracteal leaves simple, linear,
2—4(5) mm long. Inflorescence broadly paniculate; capitula ovoid,
pedunculate, drooping, 3.5—4.0 mm long and 2 mm in dia, more or less
remote. Involucral bracts hairy, with scarious margin; outer bracts
oblong-oval, inner larger, elliptical. Flowers 4-5. Flowering August to
September.

Ravines, coastal regions, often as weed in irrigated fields.—
Caucasus: Eastern Transcaucasia. Endemic. Described from the
Caucasian coast of the Caspian Sea (near Salyan). Type in Leningrad.

Economic Importance. According to A.A. Grossheim (1947), the
plant contains about 0.5-0.6% essential oil. According to I.A. Damirov
(1949), the oil contains 1.4—3.2% santonin.

Note. According to Besser, A. maritima f. hanseniana Bess. has
oblong-linear leaf lobes but the lower capitula are drooping, which
indicates its definite proximity to A. szowitziana Bess. and A. monogyna
Waldst. and Kit. but in no way to A. fragans Willd., as was believed
by Grossheim.

120. A. fragrans Willd. Sp. pl. III (1804) 1835; DC. Prodr. VI,
104, excl. syn.; Grossh. Opred. Rast. Kavk. 465.—A. fragrans Willd.
B. phyllostachys Boiss. Fl. or. III (1875) 366.—A. fragrans Willd. ssp.
gurganica Krasch. in Otch. o Rab. Pochv.-bot. Otr. Kazakhsk. Eksp.
Akad. Nauk SSSR, IV, 2 (1930) 245.—A. phyllostachys (Boiss.) Woron.
in Sched. ad Herb. Fl. cauc. X, VI (1931) No. 498.—A. maritima var.
erivanica Bess. in Bull. Soc. Nat. Mosc. VII (1834) 44; Ldb. Fl. Ross.
II, 574.—A. taurica var. erivanica DC. op. cit. 104.—A. taurica var.
hanseniana DC. op. cit. 104.—A. maritima var. v. meyeriana Bess.
op. cit. 39.—A. maritima var. fragrans Ldb. op. cit. 570.—A. maritima
var. persica f. hanseniana Ldb. op. cit. 574.—A. hanseniana Grossh.
Fl. Kavk. IV (1934) 146.—A. hanseniana var. phyllostachys (Boiss.)
Grossh. and var. divaricata Grossh. op. cit. 146.—A. meyeriana Grossh.
Opred. Rast. Kavk. (1949) 465.—A. meyeriana Grossh. var. divaricata
Grossh. op. cit. 465.—A. erivanica Grossh. op. cit. 143.—A. fedorovi
Rzazade and A. iskenderiana Rzazade p. p. and A. divaricata (Grossh.)

577

549

Rzazade and A. nachitschevanica Rzazade p. p. in Izv. Akad. Nauk
Azerb. SSR, 3 (1955) 24, 27.

Perennial. Plant 25-30-45 cm high, whitish-tomentose at beginning
of vegetative growth, later partly glabrous. Root vertical, woody, about
0.5—1.0 cm thick. Fertile stems few or more or less numerous, erect,
branched from slightly above middle, with divergent branches. Lower
cauline leaves petiolate, 2.5—-3.5 cm long, oblong, twice pinnately cut,
terminal lobes 2-5 mm long, thickish, narrowly linear or filiform-
linear obtuse; middle cauline leaves sessile, simple pinnate, with
pinnately cut auricles; bracteal leaves simple, linear, 2-4 mm long.
Capitula sessile, broadly ovoid or subglobose, 3.0-4.5 mm long and
2-3 mm (var. gurganica (Krasch.) P. Pol.), erect or divergent, in oblong,
pyramidal, paniculate inflorescence. Outer involucral bracts short, oval,
convex, inner bracts larger, oblong-linear, with wide scarious margin,
greenish dorsally. Florets yellow, 3-6, anthers with short filaments,
linear, apical appendages of anthers narrowly linear, almost cuneate,
basal appendages much smaller, somewhat subobtuse; style included;
stigma lobes short-linear, erect, truncate, ciliate apically along margin,
divergent after flowering, slightly exserted from corolla. Flowering
August.

Foothill plains as well as clayey slopes of foot-hills.—Caucasus:
Eastern and Southern Transcaucasia, Dagestan; Soviet Central Asia:
Aralo-Caspian Region. General distribution: Armenia and Kurdistan,
Iran. Described from Armenia. Type in Berlin.

Note 1. A. fragrans Willd. is very close to A. taurica Willd. The
latter species is not found in Azerbaidzhan. The epithet “A.
hanseniana,” used by Grossheim in the “Flora Kavkaza” [Flora of the
Caucasus], actually replaced Willdenow’s A. fragrans, which is
completely illegitimate because Besser’s form “A. maritima
hanseniana” can be compared only with A. szowitziana Willd. and not
in any way with A. fragrans Willd. Thus, I do not find any basis for
splitting A. fragrans Willd. into two or even three species, as was
proposed by Grossheim. The history of A. divaricata (Grossh.) Rzazade
is as follows. First, Grossheim segregated a form that seemingly con-
formed to A. maritima var. meyeriana Bess. (differing, according to
Besser, by “a slightly divergent panicle and hyaline inner involucral
bracts,”) and elevated it to the rank of a separate new species—A.
meyeriana Grossh. At the same time, the author did not provide a
description, nor did he indicate the type specimen, but only mentioned
that his new species is distinguished from A. fragrans Willd. by the
absence of grooves on the dorsal side of the leaves. Later, in the
“Opradelitel’ Rastenii Kavkaza” [Manual of Plants of Caucasus] A.A.
Grossheim split off from A. meyeriana Grossh. one more variety—var.

578

550

divaricata Grossh., which is distinguished by spreading, often flexu-
ous branches. Rzazade elevated Grossheim’s variety to the rank of
species without mentioning the type and without giving a description.

Note 2. The following varieties are distinguished:

1) Var. erivanica Bess.—distinguished by a comparatively narrow
and compact panicle, with obliquely upward directed, slightly elongate
branches, linear lobes of the leaves, and ovoid capitula; found in
Southern and Eastern Transcaucasia;

2) Var. phyllostachys Boiss.—With long (up to 20 cm), angular,
strongly divergent, sparse branches and ovoid capitula; found in
Southern and Eastern Transcaucasia.

3) Var. gurganica Krasch.—With not so strongly divergent and
elongate branches and ovoid capitula; south-western Turkmenia.

Economic Importance. According to Karakash and Zelentsova
(1938), the fresh and the dry herb of A. fragrans contains 0.87—0.92%
essential oil. Traces of alkaloids have also been reported (Mirzoyan)
and Amirzadyan, 1949). In the opinion of Magakyan and others, it is
one of the main fodder plants in Armenia and the Nakhichevan ASSR
in the fall. At this time of the year, it is relished by sheep and less so
by donkeys and horses. It is also eaten in the spring but less than
ephemerals and ephemeroids. It is not eaten by big horn cattle; in
some regions (Magakyan, 1953) it is mowed for hay. There are reports
about the prophylactic (anthelmintic) action of this wormwood on sheep
(Magakyan, 1953). According to I.A. Damirov (1949), the essential oil
contains 1.5—2.9% santonin.

According to Grossheim (1946), the plant contains 0.3-0.19%
essential oil; its main constituents are cineole, thujone and others.
Moreover, the plant contains 5—12% tannin (depending upon the stage
of development); the leaves collected in May contain 14.5 mg% of
vitamin C, while those collected in the fall contain 11.4 mg%.

121. A. taurica Willd. Sp. pl. III (1800) 1837; MB. FI. taur.-cauc.
II (1808) 291.—A. maritima var. v. taurica Ldb. FI. Ross. II, 2 (1846)
574.

Perennial. Plant whitish or gray from dense arachnoid hairs
persisting to a considerable extent to the end of vegetative growth.
Root vertical, woody, thick, with strongly reduced, sterile shoots and
rather numerous straight, stiff, sulcate, stems 15-35-40 cm high,
branched above with short or slightly elongated, obliquely upward
directed branches. Leaves of sterile shoots and lower cauline leaves
petiolate, 1.5—-2.5 cm long, lamina oval-oblong, twice or almost thrice
pinnately cut, their terminal lobes linear-filiform, 3-5 mm long,
acuminate or subobtuse; upper cauline leaves subsessile, less divided;

579

551

bracteal leaves simple, linear-filiform, 3—5(6) mm long. Capitula sessile,
ovoid, 3.0-3.5 mm long and about 2 mm in dia, upright, rather densely
spicate on branches in more or less narrowly paniculate inflorescence.
Involucre imbricate, densely covered with gray hairs; involucral bracts
convex, with scarious margin, outer bracts oval, inner larger,
oblanceolate or lanceolate. Florets 6-8. Achenes up to | mm long,
ovate, somewhat flat, grayish, finely sulcate. Flowering August.

Clayey alkali soils, and also as ruderal and weed.—European Part:
Crimea; Caucasus: Ciscaucasia. Endemic. Described from Crimea. Type
in Berlin.

Economic Importance. The essential oil of A. taurica Willd., as
reported by N.A. Lvov, consists of obeintole (C,,H,,.O) and terpene
(C,,H,,), and possesses poisonous properties. Moreover, this plant also
contains lactone tauricine (C,,H,,O,) which is related to santonin. The
oil of A. taurica Willd. causes strong irritation, spasms, and deafness
in animals. Tauricine has the same properties, but in addition it affects
the autonomic nervous system. Tauric wormwood is poisonous to cattle
and particularly to horses.

According to the data of Troitsky, in the pastures of Crimea it is
relished by sheep in the fall-winter period as well as in the early spring
when the young shoots have not yet acquired a bitter taste. According
to Larin, these shoots, as shown by analysis, contain (in %): protein up
to 27.1, fat 4.8, nitrogen-free extract 35.2, and only 12.4 cellulose. In
the opinion of Larin, poisoning from hay of this species is not observed
particularly frequently and not everywhere. However, he advises great
caution while feeding animals with hay containing a large proportion
of this wormwood.

122. A. lercheana Web. ex Stechm. Dissert. Artemisia (1775)
XXIV, XXV, excl. f. Gmel. Fl. Sib. II (1775) 114; Willd. Sp. pl. II,
3, 1838; Grossh. Opred. Rast. Kavk. 465; Poljak. in Majevski, Fl. 585;
in Tr. Inst. Khim. Akad. Nauk. Kaz. SSR. IV, 94.—A. maritima var.
lercheana Bess. in Bull. Soc. Nat. Mosc. VII (1834) 37; Ldb. FI. Ross.
II, 572; Fedtsch. Perech. Rast. Turk. IV, 193.—A. maritima var. rossica
Bess. op. cit. 38, pro parte.—A. humilis MB. Beschr. Casp. (1801) 83,
210.—A. maritima var. lercheana Ldb. FI. Ross. II, 2 (1844-1846)
572, pro parte.—A. maritima var. incana Kell. V Obl. Polupustyni, II
(1907) 110, non Druce.—A. maritima Bess. ssp. incana (Kell.) Krasch.
in Otch. o Rab. Pochv.-Bot. Otr. Kazakhsk. Eksp. Akad. Nauk SSSR,
IV, 2 (1930) 268, pro parte.—A. maritima astrachanica Kazak. (nomen)
in Zhurn. Opytn. Agronom. Yugo-Vost. IV, II (1927).—A. caspica
Kell. in Fl. Yugo-Vost. Evrop. Ch. SSSR VI (1936) 356, pro parte.—

580

552

A. astrachanica Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVI (1954) 422.

Perennial. Whole plant grayish from dense, arachnoid hairs, later
partly glabrous. Root woody, with perennial, woody, strongly reduced
and short, annual, vegetative, leafy shoots forming comparatively
small tussocks. Flowering stems rather numerous or few, erect or
ascending at base, 18-—35(40) cm high, branched in upper half, with
obliquely upright branches. Leaves of vegetative shoots and lower
cauline leaves petiolate, 2-4 cm long, lamina oval-oblong, twice or
thrice pinnately cut; their terminal lobes narrowly linear, usually 2-3
mm long; middle cauline leaves withering in summer, sessile or short-
petiolate, twice pinnately cut, with pinnately cut auricles; uppermost
leaves bracteal, simple, linear. Capitula sessile or on short peduncles,
ovate; mature capitula narrow-goblet shaped, 2.0—2.5(3.0) mm long,
usually not drooping, remote or few, spicately crowded on terminal
branches in more or less compact paniculate inflorescence. Outer
involucral bracts small, oval, densely arachnoid-hairy, inner bracts
larger, oblong, with wide scarious border, apically hairy, elsewhere
glabrous. Flowers 4—5; corolla yellow or pinkish. Flowering August to
September.

Clayey alkali soils—European part: Lower Don, Lower Volga;
Western Siberia: Upper Tobol, Orenburg Region; Soviet Central Asia:
Aralo-Caspian Region. Endemic. Described from Stalingrad District.
Type in Berlin.

Note. One should recognize var. astrachanica Poljak., with leaves
persisting in summer; it is a plant of desert-steppe sands, found in the
northern Caspian Region, Turgai, and western Kazakhstan.

According to the data of A.A. Rikhter, L.I. Kazakevich, O.I.
Sobolevskaya, and K.T. Suverova, the essential oil of var. astrachanica
Poljak. contains up to 90.8% camphor. According to M.I. Goryaev and
N.I. Bakanina, the oil contains cineole, camphor, thujone, borneole
and phenol. According to the data of R.N. Sazonova, it contains up to
0.2% santonin. In the opinion of Aleksandrovsky and Beguchev (1931),
the nutritive value of var. astrachanica is not high; it is relished more
by sheep and camels in the fall.

In the opinion of Larin (1956), this wormwood is a valuable fodder
plant in the semidesert fall-winter pastures.

123. A. nitrosa Web. ex Stechm. Dissert. Artemisia (1775) 24;
Bess. in Bull. Soc. Nat. Mosc. VII, 45; Ldb. Fl. Ross. II, 2, 576; Kryl.
Fl. Zap. Sib. XI, 2781, 2782.—A. maritima var. i. gmeliniana Bess.
op. cit. 38; Fedtsch. Perech. Rast. Turk. IV, 94; DC. Prodr. VI, 104.—
A. maritima var. lercheana f. gmeliniana Ldb. FI. Ross. II, 2 (1846)

581

553

573, quoad pl. Gmel.—A. maritima var. 6. genuina f. tt. suberecta
Ldb. Fl. ross. II, 2 (1846) 572, pro parte quoad pl. sib.—A. maritima
Bess. var. n. lercheana f. dahurica Turcz. FI. baic.-dahur. III (1856)
56.—A. maritima var. genuina Kryl. Fl. Alt. III (1904) 639.—A.
maritima Bess. ssp. incana (Kell.) Krasch. in Otch. Rab. Pochv.-Bot.
Otr. Kazakhsk. Eksp. Akad. Nauk SSSR, IV, 2 (1930) 268.—A.
maritima ssp. gmeliniana (Bess.) Krasch. in Spisok Rast. Gerb. SSSR,
X (1936) 99.—A. mongolorum Krasch. in Tr. Bot. Inst. Akad. Nauk
SSSR, ser. 1, No. 3 (1936) 350; Krascheninn. op. cit. 2782 incl. ssp.
gobica Krasch.—A. schischkint Krasch. in Sistemat. Zam. Gerb. Tomsk.
Univ., 1—2 (1940) 2; Kryl. Fl. Zap. Sib. XI, 2786.—A. lercheana auct.
non Web.: Ldb. FI. Alt. (1934) 84 quoad pl. Tschujens.—A. lercheana
var. humilis DC. Prodr. VI (1837) 104, excl. syn. — A. maritima “s”
lercheana humilis Ldb. Fl. Ross. II, 2 (1844-1846) 572, quoad plant.
Tschujens. Kryl. Fl. Alt. III (1904) 639.—Exs.: GRF No. 3191a, b, c
(sub A. maritima).

Perennial. Whole plant grayish-green from dense arachnoid-
tomentose hairs, sometimes hairs partly persistent up to end of
vegetative growth or subglabrous (var. subglabra Krasch.). Root
relatively slender, with few (2-5) short, leafy in upper part,
nonflowering annual shoots. Flowering stems erect, strong, up to 40(50)
cm high, less often 5-12 cm (var. gobica Krasch.), branched in upper
half, with somewhat divergent branches. Lower leaves long-petiolate,
3-7 cm long, lamina oval-lanceolate, twice pinnately cut, their terminal
lobes narrowly linear, usually 3-5 mm long, subobtuse or short-
acuminate; uppermost leaves undivided, sessile, linear. Capitula sessile,
oblong or ovate, 3-4 mm long and 1.5-2.5 mm in dia, remote or
crowded in groups of few on lateral branches in elongate, more or less
oblong, paniculate inflorescence. Involucral bracts more or less densely
hairy, later subglabrous, with scarious margin, outer bracts small, oval,
inner bracts larger, oblong-lanceolate. Flowers 3-5; corolla purple-
pink or yellow; anthers linear, apical appendages of anthers acute,
basal appendages small, subacute, stigma lobes linear, truncate, ciliate.
Flowering August to September.

River valleys, margins of steppe lakes, ravines as well as in alka-
line meadows, salt marshes and alkaline soils —Western Siberia: Upper
Tobol, Irtysh, Altai; Eastern Siberia: Angara-Sayans, Dauria; Soviet
Central Asia: Aralo-Caspian Region, Lake Balkhash Region (north).
General distribution: Dzhungaria-Kashgaria, Mongolia. Described from
Siberia. Type in Berlin.

Note. var. gobica Krasch. should be recognized. It is character-
ized by stems that are short (S—15 cm high), grayish-tomentose pubes-
cence on the leaves that persists to the end of vegetative growth, small

582

554

(up to 1.5 cm long) lower leaves that are usually pinnatisect or
bipinnatisect, and capitula that are racemosely or spicately borne. As
regards A. schischkini, it corresponds to this variety.

Economic Importance. According to the data of M.I. Goryaev,
A.T. Petushkova, and N.A. Sviridova, samples of A. nitrosum contain
0.1-0.5% alkaloids.

124. A. scopaeformis Ldb. Fl. Ross. II, 2 (1844-1846) 575;
Fedtsch. Perech. Rast. Turk. IV, 195.

Perennial. Plant (18)25-45 cm high, grayish, arachnoid-tomentose,
hairs sometimes partly shed at end of vegetative growth. Root rather
thick, woody, verticle, with perennial, woody, strongly reduced, creeping
or ascending nonflowering shoots forming a comparatively compact
tussock. Flowering stems erect or ascending, more or less numerous,
virgate, branched, with obliquely upright, less often drooping, 2.5 cm
long or longer branches (f. longiramosa Poljak.). Leaves of
nonflowering shoots and lower cauline leaves petiolate, (1)2—-4 cm
long, at base with simple, lanceolate or pinnate-lobate auricles, oblong-
oval or oblong, simple pinnately cut; lobes more or less deeply divided
into 2—4 lanceolate lobules with short-pointed tips or entire, middle
cauline leaves sessile, pinnate; upper leaves undivided, linear. Capitula
on short peduncles, ovoid, upright, 3—4 mm long, in narrow pyramidal
paniculate inflorescence. Outer involucral bracts small, oval, weakly
hairy, inner bracts larger, oblong-oval, dorsally brownish, subglabrous,
with wide scarious margin. Flowers 3-4; corolla yellow or red.

Ravines, lake and dry depressions, ancient river terraces, meadow
and takyrlike saline, crusty solonetz soils, sometimes on outcrops of
varicolored clays.—Soviet Central Asia: Aralo-Caspian Region, Lake
Balkhash Region, Irtysh. Endemic. Described from Chu River (Myun-
Kumy). Type in Leningrad.

125. A. halophila Krasch. in Otch. 0 Rab. Pochv.-Bot. Otr.
Kazakhsk. Eksp. Akad. Nauk SSSR, IV, 2 (1930) 272.

Perennial. Plant 20-40 cm high, white-tomentose. Root woody, up
to 3-6 mm thick, less often thicker, vertical, with perennial, woody,
short, ascending, nonflowering shoots forming small tussocks; annual,
very short shoots bear rosettes of leaves of summer vegetation.
Flowering stems few or more or less numerous, often branching in
lower half, with distinctly divergent lateral branches. Leaves thick;
leaves of nonflowering shoots and lower cauline leaves long-petiolate,
1-3-4 cm long, lamina oblong or linear, twice pinnately cut; lobes
divided into 2—4 short, broadly linear, acute lobes; middle and upper
leaves sessile, once or twice pinnately divided; uppermost leaves small,

555

SS
2
nt GN@ OS 'G/E,

—v.
SF

ase
+

SS
ones 7H
WEED

583 Plate XXVIII.
1 —A. halodendron Bess., upper part of stem; lower leaf, middle cauline leaf,
abortive pistil of disk floret, peripheral pistillate floret, outer and inner involucral
bracts; 2—A. pannosa Krasch., lower leaf, cauline leaf, achene, peripheral
pistillate floret, abortive pistil of disk floret, outer and inner involucral bracts,
3 —A. glauca Pall., upper part of plant, leaf, achene, outer and inner involucral
bracts, peripheral pistillate floret, abortive pistil of disk floret, 4 — A. commutata
Bess., upper part of plant, achene, lower cauline leaf, abortive pistil of disk
floret, peripheral pistillate floret, outer and inner involucral bracts.

58

wm

556

sessile, undivided, linear or even with 2 lateral lobes at base. Capitula
on short peduncles, ovate or oblong-cup-shaped, 3-4 mm long,
mature capitula drooping, remote or approximate at tips of branches,
2-3, in oblong-pyramidal panicle. Involucre many-rowed, imbricate;
outer involucral bracts, small, oval, greenish-gray, densely hairy, inner
bracts larger, oblong or oblong-oval, dorsally greenish or brownish
with wide, lustrous, scarious margin. Florets more often red. Flowering
August.

Highly saline soils of desert regions, outcrops of gypsiferous
clays.—Soviet Central Asia: Aralo-Caspian Region, Kzyl-Kum,
Kara-Kum, Syr-Darya. Endemic. Described from Ust-Urt. Type in
Leningrad.

Economic Importance. According to the data of M.I. Goryaev,
the yield of essential oil in this species is 0.82% of air-dry matter. The
oil contains a large quantity of camphor.

126. A. schrenkiana Ldb. FI. Ross. II, 2 (1844-1846) 575; Kryl.
Fl. Zap. Sib. XI, 2783.—A. albida auct. non Spreng. Ldb. FI. Alt. IV
(1834) 85, pro parte.—A. maritima Ldb. FI. Alt. TV (1833) 85, non L.;
Fedtsch. Perech. Rast. Turk. IV, 193 p. p.—Ic.: Ldb. Ic. pl. Fl. Ross.
V, tab. 478.

Perennial. Plant whitish throughout vegetative period from dense
arachnoid-woolly hairs. Root thick, with rather numerous, short,
nonflowering shoots, partly leafy. Flowering stems few, straight, at
base ascending, 30-50(60) cm high, branched above, with rather long
(to 10-15 cm), divergent branches. Lower leaves long-petiolate, 5—10
cm long, others sessile, with semiamplexicau!, pinnately cut auricles at
base; lamina broadly linear, 3-6 cm long, twice, or more often thrice
pinnately cut, lobes widely spaced, terminal lobes narrowly linear,
often enlarged above, spatulate and subobtuse, 2-6 mm long and 1.0-
1.5 mm wide. Capitula oblong-ovoid, 2.5—-3.0 mm long, usually erect,
sessile or on short peduncles, remote or more densely crowded on
lateral branches into broadly paniculate inflorescence. Involucre
imbricate, grayish-hairy; involucral bracts with scarious margin, outer
small, oblong-oval, inner longer, oblanceolate. Receptacle convex,
conical. Flowers 5-6; corolla yellow; anthers linear, apical appendages
of anthers narrowly linear, basal appendages shorter, subacute; style
shorter than stamens; stigma lobes short, ciliate, divergent. Flowering
August.

Steppe and desert-steppe regions, along river terraces, lake
depressions, ravines, meadow and puffy salt marshes or highly alka-
line soils.—Soviet Central Asia: Lake Balkhash Region, Irtysh, Altai,
Dzhungaria-Tarbagatai, Tien Shan. General distribution: Northwest-

586

557

ern Mongolia, Dzhungaria-Kashgaria. Described from the Sary-Su
River. Type in Leningrad.

According to the data of M.I. Goryaev (1952), the yield of
essential oil from A. schrenkiana Ldb. is 0.29%. Based on analyses,
the oil consists of the following constituents: organic acids—9%,
phenols—8.7%; camphors—10.25%; aldehydes—2.75%; cineole—
63.5%.

According to the data of M.I. Goryaev, A.T. Petushkova, and
N.A. Sviridova, plant samples of this species contain 0.1-0.5%
alkaloids.

Series 2. Compactae Poljak.—Capitula few or several on branches
in rather compact spikes or racemes forming spicate or racemose
paniculate inflorescence. Mainly alpine or montane plants.

127. A. compacta Fisch. ex DC. Prodr. VI (1837) 102; Kryl.
Fl. Zap. Sib. XI, 2784.—A. albida Ldb. Fl. Alt. IV (1833) 85, pro
parte.—A. maritima €. fischeriana 8. stephaniana, i. helmiana
Bess. in Bull. Soc. Nat. Mosc. VII (1834) 34, 36.—A. maritima €.
compacta Ldb. FI. Ross. II, 2 (1844-1846) 572; Kryl. Fl, Alt. III, 639;
Fedtsch. Perech Rast. Turk. IV, 193.—A. tianschanica Krasch. ex
Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954)
416; Poljak. in Tr. Inst. Khim. Akad. Nauk Kaz. SSR. IV (1959) 19,
20, 27, 28.

Perennial. Plant grayish-tomentose till end of vegetative growth.
Root slender, woody, with numerous, short, woody, perennial,
nonflowering shoots forming small, more or less compact tussocks.
Flowering stems few or more or less numerous, relatively slender,
erect or partly ascending, 15-40 cm high, branched above, with
obliquely upright branches. Basal and lower cauline leaves short-
petiolate, 2-3 cm long, twice- or thrice-pinnately cut, their terminal
lobes linear, subobtuse; middle cauline leaves less compound;
uppermost leaves sessile, undivided, linear, often exceeding
inflorescence. Capitula ovoid, 3-4 mm long and 2.0—2.5 mm in dia,
sessile, crowded on branches in more or less compact, short spikes,
remote or approximate and forming narrowly spicate-paniculate
inflorescence. Outer involucral bracts oval, acuminate, hairy, inner
bracts elliptical-lanceolate, subobtuse, glabrous, with scarious margin.
Florets 3-4; corolla usually purple-pink, less often yellow; anthers
linear, apical appendages of anthers acute, basal appendages shorter,
short-acuminate; style shorter than stamens; stigma lobes linear,
truncate, ciliate. Flowering August.

587

558

Clayey or rubbly-clayey slopes from foothill plains to high moun-
tains.—Western Siberia: Altai, Irtysh; Soviet Central Asia: Dzhungaria-
Tarbagatai, Tien Shan, Pamiro-Alai Region. General. distribution:
Dzhungaria-Kashgaria. Described from the Chuya River in Altai. Type
in Leningrad.

According to the data of M.I. Goryaev, A.T. Petushkova, and
N.A. Sviridova, about 0.1-0.5% alkaloids was found in plants of A.
compacta Fisch.; these figures were confirmed by M.I. Goryaev (1959).
According to the data of M.I. Goryaev, I.M. Shibanova, and L.A.
Ignatova (1953), the essential oil contains: cineole—42%, carbonyl
compounds—21.8%; the latter include 16.8% camphors and 5.62%
aldehydes.

The aldehyde mixture presumably contains phellandral and
perillaldehyde; in the higher fractions, apart from these, also present
are azulene-forming sesquiterpenes imparting a blue color, but their
content in the oil is very insignificant.

128. A. rhodantha Rupr. in Mém. Acad. Sc. Pétersb. VII, XIV,
4 (1869) 52; Fedtsch. Perech. Rast. Turk. IV, 195.

Perennial. Whole plant whitish-tomentose till end of vegetative
growth, 6—15(20) cm high. Tap root thick, with underground or aerial,
strongly reduced, upright, perennial woody shoots, rather closely
approximate and also with still shorter, annual, nonflowering shoots
forming compact tussocks. Flowering stems rather numerous or few,
4-15 cm high, sulcate, simple. Leaves of nonflowering shoots and
lower cauline leaves petiolate, 1.0-2.5(3.0) cm long, petiole 0.5—1.5
cm long, lamina roundish or oval, twice, or almost thrice pinnately cut,
lobes palmate-pinnate, terminal lobes linear, 2-3 mm _ long,
subacuminate; cauline leaves short-petiolate or sessile, twice pinnate,
at base with pinnately or palmately cut auricles; upper bracteal leaves
pinnate or in part simple, broadly linear, exceeding or as long as
capitula. Capitula sessile, ovoid, 3 mm long, in groups of few borne
in compact, oval, lax spikes, on very short peduncles or sessile,
adpressed to stem in simple racemose or capitate-spicate inflorescence.
Outer involucral bracts oblong-lanceolate, tomentose, inner bracts
narrowly lanceolate, dorsally hairy, with scarious margin. Florets 5—7;
corolla pink; anthers linear, apical appendages of anthers subulate,
basal appendages shorter, short-acuminate; style shorter than stamens;
stigma lobes linear, truncate, short-ciliate. Flowering August.

Alpine zone of mountains, on clayey and rubbly slopes.—Soviet
Central Asia: Tien Shan, Pamiro-Alai Region. Endemic. Described
from Lake Chatyrkul. Type in Leningrad.

588

559

Note. Until recently it was thought that A. rhodantha Rupr. is
found only in the inner Tien Shan. However, our expedition to the
eastern Pamirs in 1954 showed that this species is very widespread
there too, but the local botanists usually regard it as A. skorniakowii.

129. A. prolixa Krasch. ex Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVI (1954) 401.

Perennial. Tap root woody, with perennial, woody, short, almost
straight, ascending, nonflowering shoots covered with grayish-brown
bark. Flowering shoots few or more or less numerous, 50-60 cm high,
erect, white-tomentose, branched above, with long (up to 25-30 cm),
straight, obliquely upright branches. Leaves densely arachnoid-hairy;
lower cauline leaves withering before flowering, long-petiolate, 4—5
mm long, lamina oblong-oval, twice pinnately cut, terminal lobes
narrowly linear, 5—8(10) mm long, short-acuminate; middle cauline
leaves smaller, short-petiolate or sessile, with pinnate auricles at base;
upper leaves sessile, undivided, linear, sometimes with 2 lateral lobes
at base. Capitula sessile, ovate, 3 mm long, usually in groups of few
in approximate or remote heads on very short terminal branches in
long (up to 30 cm) pyramidal-paniculate inflorescence. Involucral bracts
with wide scarious margin; outer bracts smaller, oval, densely covered
with white hairs; inner larger, oblong, glabrous, smooth. Florets 5-6;
corolla yellow.

Clayey slopes of lower mountains.—Soviet Central Asia: Syr-Darya,
Pamiro-Alai Region, Tien Shan. Endemic. Described from Fergana,
from hills near the village of Dragomirovo. Type in Leningrad.

130. A. dubjanskyana Krasch. ex Poljak. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XVI (1954) 397.

Perennial. Whole plant whitish from arachnoid tomentum; root
vertical, woody, 5-8 mm thick, with short, perennial, woody,
nonflowering shoots, together with annual, leafy nonflowering shoots
forming small tussocks. Flowering shoots few, 20-30 cm high, slender,
erect, weakly branched above, with very short (1.0 to 2.5 cm long),
upright branches. Lower leaves petiolate, 2.5—4.0 cm long, lamina oval,
twice pinnately cut, lobes usually with a pair of secondary lobes, later
narrowly linear, 3—5 mm long, short-acuminate; middle cauline leaves
sessile, once or twice pinnately cut, with simple linear auricles, upper
leaves sessile, at base of branches, linear, 1.0-1.8 cm long. Capitula
sessile, oblong-ovate, up to 2.5 mm long, in spicate or spicate-racemose
inflorescence. Involucral bracts lanceolate; outer bracts white-
tomentose, inner bracts slightly longer, glabrous, only at apex pubescent,
with wide scarious margin. Florets 5; corolla yellow.

589

560

On mountain slopes.—Soviet Central Asia: Pamiro-Alai (eastern
part of Alai Range). Endemic. Described from Osh District, Type in
Leningrad.

Series 3. Knorringianae Poljak.—Low whitish-tomentose
perennials forming compact tussock; leaves once or twice pinnately
cut; capitula in simple spicate or racemose-paniculate, narrow
inflorescence. Alpine plants.

131. A. knorringiana Krasch. in Izv. Gl. Bot. Sada, XIV, 4-6
(1914) 463.

Perennial. Whole plant villous-white-tomentose till the end of
vegetative growth, 10-12 cm high. Root not very thick, vertical, with
short perennial, branched, ascending, nonflowering shoots densely
covered with remnants of leaf petioles, nonflowering shoots together
with annual and biennial, short, sterile, densely leafy shoots forming
low compact tussock; leaf rosettes at apices of shoots in several rows.
Leaves petiolate, somewhat thick, 0.5-1.5 cm long, slightly grooved
above, lamina oblong-lanceolate or more obtuse and oblong, pinnately
cut, lobes 2-3 on each side, rather approximate, usually ternately
divided, lobes cuneate, 2-3 mm long; lower cauline leaves less divided,
but upper undivided, linear, scarcely acuminate, sometimes even slightly
broadened and then carinate on inner side, with characteristic saccate
sheath at base covering capitulum from below. Peduncles many, weakly
sulcate, slender 2-3 cm high, with narrow, lax, interrupted, simple,
spicate inflorescence above. Capitula sometimes on short peduncles
enclosed by bracteal leaves enlarged at base, remote, less often 2-3,
oblong, up to 5 mm long and up to 2 mm in dia. Outer involucral
bracts broadly deltoid, with somewhat convex green keel and
scarious wings, villous-tomentose; inner bracts narrowly lanceolate,
with scarious margin, sometimes carmine-red. Florets narrowly
campanulate, sometimes constricted below, weakly lobed above, limb
fluffy, red, 1-2 mm long; stigma in young florets short, in older
florets exserted from corolla, apically reddish-brown. Achenes pyriform,
up to 2.5 mm long, circular in section, rugose-sulcate, maturing
simultaneously.

Alpine zone, on rubbly slopes of mountains.—Soviet Central Asia:
Pamiro-Alai Region (Alai Valley). Endemic. Described from Alai
Range. Type in Leningrad.

132. A. korovinii Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVII (1955) 279.

561

Perennial. Root woody, vertical with short, perennial, woody,
nonflowering ascending shoots forming small, low, lax tussock.
Flowering stems sometimes rather numerous, 15-25 cm high, slender,
angular, weakly arcuate or straight, initially whitish-tomentose, later
subglabrous, reddish below, simple or with short branches at apex,
weakly leafy. Leaves on both sides whitish-tomentose; lower most leaves
petiolate, 1.5-2.5(3.0) cm long and 0.6—0.8 cm wide; lamina oblong or
oval, once or twice pinnately cut, terminal lobes lanceolate, up to 1.5-
2.0 mm long, subacuminate; lower and partly middle cauline leaves
shorter, short-petiolate or sessile, simple pinnate, lobes undivided or
incised-toothed; upper bracteal leaves mostly simple, sessile, short-
lanceolate, acuminate, sometimes with 2 small lateral lobes at base.

*Capitula in narrow, lax raceme or racemose panicle, ovate, mature
capitula 3-4 mm long, usually sessile, very crowded. Outer involucral
bracts elliptical, hairy, inner scarcely longer, narrowly lanceolate, more
or less glabrous, with scarious margin. Florets 4-5, purple. Flowering
August.

Flat clayey places in mountains.—Soviet Central Asia: Pamiro-
Alai (Alai River valley). Endemic. Described from Alai River valley.
Type in Tashkent.

Series 4. Pauciflorae Poljak.—Flowering stems more or less
numerous, slender, brownish at the end of vegetative growth, glabrous
or white-tomentose; perennial, woody, short; nonflowering shoots
forming quite compact tussock.

133. A. pauciflora Web. in Stechm. Artem. (1775) 26; Willd. Sp.
pl. III, 3, 1828; DC. Prodr. VI, 102; Keller, V Oblasti Polupustyni,
104-109; Kryl. Fl. Alt. III, 638; Fedtsch. Perech. Rast. Turk. IV, 194;
Krascheninn. in Fl. Yugo-Vost. Evrop. Ch. SSSR, VI, 356; Kryl. FI.
Zap. Sib. XI, 2785; Poljak. in Majevski, Fl. 584.—A. maritima v.

590 stechmanniana Bess. in Bull. Soc. Nat. Mosc. VII (1834) 31.—A.
maritima var. pauciflora Ldb. FI. Ross. II, 2 (1845) 570.—A. pauciflora
Web. ssp. maikara Krasch. in Otch. o Rab. Pochv.-Bot. Otr. Kazakhsk.
Eksp. Akad. Nauk SSSR, IV, 2 (1930) 273.—A. maikara (Krasch.)
Pavl. Fl. Tsentr. Kazakhst. III (1938) 270.—Exs.: GRF Nos. 3193a,
3193b.

Perennial shrub, 10-25 cm high. Root vertical, woody, with
perennial, woody, short, prostrate or somewhat ascending nonflowering
shoots forming quite compact tussock with short, annual, leafy shoots.
Flowering stems numerous, slender, virgate, usually ascending, in
summer brown, glabrous, branched above, with upward directed slender
branches. Leaves grayish-greenish, densely hairy, later partly glabrous,

59

—

562

sometimes subglabrous, punctate-glandular; lower leaves withering’
early, petiolate, twice or thrice pinnately cut, their terminal lobes linear- —
lanceolate, very small; middle cauline leaves sessile, less divided; upper
leaves undivided, very small, linear. Capitula in narrow, compact,
paniculate inflorescence, sessile or on short peduncles, ovoid or oblong,
2 mm long, spicately crowded or somewhat widely spaced. Involucral
bracts punctate-glandular, with scarious margin; outer bracts small,
oval, more or less hairy, inner bracts larger, oblong-elliptical,
subglabrous. Florets 3; corolla yellow or purple; anthers linear, apical
appendages of anthers acute, subulate, basal appendages inconspicuous,
round; stigma lobes linear, short, truncate, short-ciliate, divergent.
Flowering August.

Abundant in steppe zone; depressions, dry ravines, river terraces
above floodline, eroded clayey foothill slopes of [mud] volcanic cones,
usually on hard crusty columnar alkaline soils—European part: Lower
Volga; Western Siberia: Upper Tobol (southern part), Irtysh (southern
part); Soviet Central Asia: Aralo-Caspian Region (northern part), Lake
Balkhash Region. Endemic. Described from the Lower Volga. Type in
Berlin.

Note. According to the observations of P.P. Poljakov, ssp. maikara,
described by I.M. Krascheninnikov, grows with A. pauciflora; and it
is distinguished only by elongated and more divergent or drooping
branches and capitula. These characters are inadequate for the rank of
subspecies and the more so for species status.

I think it is more correct to treat ssp. maikara Krasch. as an
ecological form—A. pauciflora Web. f. maikara (Krasch.) Poljak.

Economic Importance. The plants contain 0.52—0.54% essential
oil (Kazakevich and Sobolevskaya, 1928); carotene, from 4.5—5.0 (early
spring) to 11-32 (summer) mg/kg (Troitsky, 1950). On the average,
based on 10 samples, it contains (absolute dry weight, in %) Ca—
0.737; P—0.251; and on the basis of two analyses: K—0.872, Na—
1.415, Mg—0.139, S—O0.324, and CI—0.550 (Taime and others, 1948).
During the developmental stage it contains a large quantity (4.21%) of
water-soluble carbohydrates (Larin and others, 1928). According to
the data of Larin, Aleksandrovsky, Davydov, Evseev, Bannikov, and
others, it contains (based on absolute dry weight, in %) ash—7.1,
[total] proteins—12.1, soluble protein—9.3, fat—4.9, cellulose—31.3,
and nitrogen-free extractable substances—44.6. The dynamics of nu-
trients is rather well explained in the work of Shakhov and zemtsova
(1948). The nutritive value of this wormwood has been studied in
seven experiments on digestibility; of greatest interest are the
experiments of Bannikov (1940). The plants are relished by sheep,
goats, horses, and camels; they are rarely if ever eaten by cattle in
pastures (Larin, 1956).

592

563

134. A. gracilescens Krasch. and Iljin in Sistemat. Zap. Gerb.
Tomsk. Univ. 1-2 (1949) 2, 3; Kryl. Fl. Zap. Sib. XI, 2786.—A.
lercheana Ldb. Fl. Alt. IV, p. 85, quoad. pl. songor.—A. pauciflora
Kryl. Fl. Alt. III (1904) 638, partim.—Ic.: Ldb. Ic. pl. Fl. Ross. V, t.
488.

Perennial. Root thick, vertical, woody, in old plants vertically split
into several parts, with numerous, perennial, woody prostrate or weakly
ascending short vegetative shoots, forming rather compact low tussocks
together with short, annual, sterile, leafy shoots. Flowering stems usually
rather numerous, slender, 15-30 cm high, finely grayish-tomentose,
partly glabrous in fall, branched in upper half, with obliquely upright
branches, usually highly shortened. Leaves grayish-tomentose and
punctate-glandular; lower cauline leaves and leaves of nonflowering
shoots with strongly expanded base, persisting only in first half of
vegetative period, later withering, 1-2 cm long and 0.6—1.0 cm wide,
twice- or thrice-pinnately cut, with 2-3 lobes on each side, terminal
lobes filiform-linear, acuminate, or subobtuse, slightly thickened,
usually with distinct midrib beneath; middle cauline leaves less divided,
uppermost leaves undivided, small, linear. Capitula sessile, upright,
oblong, 2.0-2.5 mm long and 1 mm in dia, spicately crowded or
somewhat distant from each other, in narrowly pyramidal panicle.
Involucral bracts dorsally punctate-glandular, with scarious margin;
outer bracts small, oval-elliptical or oval, hairy, inner oblong-lanceolate,
lustrous, weakly hairy. Florets 2-5; corolla yellow; anthers narrowly
linear, apical appendages of anthers acute, basal appendages almost
undeveloped; stigma lobes short, short-ciliate, somewhat divergent.
Flowering August.

Saline, rubbly slopes and also flat lands, on alkali soils—Western
Siberia: Irtysh (southeast); Soviet Central Asia: Lake Balkhash Region
(east). General distribution: Mongolia. Described from Kulunda steppe.
Type in Tomsk.

Economic Importance. According to the data of M.I. Goryaev,
A.T. Petushkova, and N.A. Sviridova (1954), about 0.1-0.5% alkaloids
have been found in plants of this species.

135. A. terrae-albae Krasch. in Otch. 0 Rab. Pochv.-Bot. Otr.
Kazakhsk. Eksp. Akad. Nauk SSSR, IV, 2 (1930) 269; Kryl. Fl. Zap.
Sib. XI, 2787; Poljak. in Tr. Inst. Khim. Akad. Nauk Kaz. SSR, IV,
17, 18, 27, 94.—A. maritima Bess. terrae-albae Krasch. in Sistemat.
Zam. Gerb. Gl. Bot. Sada, II (1921) 187, nom.—A. suaveolens Poljak.
in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVIII (1957) 274.—
A. heptapotamica Poljak. Ibid., 278.—Exs.: GRF No. 3192.

593

564

Perennial. Whole plant white when young, later grayish-greenish
from arachnoid tomentum. Root thick, vertical, woody, in older plants
often vertically split into several parts, with perennial, woody, short,
prostrate and weakly ascending shoots forming rather compact tussock.
Annual, nonflowering shoots numerous, 3—5 cm long bearing rosettes
of leaves of summer growth. Flowering stems numerous, weak, slender,
somewhat flexuous, 15-30 cm high, branched in upper half, with usually
short branches spreading at 60—90°, or with longer, obliquely upward
directed branches. Leaves in beginning of vegetative growth wither
and fall off, once or twice pinnately cut; basal leaves 1-2 cm long,
terminal lobes small, filiform-linear, upper leaves small, undivided,
linear. Capitula pedunculate, small, 2-3 mm long, ovate, erect or
sometimes drooping, remote in lax, rather broad panicle. Involucral
bracts many-whorled, imbricate, outer bracts very small, succulent,
dorsally convex, covered with short tomentum, oval, inner much longer,
weakly convex, glabrous, with scarious border, oblong-lanceolate,
subobtuse or somewhat acuminate. Florets (4-5) at maturity with
spreading corolla, purple-pink or yellow; staminal filaments shorter
than anthers; stigma lobes short, after anthesis divergent, at apex with
dense tuft of erect cilia. Flowering August.

Mainly on clayey, brown, leached desert soils —Soviet Central
Asia: Lower Volga (Caspian Sea Region), Aralo-Caspian Region (Ust-
Urt), Kyzl-Kum, Lake Balkhash Region. General distribution:
Mongolia. Described from Betpak-Dal desert, the Sarysuk River. Type
in Leningrad.

Note. The following varieties should be recognized:

1) Var. massagetovii Krasch. (Kryl. Fl. Zap. Sib. XI, 2787)—
distinguished by relatively thicker, stronger and higher (30-40 cm)
stems, broadly oblong paniculate inflorescence, and large capitula; Lake
Balkhash Region (Muyun-Kum, Balkhash-Alakul, Zaisan);

2) Var. semiarida Krasch. and Lavr.—distinguished by fewer and
rather tall (20-35 cm) stems, narrowly pyramidal panicle, and obliquely
upright branches; desert-steppes: Lake Balkhash Region (Karaganda
Region, Zaisan);

3) Var. suaveolens Poljak.—distinguished by short (8-15 cm high)
stems, large capitula that are 3-4(5) mm long and 2-3 mm wide, usually
approximate, upright; southern Lake Balkhash Region, Ili River valley;

4) Var. heptapotamica Poljak.—distinguished by narrow, rather
dense, pyramidal panicle; northern Tien Shan, intermontane plains.

5) Var. kurdaica Poljak.—distinguished by thick stems, short
branches, and dense pubescence on entire plant; according to the data
of R.N. Sazonova (1959), the plants collected near Kurdai Pass (Trans-
Ili Alatau) contain 2.04% santonin; the yield of essential oil was 1.8%;
the oil contains aldehydes and cineole.

594

565

Economic Importance. According to the data of M.I. Goryaev
(1954), the essential oil of A. terrae-albae Krasch. contains cineole-
8%, 1-camphor—37%, cumaldehyde—6%; borneole—7%, and
presumably artemisiaketone* —about 14%. According to the data of
E.I. Saudarova, the essential oil of A. terrae albae var. massagatovii
Krasch. contains the following components: 1) carbonyl compounds
(camphors 95.36% by weight of the oil, and carvone comprising 1.19%
by weight of the oil; this substance was found for the first time in
wormwood essential oils); 2) alcohols and complex esters: Cis-
pikomfeol’—0.6%; Trans-pikomfeol’—0.79%; bornyl ester of isova-
leric acid—0.55%; crystalline alcohol of unknown structure with a
melting point of 47%[sic.], comprising 0.48% of the oil by weight; 3)
phenol—0.05%, and 4) acid 0.09%. According to the data of R.N.
Sazonova, it contains up to 0.37% santonin.

According to the data of Makhmudov, the nutritive value of this
wormwood, during the vegetative growth phase, is close to that of oats
(Larin). One hundred kilograms of absolutely dry fodder contains 9.4
kg of digestible protein and 93.8 feed units. At the flowering stage, the
fodder contains 2.8 kg of digestible protein and 46.5 feed units. In
winter its nutritive value is low: 2.5 kg of digestible protein and 24.7
feed units. In the opinion of Larin, this wormwood is greatly relished
by sheep in the spring, satisfactorily to poorly in the summer, and
greatly in the fall and winter. Horses and camels relish it less. The
data about its feed value for cattle are inadequate.

According to the studies of M.I. Goryaev, E.I. Glyadkovskaya and
I.M. Shabanov (1951), a mixture of the three species (A. terrae albae
Krasch., A. serotina Bge., and A. leucodes Schrenk), steam-processed
to extract the essential oil, does not lose its fodder value and is eaten
by big horn cattle better than the mixture containing the essential oil.
Steam treatment removes from the fodder the substances that impart
poor taste for the animals. An experiment of making silage from the
green mass (after removal of the essential oil) of succulent feeds
(bagasses, watermelon, pumpkin) showed that the fodder is entirely of
good quality for cattle.

Series 5. Lessingianae Poljak.—Plants grayish from pubescence,
later subglabrous, sometimes completely glabrous or even whitish-
tomentose; lower leaves pinnatisect or twice pinnately cut, terminal
lobes narrowly linear, acute.

*The name of the compound could not be confirmed — General Editor.

Nn

566

136. A. lessingiana Bess. in Linnaea, XV (1841) 90, 103; Fedtsch.
Perech. Rast. Turk. IV, 195; Krascheninn. in Fl. Yugo-Vost. Evrop.
Ch. SSSR, VI, 356.—A. taurica Less. in Linnaea, IX (1834) 170, 190,
non Willd.; Ldb. Fl. Ross. II, 2 (1846) 575, pro parte, quoad plant.
Lessing. (v. spec. auth.).—Exs.: GRF No. 3194a, b.

Perennial. Plant pubescent, later subglabrous. Root thick, woody,
with perennial, weakly ascending, woody, short, nonflowering shoots
forming rather dense, lax, small tussocks with short, young, leafy shoots.
Flowering stems rather numerous, ascending or almost erect, 18—30(40)
cm high, branched above, with short, upright branches. Most leaves
weakly hairy or glabrous, or arachnoid-tomentose (f. canescens
Krasch.); leaves of vegetative shoots and lower cauline leaves petiolate,
2-5 cm long, their laminas oblong-oval, once or twice pinnately cut
terminal lobes linear-filiform, 5-10 mm long, keeled at base, green;
upper cauline and bracteal leaves smaller, simple, narrowly linear or
with a pair of small lateral lobes at base. Capitula sessile or on short
peduncles, oblong-ovoid, 3-4 mm long, in narrowly racemose-paniculate
inflorescence. Outer involucral bracts small, oval-lanceolate; inner
bracts longer, linear-lanceolate, with scarious margin. Florets 5-6,
corolla purple-pink or yellow; anthers linear, apical appendages of
anthers subulate, basal appendages short-acuminate; style at maturity
of achenes as long as stamens; stigma lobes linear, erect, truncate,
short-ciliate. Flowering August.

Steppe region, on rubbly slopes of [mud] volcanic cones, chalk
outcrops, sometimes on chestnut, solonetzic and clayey soils.—West-
ern Siberia: Upper Tobol (southern part of the Urals and the whole of
Syrt.); Soviet Central Asia: Aralo-Caspian Region (Mugodzhary). En-
demic. Described from Orenburg Region. Type in Kiev.

According to the data of M.I. Goryaev, T.E. Serkebaeva, L.A.
Ignatova, and I.M. Shabanov (1951), the yield of essential oil from the
air-dried matter is 0.46%. The oil contains 59.4% carbonyl compounds,
of which camphors constitute 43.8%; the carbonyl liquid compounds
of aldehyde nature are tentatively related to methylheptane.

137. A. aralensis Krasch. in Spisok Rast. Gerb. Fl. SSSR, X (1936)
105.—Exs.: GRF No. 3196.

Perennial. Plant 30-50 cm high, grayish-green in early vegetative
growth period from more or less dense, arachnoid hairs, later glabrous,
with distinct lemony fragrance. Root rather thick, woody, vertical, with
perennial, short, branched, woody, nonflowering shoots covered with
grayish bark. Flowering shoots numerous, slender, virgate, (20)25—40
cm high, in lower part stramineous-brownish smooth, in upper half
green, branched with short, more or less divergent branches. Leaves

596

567

hairy, later subglabrous, densely glandular, yellowish-green, 2-3 cm
long, lamina oblong-lanceolate, twice pinnately cut, terminal lobes
remote, succulent, 4-5 mm long and 0.5—1.0 mm wide, linear, short-
acuminate; leaves of vegetative shoots and lower cauline leaves
petiolate; middle cauline leaves sessile; upper leaves undivided, linear.
Capitula pedunculate or sessile, ovate-oblong, 2-3 cm long, remote or
on apices of branches, 2-3, drooping or even divergent, in oblong-
pyramidal panicle. Involucre many-whorled, strongly glandular; outer
involucral bracts small, round, dorsally convex, pubescent, inner bracts
lanceolate or oval-oblong, glabrous, greenish, with scarious border.
Florets 3-4; corolla yellow; anthers linear; apical appendages of anthers
acute, basal appendages smaller, acute; style short; stigma lobes linear,
erect, apically truncate, short-ciliate. Flowering August.

Clayey, sandy loam, solonetzic soils.—Soviet Central Asia: Aralo-
Caspian (northern Aral) Region. Endemic. Described from southeastern
part of Aktyuba Region, Kara-Chokat. Type in Leningrad.

138. A. camelorum Krasch. in Otch. 0 Rab. Pochy.-bot. Otr.
Kazakhsk. Eksp. Akad. Nauk SSSR, IV, 2 (1930) 272.

Perennial shrub 12—15(30) cm high, more or less hairy, later
subglabrous. Root 4-8 mm thick, woody, with perennial, woody,
prostrate or ascending, strongly branched, nonflowering shoots forming
small tussocks. Flowering shoots numerous, slender, strong, straight or
basally arcuate, branched above, with short, divergent branches. Lower
leaves petiolate, 1-2 cm long, oblong-linear, twice pinnately cut, lobes
with 2-3 linear, short, 1-2 mm long, acuminate secondary lobes; middle
cauline leaves smaller, simple pinnate, upper leaves undivided, linear,
sometimes with a pair of small lateral lobes at base. Inflorescence
oblong, paniculate. Capitula on short peduncles or sessile, narrow-
oblong, 2-3 mm long, at maturity divergent, more or less remote.
Involucre 3-rowed; outer involucral bracts small, round, grayish-
pubescent, inner bracts oblong or prolate, with scarious, brownish-red,
lustrous margin. Receptacle rather small, convex. Florets 2—4; corolla
yellow; anthers linear, apical appendages of anthers obtuse, basal
appendages weakly developed, roundish; style shorter than stamens;
stigma lobes linear, erect, truncate, short-ciliate. Flowering August.

Depressions, ravines, old river terraces, puffy salt marshes, cloddy
solonetz soils, as well as on outcrops of colored saline clays.—Soviet
Central Asia: Aralo-Caspian Region, Lake Balkhash Region (western
part). Endemic. Described from Turgai (Aktyuba Region). Type in
Leningrad.

Series 6. Sublessingianae Poljak.—Leaves simple pinnate or usu-
ally twice or even thrice pinnately cut into narrowly linear or linear

597

568

lobes; inflorescence narrowly paniculate; plants glabrous or subglabrous
only at end of vegetative growth.

139. A. sublessingiana (Kell.) Krasch. ex Poljak. in Bot. Mat.
Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954) 395; Krascheninn. in
Spisok Rast. Gerb. Fl. SSSR, X, No. 3195; Kryl. Fl. Zap. Sib. XI,
2788.—A. maritima var. sublessingiana Kell. Bot. Geogr. Issled. Zais.
u. Semipalat. Obl. II (1912) 89-93, nomen.—A. lessingiana Kryl. FI.
Alt. III (1904) 640, non Bess.—A. gorjaevii Poljak. op. cit. 420.—A.
polysthicha Poljak. ibid., 420.—Ic.: Tr. Inst. Khim. Akad. Nauk Kaz.
SSR, IV (1959) 92.—Exs.: GRF No. 3195.

Perennial. Tap root woody, with short, perennial, ascending, woody,
nonflowering shoots covered with brownish bark shredding into fibers.
Flowering shoots erect, usually numerous, 30-45 cm high, arachnoid-
hairy, later subglabrous, particularly in lower part, branches above,
with somewhat divergent branches. Leaves grayish, hairy, later
subglabrous; lower leaves long-petiolate, 3-4 cm long, their laminas
oblong-oval, twice pinnatisect, terminal lobes linear-filiform, 5-12 mm
long, subacuminate; middle cauline leaves sessile, pinnate, with
palmatisect, sometimes pinnatisect auricles; uppermost bracteal leaves
undivided, linear. Capitula sessile, oblong-ovate, 2-3 mm long, mostly
approximate, in groups of few in rather narrow panicle. Involucral
bracts oblong, slightly convex, dorsally densely hairy; inner bracts
with wide scarious border. Florets 3-8; corolla yellow, anthers on
short filaments, linear, apical appendages of anthers acute, basal
appendages shorter, acute; style short. Receptacle quite small, convex.
Flowering August.

Rubbly and stony slopes of [mud] volcanic cones.—Western
Siberia: Irtysh (south), Altai (southwest); Soviet Central Asia: Lake
Balkhash Region, Aralo-Caspian (only northeastern) Region,
Dzhungaria-Tarbagatai. Endemic. Described from southern Lake
Balkhash Region. Type in Leningrad.

According to the data of M.I. Goryaev, A.T. Petushkova, and N.A.
Sviridova, about 0.1-0.5% alkaloids were found in samples of A.
sublassingiana.

Note. The following varieties should be recognized:

1) Var. gorjaevii Poljak.—distinguished by slender stems that are
usually 20-30 cm high and leaf lobes that are narrowly linear, short
(1-3 mm) and triangular; foot-hills of Dzhungarian Alatau and
Tarbagatai. According to the studies of R.N. Sazonova, it contains up
to 1.24% santonin; the essential oil contains up to 70% cineole and
many azulene-forming sesquiterpenes.

598

569

2) Var. polysthichum Poljak.—distinguished by the lower cauline
leaves that are smaller (up to 2 cm and not 3-4 cm), small leaf lobes,
and 3, rarely 2, stigmatic lobes; Chu-Ili Mountains.

140. A. transiliensis Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVI (1954) 417; Poljak. in Tr. Inst. Khim. Akad. Nauk
Kaz. SSR, IV (1959) 70.—Ic.: Poljak. ibid., 71.

Perennial. Root woody, thick, with short, ascending, perennial,
woody nonflowering shoots covered with brownish cracking bark.
Flowering shoots 40-65(75) cm high, rather numerous, erect, grayish,
arachnoid-hairy, later, particularly in lower part, subglabrous, brownish,
branched above, with moderately divergent long branches. Leaves
grayish-green, hairy; lower cauline leaves and leaves of vegetative
shoots withering early, long-petiolate, 3.5—6.0 mm long, oblong, twice
pinnately cut, terminal lobes narrowly linear, 4-8 mm long, with short
stiff apices; middle cauline leaves smaller, short-petiolate, with pinnately
cut auricles; upper bracteal leaves simple, very small, linear. Capitula
pedunculate, upright or divergent, oblong-ovoid, 3 mm long, somewhat
remote or in groups of few in narrow, dense panicle. Outer involucral
bracts small, oval, densely hairy, inner larger, oblong, green, with
occasional hairs, mixed with white cystiform hairs, with broad scarious
margin. Receptacle about 1 mm, obtusely conical. Florets 3-5; corolla
pink or yellow; stamens and style at maturity exserted; anthers linear,
apical appendages of anthers subulate, basal appendages shorter,
subobtuse, stigma lobes weakly divergent, with tuft of short erect cilia
at apex. Achenes somewhat flat, pyriform, up to 1.5 mm long. Flowering
August.

Loessial meadow-steppe slopes of foothills.—Soviet Central Asia:
Tien Shan (Trans-Ili Alatau). Endemic. Described from vicinity of
Alma-Ata. Type in Leningrad.

Note. Var. boamensis Poljak., characterized by taller stems and a
narrowly pyramidal panicle, should be recognized. According to the
data of R.N. Sazonova, the plants of this variety growing in the Boam
gorge (Kirghiz Alatau) contain up to 5.124% santonin.

Economic Importance. According to the data of M.I. Goryaev
(1952), the yield of essential oil was 0.534% in terms of absolute dry
matter. The oil contains cineole—68%, free organic acids—7%,
phenols—4.65%, aldehydes and ketones—16%. In samples of this
wormwood, about 0.1-0.5% alkaloids was found (Goryaev, Petushkova,
and Sviridova). The santonin content in this species is not any lower
than in A. cina (R.N. Sazonova).

According to the data of M.I. Goryaev and M.G. Pugachev (1955),
the composition of the essential oil is: formic acid (0.1%), isobutyric

599

570

acid (0.086%), m-cresol (1.5%), a-pinene (traces), cineole (55-56%),
thujyl alcohol (4-5%), complex ester of thujyl alcohol and an
unidentified acid (6—-7%), cumaldehyde (7.5%), thujone (14.0—17.5)
(mixture of a- and B-forms), and sesquiterpenic alcohol (not identified).
The oil extracted from the distillate contains formic acid (12.33%),
isobutyric acid (27.27%), phenols (56.7%), and cineole (5.2%). In the
multipurpose exploitation of the plant, its essential oil may be used for
extracting cineole and as a surface-active substance.

141. A. tenuisecta Nevski in Tr. Bot. Inst. Akad. Nauk SSSR, ser.
1, No. 4 (1937) 284.—A. grigorievi Krasch. ex Nevski, nomen in ibid.,
284.—A. karatavica Krasch. and Abol. ex Poljak. in Bot. Mat. Gerb.
Bot. Inst. Akad. Nauk SSSR, XVI (1954) 397.—A. glaucina Krasch.
ex Poljak. ibid., 413.—A. mucronulata Poljak. ibid., 419.—Ic.: Tr.
Inst. Khim. Akad. Nauk Kaz. SSR, IV (1959) 90, sub A. glaucina.

Perennial. Semishrub, 35—60(70) cm high, with thick, strongly
branched, short, perennial, woody, nonflowering shoots covered with
brownish-gray cracking bark. Flowering shoots numerous, erect or
ascending, slender, virgate, 15—-40(50) cm high, grayish from dense
arachnoid hairs, later partly glabrous, branched above, with obliquely
spreading branches. Leaves grayish, densely arachnoid-hairy, later more
or less glabrous, green; lower cauline leaves short-petiolate, 1.5—2.5
cm long, lamina broadly oval, thrice pinnately cut, terminal lobes
narrowly linear, 2.0-3.5 mm long; upper leaves sessile, once or twice
pinnately cut, uppermost leaves undivided, linear. Capitula sessile or
subsessile, ovoid-oblong, up to 3 mm long, remote or spicately crowded
on branches in narrow panicles. Involucral bracts pale brownish, smooth,
with scarious margin. Florets 3—7; corolla yellow or pinkish; anthers
on shorter filaments, linear, apical appendages of anthers narrow, almost
subulate, basal appendages very short, obtuse; stigma lobes short, linear,
obtuse, with erect cilia. Flowering August.

Clayey and rubbly-clayey mountain slopes.—Soviet Central Asia:
Tien Shan, Pamiro-Alai Region. Endemic. Described from southeastern
Turkmenia, Kugitang Range. Type in Leningrad.

Note. The following varieties should be recognized:

1) Var. karataviensis Poljak.—distinguished by bi- or almost
tripinnatisect lower leaves, longer (4-6 mm long and not 2.0—-3.5 mm)
terminal leaf lobes and a relatively weak pubescence over the whole
plant; found in Syr-Darya Region of Karatau. According to the data of
M.I. Goryaev and R.N. Sazonova (1959), the plant contains 0.22-
0.33% santonin. It is interesting that some samples of this variety did
not contain santonin. The yield of essential oil varies from 0.6 to
1.7%. The oil contains a large percentage of camphors as well as

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ant

aldehydes, ketones, phenols, azulene-forming sesquiterpenes, alcohols,
acids, and complex esters. Variation in the contents of organic acids
and esters is especially noticeable.

2) Var. glaucina Poljak.—distinguished by bipinnatisect lower
leaves and sparse pubescence; Tien Shan (Chatkal Range), Pamiro-
Alai (Turkestan Range). According to the studies of M.I. Goryaev and
R.N. Sazonova (1959), the plant contains 1.2—1.9% santonin. The
essential oil contains up to 70% cineole. Alkaloids are also present
(M. Goryaev, G.K. Kruglykhina, and E.I. Satdarova, 1959).

3) Var. mucronulata Poljak.—distinguished, in general, by shorter
stems (15-25 cm high), capitula with fewer (3-4) florets, and abundant
pubescence of the whole plant, persisting up to the end of the vegetative
phase; typical of the highest mountains of the Syr-Darya part of Karatau.

142. A. kaschgarica Krasch. in Tr. Bot. Inst. Akad. Nauk SSSR,
Ser. 1, No. 3 (1936) 350.

Perennial. Plant grayish from dense arachnoid hairs in early
vegetative phase, later subglabrous. Root vertical, woody, rather thick,
with perennial, woody, nonflowering shoots. Flowering shoots more or
less numerous, stiff, erect, up to 30-35 cm high, branched from base
or in lower part, with slender, virgate, long (lower branches up to 10-
25 cm), more or less arcuate branches. Leaves stiff, somewhat thick,
highly variable; lower and middle cauline leaves 1.5—2.0 cm long,
once or twice pinnately cut, sometimes simple, linear, sub-obtuse or
divided into 2—3 lobes; upper leaves simple, linear. Capitula sessile or
on short peduncle, ovoid, cuneately narrowed at base, 4-5 mm long,
in rather wide, subglobose panicle. Involucral bracts green at maturity,
strongly recurved and easily detached, with scarious border; outer bracts
small, oval, dorsally convex, inner larger, oblong-lanceolate or
lanceolate. Florets 4-5; corolla purple-red; anthers linear, on more or
less long filaments, apical appendages of anthers almost subulate,
acuminate, basal appendages smaller, subobtuse; stigma lobes linear,
truncate, short-ciliate, after anthesis divergent. Flowering August.

Stony-rubbly slopes of lower mountains, rubbly desert plains.
Sometimes on gravel beds of rivers.—Soviet Central Asia: Lake
Balkhash Region (near Zaisan, Takhta and Dzharkent). General
distribution: Dzhungaria-Kashgaria. Described from northern Kashgar.
Type in Leningrad.

Economic Importance. According to the data of M.I. Goryaev
and R.N. Sazonova (1959), it contains 1.5% essential oil; aldehydes,
ketones, cineole, and azulene-forming sesquiterpenes were found as
constituents of the oil.

60

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572

143. A. amoena Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVI (1959) 421.—Ie.: Tr. Inst. Khim. Akad. Nauk Kaz. SSR,
IV (1954) 91.

Perennial. Plant 18-30 cm high, arachnoid white-tomentose up to
the end of vegetative phase. Root thick, woody, with perennial, short,
prostrate or ascending, woody, non-flowering shoots forming low, rather
compact tussocks together with short, young, leafy shoots. Stems usually
10-15, erect, 15—28 cm high and up to 1.5 mm thick, branched above,
with short (up to 2-3 cm long), upright branches. Leaves of
nonflowering shoots and lower cauline leaves long-petiolate, 4-8 cm
long, lamina oval, once or twice pinnately parted; lobes 3-5 pairs;
terminal lobes linear, 4-12 mm long and 1 mm wide, flat with
cartilaginous apex; middle cauline leaves smaller, sessile or short-
petiolate, simple pinnatipartite, with simple linear auricles; uppermost
leaves linear, up to 4 mm long. Capitula sessile, ovoid, up to 4 mm
long and 2 mm in dia, borne in narrowly spicate or spicate-paniculate
inflorescence. Involucral bracts with cystiform hairs, with scarious
border; outer bracts oval, densely hairy, inner larger, oblong-elliptical,
somewhat flat, green dorsally, subglabrous, only with occasional hairs
at apex. Florets 4—5, corolla yellow or pink; anthers on long filaments,
linear, apical appendages of anthers subulate, basal appendages almost
undeveloped; stigma lobes and style thickened along margin, divergent,
with tuft of strongly divergent cilia at apex. Flowering August.

Stony slopes of granitic lower mountains.—Soviet Central Asia:
Lake Balkhash Region (southern edge of Altai Mountains). Endemic.
Described from Marka-Kul District. Type in Leningrad.

Economic Importance. According to the data of M.I. Goryaev,
A.I. Petushkova, and N.A. Sviridova, he plant contains about 0.1-
0.5% alkaloids. According to the studies of R.N. Sazonova (1952), the
plant contains 0.79% santonin. The essential oil has a high cineole
content; the yield of essential oil is 0.79%.

144. A. fedtschenkoana Krasch. in Tr. Bot. Inst. Akad. Nauk
SSSR, Ser.1, No. 3 (1936) 351; Bot. Mat. Gerb. Gl. Bot. Sada, II, 44,
186, nomen.—A. issykkulensis Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVII (1955) 415.

Perennial. Root woody, vertical, 3—8(15) mm thick, with short,
perennial, woody, ascending nonflowering, shoots forming tussocks.
Flowering shoots very numerous, 16—30(35) cm high, slender, virgate,
erect, grayish, hairy, later glabrous, branched above, with short, usually
up to 2-6 cm long, slender, more or less weakly divergent branches.
Leaves of nonflowering shoots and lower cauline leaves hairy, petiolate,
1.5-3.0 cm long, lamina oval or oblong, 10-12 mm wide, twice

573

pinnately cut, with 2—4 pairs of lobes, terminal lobes linear or linear-
spatulate, somewhat succulent, 2-3 mm long, with short, cartilaginous
apex; middle cauline leaves smaller, sessile, pinnatisect or simple,
linear; upper leaves small, linear. Capitula sessile, ovate, 2.3-3.0 mm
long, erect, remote, in narrowly paniculate inflorescence. Involucral
bracts greenish dorsally, hairy or glabrous, lustrous, with brownish
scarious border; outer bracts small, oval-deltoid, inner bracts larger,
oblong-lanceolate. Florets 2-3-5. Achenes brown, 1.0—0.5 mm long.

Alkaline steppes, meadows.—Soviet Central Asia: Tien Shan (Tersk
Alatau). General distribution: Dzhungaria—Kashgaria. Described from
southeastern Kashgaria. Type in Leningrad.

Note. Var. issykkulensis Poljak., which is distinguished by some-
what thickened stems, a many-headed, narrowly pyramidal panicle,
capitula with 3—5 (and not 2-3) flowers and large achenes (up to 1 mm
long and not 0.5 mm), should be recognized. Central Tien Shan (Issyk-
Kul Basin).

145. A. cina Berg. ex Poljak. in Tr. Inst. Khim. Akad. Nauk Kaz.
SSR, IV (1959) 69.—A. cina Berg. in Darstell. IV (1863) tab. 29°,
nomen provisorium; Fedtsch. Perech. Rast. Turk. IV, 194.—A. contra
Schachn. Iss]. Tsitv. Polyni (1899).—A. maritima Bess. f. fragrans
Rgl. in herb.—A. mogoltavica Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVII (1955) 416.—Ic.: Shakhnazarov (1899) 13;
Tr. Inst. Khim. Akad. Nauk SSSR, IV, 88.

Perennial. Plant 25—35(40) cm high, arachnoid-hairy in early
vegetative phase, later glabrous. Root vertical, woody, with short,
perennial, woolly, ascending; non-flowering shoots covered with
brownish-gray cracking bark. Flowering shoots numerous, 18-—30(40)
cm high, rather slender, virgate, erect, sometimes basally arachnoid-
hairy, in second half of summer brown, glabrous, lustrous, branched
above, with very slender, erect branches. Leaves of vegetative shoots
and lower cauline leaves withering early, long-petiolate, 2.5-4.0 cm
long, lamina oval, twice pinnately cut, terminal lobes narrowly linear,
2-5 mm long, short-acuminate; middle cauline leaves subsessile, less
divided, with pinnately cut auricles; uppermost leaves sessile, undivided,
linear, small. Capitula sessile, erect, oblong-ovoid, up to 3 mm long,
spicately crowded on branches in narrow, compact panicle. Involucre
many-rowed, imbricate; involucral bracts smooth, lustrous, green, with
scarious margin; outer bracts small, oval; inner bracts larger, oblong.
Florets 3-5; corolla yellow or purple; anthers on short filaments, linear,
apical appendages of anthers acuminate, basal appendages shorter,
acute; stigma lobes oblong, weakly divergent, with tuft of erect cilia
at apex. Achenes about 1-2 mm long, ovoid, somewhat flat. Flowering
August.

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574

Flat plains, ravines, terraces above the floodplains, saline soils.—
Soviet Central Asia: Syr-Darya, Tien Shan. Endemic. Described from
southern Kazakhstan. Type in Leningrad.

Note 1. The name A. cina Berg. is not valid because it is not
accompanied by a description and is indicated as a provisional name
(nomen provisorium) uniting several species of Seriphidium with
capitula having anthelmintic properties. Moreover, in the words of the
author, Artemisia cina Berg. unites three trade cultivars:

1) Levant Stock.—Cinae levantici—a medicinal cultivar of
unknown origin. It grows in Persia and Bukhara; it was collected there
and brought to Petersburg or Moscow by the caravans of Russian
traders. This cultivar has been considered as one of the best.

2) Russian santonica stock—Cinae rossici— collected from the
steppes along the banks of the Volga River near Sarepta and Saratov.
Two varieties are distinguished: a) capitula of Artemisia pauciflora
and A. monogyna; and b) capitula of A. lercheana.

3) Cinae barbarici, consisting of A. ramosa which grows in north-
eastern Africa.

Besides similar explanations, in the particular work on plate No.
29 there are illustrations of a transverse section of the capitulum and
of the inner involucral bract, referred by the author to A. cina. However,
these scanty details, in my opinion, are not sufficient criteria to accept
the above name. On the basis of these considerations, I myself compiled
the description of this, the santonica wormwood.

Economic Importance. It is the main raw material for obtaining
an anthelmintic preparation, viz. santonin.

According to the data of M.I. Goryaev, L.A. Ignatova, and I.M.
Shabanov (1953), the presence of 6% carbonyl compounds, mainly as
a mixture of a- and B-thujone, has been established for the first time
in the essential oil of A. cina (Berg.) Poljak. The presence or absence
of a small quantity of thujone in some oil samples is possibly because
its accumulation occurs at a much later stage of the plant’s development,
when the processes of assimilation and growth slow down considerably.
The oil also contains 5—7% of thujyl alcohol. Cineole is the major
constituent (up to 80%) of the oil. It contains a poisonous substance—
santonin C,.H,,O,; traces of alkaloids have also been found
(Massagetov, 1947). The dried capitula under the name santonica seed
(from the Italian “Semen Zina’) are used in medicine as an excellent
anthelmintic. The reports about the edibility of this wormwood for
cattle are contradictory.

Note 2. Var. mogoltavice Poljak. should be recognized. It is
distinguished by more or less elongate and divergent shoots, usually
somewhat smaller lower leaves and hairy involucral bracts. It grows

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575

near the Mogoltau Mountains (Mirzarobad natural boundary) and con-
tains 2.6—2.9% santonin. According to the studies of M.I. Goryaev,
I.M. Shabanov, and L.A. Ignatova, the essential oil of var. mogoltavica
Poljak. contains: 1) 30.77% carbonyl compounds, of them 17.44%
camphors and the rest consisting of a mixture, which presumably
includes thujone; 2) cineole—12.3%; 3) free alcohols 8.72% and bound
alcohols as esters—10.04%. It contains almost the same quantity of
santonin as A. cina (Berg.) Poljak. Thickets of this variety were used
until 1952 as raw material for obtaining santonin. At present, their
area has shrunk considerably.

Note 3. N.V. Pavlov (Rastitelnye Resursy Yuzhnogo Kazakhstana
[Plant Resources of Southern Kazakhstan], Alma-Ata, 1947) reports a
new species, A. cinoides Pavl., collected from the Chu-Ili Mountains,
which presumably contains 0.24% santonin by dry weight. This report
remains a puzzle, since herbarium specimens of this plant are unknown,
even a description of it is missing.

146. A. ciniformis Krasch. and M. Pop. ex Poljak. in Bot. Mat.
Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954) 403.

Perennial semishrub 30-45 cm high, subglabrous. Tap root, thick,
woody, with few short, perennial, woody, ascending, nonflowering
shoots covered with grayish-brown cracking bark. Flowering shoots
10-20 and more, erect or basally arcuate, hairy, later glabrous, smooth,
stramineous, branched above, with short (mostly 2-5 cm long),
somewhat divergent branches. Leaves of nonflowering shoots and lower
cauline leaves most often 1.5-3.0 cm long, with petioles thickened at
base, lamina broadly round, twice or thrice pinnately cut, terminal
lobes filiform-linear, 2-8 mm long, short-acuminate with pinnately or
palmately cut, or even linear auricles; middle cauline leaves subsessile,
twice pinnately cut; upper leaves undivided, linear. Capitula sessile,
upright or weakly divergent, ovoid, 3-4 mm long, loosely spicate on
branches in narrowly paniculate inflorescence. Involucre many-whorled,
imbricate; outer bracts small, hairy, inner larger, oblong, glabrous,
with wide, scarious margin. Florets 4-5. Flowering August to
September.

Clayey, rubbly and stony mountain slopes as well as foothills.—
Soviet Central Asia: mountainous Turkmenia. Described from Kopetdag.
Type in Leningrad.

According to the data of V.A. Vyshensky, 0.4—0.5% essential oil
was obtained from plants up to the flowering stage. The total quantity
of aldehydes in the oil was up to 16%. A certain quantity of santonin
is also present. According to I.V. Larin, it is grazed in pastures during
the fall and winter. It contains some quantity of santonin.

576

147. A. glanduligera Krasch. ex Poljak. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XVI (1954) 402. .

Perennial. Plant 40-65(75) cm high, usually hairy in early
vegetative phase, later glabrous, smooth, punctate-glandular. Root
vertical, woody, up to 1-5 cm thick, with perennial, erect, woody,
nonflowering shoots up to 15 cm long, covered with brown, cracking
bark. Flowering shoots 30—50(60) cm high, usually numerous, erect,
strong, virgate, stramineous, branched above, with few divergent, short
or long branches. Leaves of nonflowering shoots and lower cauline
leaves petiolate, 2-3 cm long, lamina oblong, twice or thrice pinnately
cut, terminal lobes (2)3—6(7) mm long, linear, short-acuminate; middle
cauline leaves subsessile, once or twice pinnately cut, with pinnately
cut auricles; uppermost leaves undivided, linear-lanceolate, usually
exceeding capitula, keeled. Capitula sessile, at maturity oblong, up to
3 mm long and 2 mm in dia, upright, approximate, or somewhat remote
in narrow, compact paniculate inflorescence. Involucre many-whorled,
imbricate, outer bracts small, oval, hairy, inner bracts larger, oblong-
oval, green dorsally, with wide scarious border. Florets 5; corolla
yellow; anthers linear, apical appendages of anthers subulate, basal
appendages shorter, sub-acute; style shorter than stamens; stigma lobes
short, linear, truncate, ciliate. Flowering September.

Rubbly and clayey mountain slopes.—Soviet Central Asia: Pamiro-
Alai Region. Endemic. Described from Alai Range. Type in Leningrad.

Series 7. Validae Poljak.—Plants usually not forming tussock,
grayish, densely arachnoid-hairy in early vegetative phase, later
subglabrous; stems branched with more or less long, divergent branches,
lower leaves mostly 3-8 cm long, twice pinnately cut, their terminal
lobes linear, usually 3-8 mm long.

148. A. ferganensis Krasch. ex Poljak. in Bot. Mat. Gerb. Bot.
Inst. Akad., Nauk SSSR, XVI (1954) 409; Poljak. in Tr. Inst. Khim.
Akad. Nauk Kazakh SSR, IV (1959) 119.

Perennial. Plant grayish from woolly tomentum, partly glabrescent
by end of vegetative growth. Root thick, woody, vertical with rather
short, perennial, woody non-flowering shoots. Flowering shoots few,
sometimes very numerous, basally ascending or erect, 40—60(70) cm
high, branched above, with long (up to 10-15 cm), strongly divergent
branches. Lower leaves long-petiolate, 4-7(8) cm long, lamina oblong,
twice pinnately cut, lobes remote, terminal lobes flat, linear, (3)6—
9(12) mm long and 0.5—1.0(1.5) mm wide, apically acute or with 2—
3 teeth; middle and upper leaves sessile, less divided, with pinnate
auricles; upper leaves undivided, linear. Capitula ovate-oblong,

605

way
SS vie
LU

Plate XXIX.
1 — Artemisia demissa Krasch., lower cauline leaf, middle and upper cauline
leaves, outer involucral bract, inner involucral bract, peripheral pistillate floret,
rudiment of pistil of disk floret, achene. 2— A. ledebouriana Bess., upper part
of plant, achene, outer involucral bract; inner involucral bract, lower cauline
leaf. 3—A. borealis Pall., habit, staminate disk floret, rudiment of pistil of
disk floret, peripheral pistillate floret, achene, inner involucral bract; outer
involucral bract, lower cauline leaf, middle cauline leaf, upper cauline leaf.

608

578

3.0-3.5(4.0) mm long, usually upright, remote or approximate or
branches in broadly pyramidal panicle. Involucre imbricate, densely
whitish-pubescent; involucral bracts with scarious margin, outer small,
oval, inner much larger, elliptical or oblanceolate. Florets 5-6 pink.

Plains as well as foothills and montane regions along river valleys,
on saline clayey soils, sometimes on sands, gravelly or rubbly-clayey
deposits.—Soviet Central Asia: Kyzl-Kum, Amu-Darya, Syr-Darya, Tien
Shan, Pamiro-Alai. Endemic. Described from Dzhalalabad District.
Type in Leningrad.

Economic Importance. As reported by Agababyan, Kasimenko,
and Granitov (1934), it is the main pasture fodder for sheep and cam-
els in fall and winter but poor fodder for other types of stock.

According to the data of M.I. Goryaev, G.K. Kruglykhina, and
E.I. Satdarova (1959), the plant contains alkaloids.

149. A. porrecta Krasch. ex Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVI (1954) 410; Poljak. in Tr. Inst. Khim. Akad.
Nauk Kaz. SSR, IV, 18, 27; Nevski in Tr. Bot. Inst. Akad. Nauk
SSSR, ser. 1, No. 4, 284, No. 201, nomen.

Perennial. Plant grayish to grayish-glaucous-green from dense
pubescence in early vegetative period, later glabrescent. Tap root thick,
woody, with short, perennial woody shoots covered with brown cracking
bark. Flowering shoots very numerous, 45—60(70) cm high and up to
2—4(5) cm thick, strong, erect or basally ascending, arachnoid-hairy,
later partly subglabrous, stramineous, branched above, with long, up
to 7-15 cm long, obliquely upright, straight branches. Lower cauline
leaves withering early, long-petiolate, up to 5 cm long, lamina oval-
oblong, twice or thrice pinnately cut, terminal lobes flat, linear, 3—5(7)
mm long, scarcely pointed, with prominent midrib; middle cauline leaves
usually sessile, with pinnate auricles, their laminas once or twice
pinnately cut, terminal lobes linear, 5—10(15) mm long, uppermost
leaves undivided, linear, usually 2-3 mm long. Inflorescence broadly
pyramidal panicle; capitula sessile, ovoid, 3—4 mm long, approximate,
less often somewhat remote. Involucre imbricate; involucral bracts
convex, punctate-glandular and whitish-pubescent, with scarious margin;
outer bracts small, oval, inner considerably longer, oblanceolate,
apically roundish. Florets 4—5, pink or yellow.

Note. The species is similar to A. ferganensis Krasch., distinguished
by the narrower, always simple, lobes of the lower cauline leaves.

Economic Importance. According to the data of M.I. Goryaev,
I.M. Shabanov, and L.A. Ignatova (1953), the essential oil contains:
36.9% levorotatory camphor and 47% cineole. Moreover, the oil also
contains 6.9% free terpenic alcohols and 8.17% bound alcohols as

609

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esters. The alcohol is mostly borneol, which is free form and also as
an ester with a predominance of bornyl-isovalerate. Among acids only
isovalaric acid was identified. Phenols were present in the oil in small
quantity, and their nature has not been studied yet. The oil also contains
sesquiterpenic compounds, partly carbonyl, in which, based on the
recovery of 2.4-dinitro-phenylhydrazones, we may presume the presence
of a-ciperone and tetrahydro-a@-ciperone. The presence of o-thujone
was confirmed from the constants of the fractions of the above-
mentioned compounds.

According to the data of M.I. Goryaev and R.N. Sazonova (1959),
the yield of essential oil is 1.5%; the oil contains aldehydes, ketones,
camphor, azulene-forming sesquiterpenes, and sometimes also phenols.
Alkaloids are also present.

150. A. serotina Bge. in Mém. Acad. Sc. Pétersb. VII (1854) 341;
Boiss. Fl. or. III, 367; Fedtsch. Perech. Rast. Turk. IV, 195.—Poljak.
in Tr. Inst. Khim. Akad. Nauk Kaz. SSR, IV, 21, 27, 94, 119.—A.
maracandica Bge. in Mém. Acad. Sc. Pétersb. VII (1854) 345.—A.
fragrans var. Y. maracandica Boiss. Fl. or. HI (1875) 366; Fedtsch.
op. cit. 195.—A. fragrans Bornm. in Sintenis, exs. No. 1999.

Perennial. Plant 35-65 cm high, grayish in early vegetative phase
from dense arachnoid hairs, later weakly hairy or almost glabrous. Tap
root woody, with short, perennial, woody shoots and annual, rather
short, non-flowering shoots with rosette of leaves of early summer
growth. Flowering shoots few, erect, hairy, later subglabrous, smooth,
stramineous, branched above, with obliquely upright, arcuate or straight,
most often up to 8-12 cm long branches. Lower leaves long petiolate,
withering early, mostly 3-5 cm long, with pinnatisect auricles; their
laminas oblong to broadly oval, twice pinnately cut, terminal lobes
linear, 2-4 mm long, short-acuminate; middle cauline leaves short-
petiolate or sessile, once or twice pinnate; upper leaves undivided,
linear. Capitula sessile or pedunculate, ovate, up to 3 mm long, loosely
spicate on branches in rather wide pyramidal panicle. Involucral bracts
with scarious border; outer bracts small, oval, sub-acuminate, densely
pubescent; inner larger, oblong oval, subglabrous. Florets 5—7; corolla
yellow or red; filaments of stamen as long or somewhat shorter than
anthers; style at maturity of ovary slightly exserted from corolla tube;
stigma lobes long, recurved, with tuft of erect cilia at apex. Flowering
August.

Clayey steppe slopes of foothills, loessial ridges, as also on terraces
of river valleys and dry ravines. Weed of non-irrigated, abandoned,
arable lands and pastures. — Soviet Central Asia: Kyzyl-Kum, Aralo-
Caspian Region (southeastern part), Lake Balkhash Region (southern
part), Syr-Darya. Endemic. Described from Uzbekistan. Type in Paris.

610

580

Note. Var. maracandica Bge., distinguished by longer, usually up
to 15-18 cm-long, often almost horizontal branches, and 3-4 mm-long
(and not 3 mm) capitula, should be recognized.

According to the studies of E.I. Satdarova, the essential oil of A.
serotina Bge. contains the following components: 1) carbonyl
compounds—a) a-thujone—53.9% by oil weight, b) carvone—5.73%
by oil weight; c) camphor—2% by oil weight; d) unidentified carbonyl
compounds—2.4%; 2) alcohols (thujyl alcohol—23.2%); 3) terpenic
oxides (1.8=cineole=6.6% by oil weight); sesquiterpenic compounds
similar to aremadendrone; its dehydrogenation produced azulene; 5)
phenols (allylpyrocatechin was found in traces); 6) acids (isobutyric
acid). According to the data of M.I. Goryaev, A.T. Petushkova, and
N.A. Sviridova samples of A. serotina Bge. contain approximately
0.1-0.5% alkaloids, as was reported for the first time by P.S.
Massagetov. Carotene content in the plant varies significantly: in
drought years from 4.5 to 62.2 mg/kg and in wet years from 10.2 to
62.5 mg/kg; its minimum content being in the early spring, fall, and in
the driest part of summer, while the maximum level shows two peaks:
in April-May and August-September (Karmanovskaya, Berlyaeva, and
Ryabina, 1951). The nutritive value of this wormwood, according to
the above authors, is very high: 100 kg of absolute dry fodder
contains 8.0-14.1 kg digestible protein and 57.6-106.6 fodder units.
Despite high indices, the animals either do not eat this plant at all
or very reluctantly. According to P.P. Poljakov’s observations,
this species is a rather persistent pasture weed, widely distributed in
southern Kazakhstan and to some extent in the northern parts of
Uzbekistan.

According to the data of M.I. Goryaev, G.I. Kryglykhina, and E.I.
Satdarova (1959), var. maracandica Bge. contains alkaloids.

151. A. valida Krasch. ex Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVI (1954) 398; Poljak. in Tr. Inst. Khim. Akad.
Nauk Kaz. SSR, IV, 29.

Perennial. Root woody, thick, with rather numerous, ascending,
perennial, woody, short, nonflowering shoots forming small tussocks.
Flowering shoots numerous, less often few, strong, virgate, erect,
sometimes basally ascending, 20-35 cm high, whitish, arachnoid-hairy
when young, but partly glabrescent in fall, branched above, with
somewhat divergent branches. Lower leaves withering early; petiolate,
usually 2-4 mm long, lamina oval, twice pinnately cut, terminal lobes
1.5—3.0 mm long, somewhat succulent, linear; middle cauline leaves
less compound, sessile; bracteal leaves small, linear or lanceolate.
Capitula sessile or pedunculate, up to 3 mm long, spicately crowded

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on branches in relatively narrow, dense, paniculate inflorescence.
Involucre imbricate, whitish, densely pubescent; inner involucral bracts
larger, outer bracts oval-oblong or oval, with broad, scarious border.
Florets 3-5 corolla yellow. Flowering August.

Depressions, dry flat[beds of] ravines, clayey desert-like, salt
marshes; quite abundant.—Soviet Central Asia: Syr-Darya. Endemic.
Described from southern Kazakhstan, in vicinity of the village of
Kobulsai. Type in Leningrad.

Economic Importance. According to the date of M.I. Goryaev
and R.N. Sazonova (1959) the plant contains 0.5% essential oil; a
little cineole and a large quantity of phenols were found in the oil.
Phenols crystalize from the oil during steam distillation. Based on the
analytical results, the phenol is tentatively related to the
fluoracetophenon-dimentyl ester with the chemical formula C,,H,,O,.
It is interesting to note that the quantity of phenols in the oil increases
as the plant becomes older.

152. A. gypsacea Krasch., M. Pop. and Lincz. ex Poljak. in Bot.
Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954) 409.

Perennial, semishrub 60-80 cm high, with perennial woody, long,
erect, up to 6-8 mm thick nonflowering shoots covered with brownish-
gray cracking bark. Flowering shoots few, straight, erect, 45-65 cm
high, grayish, arachnoid-hairy, later almost glabrous, particularly in
lower part, stramineous, branched, with short (2-6 cm long) or long,
up to 12 cm, weakly divergent branches. Lower leaves long-petiolate,
5-8 cm long, densely hairy, later subglabrous, oblong, twice pinnately
cut, terminal lobes linear, 7-12 mm long and 1 mm wide, with prominent
veins, scarcely acuminate; middle cauline leaves sessile, with pinnate
auricles; upper leaves undivided, linear, often with 2 short lobes at
base. Capitula sessile or pedunculate, ovoid, up to 4 mm long, upright,
remote, 2-3, only at apices of branches, in narrow panicle. Involucral
bracts light brownish, with wide scarious border; outer bracts small,
oval, scarcely acuminate, weakly pubescent, inner larger, oblong,
glabrous. Florets 4—5, corolla up to 3 mm long; anthers at maturity
scarcely exserted from corolla tube or as long as tube; stigma lobes
one-fourth as long as style, with tuft of erect cilia at apex.

Southern clayey slopes, sometimes on white clay out-crops.—Soviet
Central Asia: Mountainous Turkmenia. General distribution: Northern
Iran. Described from the Chul. Type in Leningrad.

According to the data of Vyshensky, 0.5-0.7% essential oil
containing camphor was obtained from the plant up to flowering time.
I.A. Linczevsky (1935) relates this species to fodder plants in Kopetdag.

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582

153. A. namanganica Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVIII (1957) 275; Poljak. in Tr. Inst. Khim. Akad. Nauk
Kaz. SSR, IV, 119.—A. uzbekistanica Poljak. op. cit. 277.

Perennial, grayish arachnoid-hairy, later subglabrous. Root vertical,
woody with short, perennial, woody, nonflowering ascending shoots
forming small tussock. Flowering shoots very numerous, 35-60 cm
high, straight or weakly curved, virgate, strongly branched, long, more
or less divergent. Leaves of vegetative shoots and lower cauline leaves
petiolate, 1.5-2.5 cm long and 0.7—1.0 cm wide, twice pinnately cut,
terminal lobes 1.5—4.0 mm long, linear, subacuminate or round; middle
cauline leaves sessile, once or twice pinnately cut, with pinnate auricles;
upper cauline leaves smaller, simple pinnate, uppermost leaves
undivided, linear, obtuse, sometimes with a pair of lateral lobes at
base. Inflorescence pyramidal panicle; capitula sessile, ovoid, 3-4 mm
long, upright, approximate or somewhat remote. Involucral bracts
convex, with wide scarious border; outer bracts oval, herbaceous,
arachnoid-hairy, inner larger, elliptical or oblanceolate, more or less
glabrous. Florets 7-8; corolla yellow. Flowering August.

Rubbly-clayey desert, sometimes on rubbly-gravel deposits.—Soviet
Central Asia: Syr-Darya (Fergana valley). Endemic. Described from
Namangan District. Type in Leningrad.

Note. Var. usbekistanica Poljak. should be recognized; it is dis-
tinguished by smaller capitula (2-3 mm and not 3-4 mm long) with
fewer florets (florets usually 3-7 and not 7 or 8); vicinity of Namangan
Town.

Economic Importance. According to the data of M.I. Goryaev,
G.K. Kruglykhina, and E.I. Satdarova (1959) it contains alkaloids.

Series 8. Turanica Poljak.—Involucral bracts usually many-
whorled, imbricate, convex; flowering shoots whitish-hairy, later
subglabrous, brown or blackish-brown, smooth; cauline leaves withering
relatively early.

154. A. turnica Krasch. in Otch. 0 Rab. Pochv. Bot. Otr. Kazakhsk.
Eksp. Akad. Nauk SSSR, IV, 2 (1930) 270.—A. pallida Poljak. and
Krasch. in Byull. Gl. Bot. Sada Akad. Nauk SSSR, 6 (1950) 66.—A.
eremophila Krasch. ex Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVI (1954) 401.—A. diffusa Krasch. ex Poljak. ibid.,
400; Poljak. in Tr. Inst. Khim. Akad. Nauk Kaz. SSR, IV, 119.

Perennial. Tap root thick, woody, in old specimens vertical split
into several parts, with very numerous, prostrate or ascending, strongly
condensed, perennial, woody, nonflowering shoots covered with grayish
cracking bark. Flowering shoots numerous, slender, 20-40 cm high,

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583

whitish arachnoid-hairy, later completely glabrous, intensely violet-
brown or almost black, lustrous, branched above with almost horizontal
branches (cf. the note). Leaves withering early, green; lower leaves
petiolate, up to 1.5-2.0 cm long, their laminas oval, twice or thrice
pinnately cut, terminal lobes linear, 3-5 mm long; middle cauline leaves
sessile, less compound, with pinnate auricles; uppermost leaves
undivided, linear, sometimes with 2 lateral lobes at base. Capitula
sessile, remote, ovate or oblong, small, 2 mm long, in broad lax panicle.
Involucral bracts 5—7-rowed, imbricate, grayish-tomentose, later
subglabrous; outer bracts small, fleshy, oval, convex, humped, inner
larger, oblong or oblong-linear, with wide scarious, lustrous border.
Florets 3—5, corolla yellow; anthers linear, apical appendages subulate,
basal appendages much short, round; stigma lobes short, linear-oblong,
truncate, short-ciliate, after anthesis weakly divergent. Flowering
September.

Clayey-rubbly desert soils, along edges of sands, sandy gravelly
alluvium, loams, as well as on outcrops of sandstones, limestones and
conglomerates.—Soviet Central Asia: Kara-Kum, Kyzyl-Kum, Aralo-
Caspian Region (southeastern part), Lake Balkhash Region (Betpak-
Dala, Muyun-Kums), Syr-Darya, Amu-Darya, Pamiro-Alai (southeastern
part). Endemic. Described from central Kazakhstan. Type in Leningrad.

Note. The following varieties should be recognized:

1) Var. pallida Poljak. ex Krasch.—distinguished by slender stems
that are initially densely hairy, later almost glabrous, brownish,
branched, with upwardly directed branches; found in Tien Shan (foothill
plain of Syr-Darya part of Karatau).

2) Var. diffusa Krasch. ex Poljak.—With strong branched stems
that are up to 3 mm thick at base, with leaves grayish-tomentose almost
up to the end of vegetative growth or partly glabrescent, brownish or
violet-brown in fall; found in Kyzyl-Kum and Kara-Kum.

3) Var. eremophila Poljak.—distinguished by few, comparatively
low (15-25 cm high), weakly branched stems, with arcuate or almost
horizontal branches; fewer capitula that are remote on branches; oc-
casional in the foothill plain near Turkestan Range.

Economic Importance. According to the data of Z.M. Manulkin,
the yield of essential oil is 0.69% (0.34% by weight of the whole
plant). The oil has a characteristic camphor smell; the test for aldehydes
and ketones is negative.

According to the data of Nechaeva (1949), var. diffusa has a
strongly branched (70-80 cm thick[sic.]*) tap root in the upper 30-40
cm of the soil; besides, the major part of the roots and rootlets is

*Presumably ‘long’ intended—General Editor.

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584

concentrated in the upper 20 cm of the soil. Because of the shallow
root system the plant suffers greatly from summer drought.

The chemical composition of var. diffusa was studied by Pelt,
Amelin, Morozova, and Morozov. They found in this variety a rather
large quantity of ash, varying from 6.3 to 10.9%, depending on the
stage of vegetative growth; fat—2.9-7.3%; cellulose—33.0—46.8%;
nitrogen-free extractible substances—37.5—43.1%. Traces of alkaloids
were found in the young shoots and capitula (Lazurevsky, 1937).
According to the observations of Borovsky, Zinovev and others, it is
a good fall-winter fattening fodder for sheep, camels, and, to some
extent, horses; similarly it is relished in early spring. According to the
data of I.V. Larin, the highest nutritive value of var. diffusa is found
in spring (May) and is comparatively lower in late fall. According to
the studies of M.I. Goryaev and R.N. Sazonova (1959), the plant
contains 3.826% santonin; the yield of essential oil is 1.1%; the oil
contains aldehydes, phenols, ketones, cineole, and azulene-forming
sesquiterepenes.

According to the data of M.I. Goryaev, the yield of essential oil
in var. pallida Poljak. is 1.82% on the basis of absolute dry weight;
the test for phenols produces a weak, brownish-green color; the oil has
a camphorous smell.

155. A. kemrudica Krasch. in Otch. 0 Rab. Pochv. Bot. Otr.
Kazakhsk. Eksp. Akad. Nauk SSSR, IV, 2 (1930) 270.—A. herba-alba
Asso ssp. kemrudica Krasch. in herb.—Ic.: Krascheninn. op. cit., Plate
|e

Perennial. Whole plant white when young, later grayish-greenish
from arachnoid hairs. Root thick, vertical, woody, in older specimens
often vertically split into several parts, with short, perennial, ascending
or erect, woody, nonflowering shoots, forming rather compact, large
tussock; short annual shoots with rosette of leaves in early summer;
flowering shoots numerous, very slender, straight or weakly flexuous,
12-25 cm high, branched above, with upwardly directed, sometimes
drooping or horizontal, usually 2-5 cm long branches. Leaves of
nonflowering shoots and lower cauline leaves withering in summer,
petiolate, 8-15 mm long, lamina oblong-lanceolate, once or twice
pinnately cut, lobes divided into 2-3 linear secondary lobes, or simply
linear; middle cauline leaves sessile, much shorter, pinnately cut;
uppermost leaves undivided, linear. Capitula pedunculate, more or less
divergent, sometimes drooping, ovoid or ovoid-conical, that is, shortly
narrowed toward base, 3.0-4.5 mm long and 1.5-2.0 mm in dia,
approximate or remote, in comparatively narrow paniculate
inflorescence. Involucral bracts punctate-glandular, outer small, oval,

615

585

dorsally humped-convex, hairy; inner much larger, oblong, brownish,
subglabrous, with scarious margin. Florets 2-3. Achenes somewhat
flat, ovate, brown, up to 1.5 mm long and 0.7 mm wide.

Soviet Central Asia: Aralo-Caspian Region (Mangyshlak Peninsula,
western Ust-Urt), Kara-Kum (northern part). Endemic. Described from
eastern coast of Caspian Sea. Type in Leningrad.

Note. Var. karabugasica Krasch. (Otch. o Rab. Pochv. Bot. Otr.
Kazakhsk. Eksp. Akad. Nauk SSSR, IV, 2 (1930) 270) differs from the
typical form by having short stems (S—8 cm high), dense white-arachnoid
pubescence, small simple pinnate leaves 2-4 mm long, and small sessile
capitula. It is found on the eastern coast of the Caspian Sea, near
Karabugaz Gulf.

156. A. scotina Nevski in Tr. Bot. Inst. Akad. Nauk SSSR, Ser.
1, No. 4 (1937) 284.—A. albicaulis Nevski, ibid., No. 200.

Perennial. Plant 30—35(40) cm high, grayish below in early
vegetative phase, densely hairy, later partly glabrescent. Root vertical,
woody, very thick, with short, perennial, ascending, less often prostrate,
woody, nonflowering shoots forming tussock. Fertile shoots numerous,
20-30 cm high, straight or weakly flexuous, basally often arcuate,
branched above, with long, less often short, upwardly directed branches.
Leaves grayish-green, lower cauline leaves and those of nonflowering
shoots petiolate, their laminas oval, 1-2 cm long, twice pinnately cut;
lobes pinnate, with 1-2 pairs of terminal linear-spatulate or linear
lobes 3-6 mm long; middle cauline leaves sessile, less divided, with
linear, sessile auricles at base; upper leaves undivided, linear-oblong,
subacuminate. Capitula usually sessile, ovate, 3-4 mm long, remote or
approximate in more or less oblong panicle. Involucre many-rowed,
imbricate; involucral bracts punctate-glandular; outer bracts small, oval,
densely hairy; inner larger, oval-elliptical or more or less lanceolate,
hairy in upper part, elsewhere glabrous, light-green, with wide, scarious
margin. Florets 5—6; corolla yellow; stigma lobes short, more or less
divergent, apically with tuft of upright cilia. Flowering September.

Rubbly and stony slopes of lower mountains.—Soviet Central Asia:
Pamiro-Alai Region. Endemic. Described from Kugitang Range. Type
in Leningrad.

157. A. kochiiformis Krasch. and Lincz. ex Poljak. in Bot. Mat.
Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954) 411-412.

Perennial. Plant 35—60(70) cm high, whitish from dense arachnoid-
hairs, partly glabrescent by end of vegetative growth. Tap root, woody,
with brown, short, ascending, perennial, woolly, nonflowering shoots
forming tussock. Flowering shoots 12—20 and more, straight, robust,

586

whitish-tomentose, later sub-glabrous, stramineous-brown, smooth,
branched above, with upwardly directed branches, up to 12 cm long.
Leaves of vegetative shoots and lower cauline leaves long-petiolate,
4-7 cm long, their laminas oblong, twice or thrice pinnately cut, terminal
lobes narrowly linear, 7-12(20) mm long and 0.3 mm wide, short-
acuminate; middle cauline leaves sub-sessile, once or twice pinnately
cut, with pinnate or undivided linear auricles at base; uppermost leaves
undivided, linear. Capitula sessile, upright, oblong-ovoid, narrowed
toward base, cuneate, 3—4 mm long, in oblong paniculate inflorescence.
Involucre many-rowed, imbricate; outer bracts small, oval, convex,
inner larger, oblong, with wide, scarious margin. Florets 5; corolla
yellow; anthers linear, apical appendages of anthers subulate, basal
appendages short, subacuminate, style shorter than stamens; stigma
lobes linear, exserted, short-ciliate. Achenes plano-convex, ovoid, up
to 1 mm long and 0.5 mm wide.

Clayey, sometimes on rubbly slopes of mountains, river terraces
above floodplain and gypsiferous outcrops.—Soviet Central Asia:
Pamiro-Alai Region. Endemic. Described from southern Tadzhikistan
(Karatau Range). Type in Leningrad.

158. A. sogdiana Bge. in Mém. Acad. Sc. Pétersb. VII (1854)
343; Boiss. Fl. or. III, 367; Fedtsch. Perech. Rast. Turk. III, 195.

Perennial. Plant grayish from dense arachnoid hairs, partly glabrous
at anthesis. Root thick, woody, vertical, with woody, strongly shortened,
ascending, nonflowering shoots covered with light brown peeling bark,
annual shoots bearing rosettes of early summer leaves; flowering shoots
30-35 cm high, numerous, slender, erect, virgate, branched slightly
above base, with somewhat divergent branches, mostly 1 to 3 cm long.
Leaves of vegetative shoots and lower cauline leaves short-petiolate,
up to 1.0-1.5 cm long, their laminas oblong, twice pinnately cut,
terminal lobes oblong-linear, subobtuse; higher leaves usually sessile,
pinnate, with 2 pairs of lateral lobes; uppermost leaves undivided,
oblong-linear, with short oblong lobes at base. Capitula pedunculate,
oblong-ovoid or narrowly campanulate, up to 3 mm long, spicately
aggregated, less often solitary and remote in narrowly branched
inflorescence. Involucral bracts with scarious border, outer bracts small,
oval, inner long, linear-oblong. Florets 3; corolla pink; anthers linear,
apical appendages of anthers narrowly linear, almost subulate, basal
appendages shorter, subacuminate; style shorter than stamens; stigma
lobes narrowly linear, truncate, short-ciliate, after anthesis divergent.

Gravelly-clayey deposits in desert zone.—Soviet Central Asia: Syr-
Darya. Endemic. Described from Zerav-shan. Type in Leningrad.

617

587

Note. The report by B.A. Fedtschenko (Perech. Rast. Turk., IV
(1911) 196) on the occurrence of Artemisia oliveriana Gay ex Bess.
in the Tien Shan and the Samarkand Region is not confirmed by
herbarium material. This species, described from northern Iran, has
cylindrical capitula in spicate-paniculate inflorescences with few, purple
florets and shiny involucral bracts. On this basis one can say that it
probably is close to A. sogdiana Bge.

159. A. stenocephala Krasch. ex Poljak. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XVI (1954) 404.

Perennial. Root woody, vertical. Flowering shoots few or solitary,
at base woody, erect, 40-80 cm high, arachnoid-hairy, later subglabrous,
light brown, branched slightly above base, with numerous, obliquely
upright, rather long, slender, virgate, straight, flowering branches. Lower
leaves withering early, petiolate, about 1 cm long, twice pinnately cut,
their terminal lobes linear, 2-3 mm long, subobtuse; upper leaves
sessile, undivided, linear, mostly 1.5-3.0 mm long, obtuse, round.
Capitula in dense, more or less broad, paniculate inflorescence, sessile,
narrowly oblong, 3—4 mm long and up to 1.5 mm in dia, somewhat
remote or almost spicately aggregated. Involucral bracts many-rowed,
with scarious border; outer bracts oval, convex, grayish-green,
pubescent, inner long, oblong-lanceolate, subglabrous. Florets 5-6
corolla purple-pink; anthers linear, apical appendages of anthers
narrowly angular, round, basal appendages acute; stigma lobes linear,
exserted. Flowering September.

Gravel streambeds.—Soviet Central Asia: mountainous Turkmenia
(Kopetdag). General distribution: Afghanistan. Described from
Bukinich’s collections from Afghanistan (between Ghazni and
Kelat[Kalat, Khelat]). Type in Leningrad.

160. A. kopetdaghensis Krasch. ex Poljak. in Bot. Mat. Gerb.
Bot. Inst. Akad. Nauk SSSR, XVI (1954) 406.

Perennial. Plant 35-45 cm high, whitish, later grayish-greenish
from arachnoid hairs. Root vertical woody, thick, with short perennial,
upright, woody, nonflowering shoots covered with brownish-gray
peeling bark.

Flowering shoots numerous, 25-40 cm high, 1.0-1.5 mm thick,
usually erect, virgate, branched in upper third, with upwardly directed
branches, mostly up to 4-7 cm long. Leaves on most part of stem
withering, early, with pinnately cut auricles; lower leaves petiolate,
1.5—2.0 cm long, their laminas oval, twice pinnately cut, terminal lobes
narrowly linear, 4-6 mm long, subacuminate, middle cauline leaves
subsessile, less compound, upper sessile, undivided, linear, with 2

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588

lateral, narrow lobes at base. Capitula sessile, upright, 3.0-3.5 mm
long, somewhat remote or loosely spicate on branches, in oblong,
paniculate inflorescence. Involucre many-rowed, imbricate; involucral
bracts white-pubescent, punctate-glandular, oval, with scarious border,
outer bracts shorter than inner. Florets 4—5.

Clayey or rubbly clayey slopes of hills.—Soviet Central Asia:
mountainous Turkmenia. Described from Kara-kala District. Type in
Leningrad.

According to the studies of V.I. Nilov and N.P. Kiryalov, the yield
of essential oil varies according to the stage of vegetative growth from
0.2 to 1.2%. After analysis of the fractions the presence of a-pinene
—2.2%, cineole—35—40%, and ketones—47-79% was determined;
camphor and mainly a-thujone, as well as a small quantity of alde-
hydes and a similar unstudied phenol, were found among them.

161. A. balchanorum Krasch. in Spisok Rast. Gerb. Fl. SSSR, X
(1936) 104, 105; Bobrov in Tr. Bot. Sada Akad. Nauk SSSR, XLIV,
27, 75.—Exs.: GRF No. 3197.

Perennial semishrub, 40-50 cm high, pleasantly fragrant. Root
woody, with short, ascending, perennial, woody, nonflowering shoots
covered with gray bark. Flowering shoots hairy, later subglabrous,
light brown, branched above, with weakly divergent branches. Leaves
grayish, later green, densely punctate-glandular, lower cauline leaves
withering early, petiolate, 1-3 cm long, lamina oblong, twice or thrice
pinnately cut, terminal lobes somewhat thick, remote, linear, 3-5 mm
long; middle cauline leaves subsessile, less divided; uppermost leaves
linear. Capitula sessile, upright, ovate, 3-4 mm long, remote or 2-3,
in long, compact panicle. Involucre many-rowed, imbricate; involucral
bracts hairy and densely punctate-glandular, outer bracts small, round,
carinate, inner large, oblong-oval, green, with broadly scarious border.

_Florets 4-5; corolla yellow, punctate-glandular.

Clayey and stony slopes, foothill plains, limestone outcrops.—
Soviet Central Asia: mountainous Turkmenia (Greater Balkan
Mountains). Described from Greater Balkans. Type in Leningrad.

According to the data of R.Ya. Ratanova, the essential oil of wild
A. balchanorum Krasch. contains about 16% citral, about 35% geraniol,
35-40% levorotatory, linalool [coriandrol], about 0.5% cresols, about
1% valeric and butyric acids. Cultivated specimens of this species
contained: 5% citral, 40-50% linalool, 25-30% geraniol, about 0.5%
cresols, and acids (probably butyric and valeric) in insignificant
quantity.

The plant is cultivated for its essential oil.

589

162. A. turcomanica Gndg. in Bull. Soc. Bot. de France, XVIII
(1918) 38.—Exs.: Sintenis, It. transcasp. pers. 1900-1901, No. 1035.

Perennial semishrub, 30-50(60) cm high. Tap root woody, with
short perennial, woody, erect or ascending, nonflowering shoots covered
with brownish-gray bark. Flowering shoots very numerous, 25—40(45)
cm high, erect or at base arcuate, strong, virgate, in early vegetative
growth phase grayish-tomentose, later partly glabrous, light brown,
branched above, with more or less short, somewhat divergent branches.
Leaves whitish-tomentose, later grayish-green, punctate-glandular; lower
cauline leaves and leaves of nonflowering shoots petiolate, usually
1.2-1.5 cm long, lamina oval or round, twice pinnately cut, terminal
lobes linear, lobate, 1-2-3 mm long, somewhat thick, subacuminate,
middle cauline leaves subsessile, less divided, with pinnate or simple,
linear auricles at base; uppermost leaves linear, sometimes with 2 lateral
lobes at base. Capitula sessile, oblong, 3-5 mm long, loosely spicate
or more or less remote in narrow pyramidal panicle. Involucre many-
rowed, imbricate; outer involucral bracts small, oval, densely hairy,
inner considerably larger, lanceolate-oblong, smooth, with wide scarious
border. Florets usually 5; anthers with very long filaments, linear, apical
appendages of anthers narrow, subulate, basal appendages much shorter,
obtuse; style shorter than stamens; stigma lobes linear, sulcate, exserted,
truncate, with ciliate margin.

Stony slopes of lower mountains—Soviet Central Asia:
mountainous Turkmenia. Endemic. Type published.

163. A. badhysi Krasch. and Lincz. ex Poljak. in Bot. Mat. Gerb.
Bot. Inst. Akad. Nauk SSSR, XVI (1954) 405.—A. arenicola Poljak.
ibid. 408.

Perennial semighrub, 30-45(60) cm high. Tap root woody with
short, erect or ascending, perennial, woody, nonflowering shoots
covered with brownish bark. Flowering shoots numerous, 20—35(50)
cm high, whitish-tomentose, later partly glabrous, light brown, strong,
virgate, branched above, with long, erect or divergent branches. Leaves
grayish-green, densely arachnoid-hairy; lower leaves withering early,
long-petiolate, 1-2-3 cm long, lamina round or broadly round, twice
pinnately cut, terminal lobes linear, 2-5 mm long, subacuminate or
with round apex, middle cauline leaves short-petiolate, less compound
with pinnate auricles; upper leaves sessile, pinnate or broadly linear,
usually with 2 lateral lobes at base. Capitula sessile or pedunculate, 2—
3 mm long, ovate, remote or 2—5 on branches, or crowded in narrow
spicate-paniculate inflorescence. Involucral bracts oval, densely hairy
and punctate-glandular, with scarious border, outer shorter than inner
ones.

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590

Stony mountain. slopes.—Soviet Central Asia: mountainous
Turkmenia, Kara-Kum (southeast). Endemic. Described from Gyaz-
Gyadyk (Badkhyz) Range. Type in Leningrad.

Note 1. Var. arenicola Poljak should be recognized. It is distin-
guished by more or less long, usually almost horizontal branches and
ovoid-oblong, somewhat larger capitula (3.5-4.0 mm long). Kara-
Kum.

Note 2. According to the observations of E.A. Mokeeva and E.lI.
Rachkovskaya, the main mass of the roots of A. badhysi is concen-
trated at a depth of 24 cm (84%); ephemeral rootlets develop on the
upper part of the root using the spring moisture and dying out toward
the dry summer period. The superficial root system enables the plant
to use only rainfalls, thereby making it mostly dependent on precipi-
tation. The vegetative growth of A. badhysi begins in March; rapid
shoot growth commences in April, and the capitula develop in May.
However, in the opinion of the authors, this development is limited to
the formation of the involucre and only in the summer do rudiments of
florets develop slowly, while in the fall the plant flowers and bears
fruits.

In M.I. Goryaev’s work: (Materially po Khimii Efirnogo Masla
[Data on the Chemistry of Essential Oils], Alma Ata, 1952), chemical
data pertaining to [the oil of] A. balchanorum, are erroneously attributed
to A. badhysi.

Economic Importance. According to I.V. Larin, the edibility of
this wormwood for sheep varies with the season of the year as well-as
from year to year, depending on the quality of the pastures. When
there is good growth of the ephemeral vegetation in the spring and
summer, it is not eaten much, but in bad years it is eaten readily.

164. A. prasina Krasch. ex Poljak. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVII (1955) 413.

Perennial semishrub. Root vertical, woody, thick, with short,
perennial, woody, nonflowering shoots. Flowering shoots rather
numerous, sometimes fewer, erect, 20-30 cm high, whitish-tomentose,
later subglabrous, dark brown, branched above, with short or long,
upwardly directed branches. Leaves grayish, densely hairy; lower
cauline leaves and leaves of vegetative shoots petiolate, 1.5—2.0(2.5)
cm long; lamina oval or round, twice pinnately cut, terminal lobes 2—
4 mm long, linear, subobtuse, middle leaves sessile or short-petiolate,
pinnately cut, with palmate or pinnate auricles at base; upper leaves
small, linear, sometimes with 2 lateral lobes at base. Panicle narrowly
pyramidal; capitula sessile, ovoid, 3~4 mm long, upright, approximate
or somewhat remote. Involucral bracts oval, convex, bright, dark bluish-

62

_

591

green, pubescent and punctate-glandular, inner bracts somewhat longer
than outer with scarious border. Florets 5; anthers with shorter filaments,
linear, apical appendages of anthers acute, almost subulate, basal
appendages much shorter, sub-obtuse; stigma lobes short, linear, weakly
sulcate, thickened along edges, apically truncate, with erect cilia.

Clayey slopes of foothills —Soviet Central Asia: Western Pamiro-
Alai Region. Endemic.

Note. A doubtful species, very similar to A. badhysi Krasch.
Additional material is necessary to resolve finally the question of its
separate status.

165. A. spicigera C. Koch in Linnaea, XXIV (1851) 345; Grossh.
Fl. Kakv. IV, 142.—A. araxina Takht. in Tr. Armyansk. Fil. Akad.
Nauk SSSR, Ser. Biol. II (1937) 209.—A. issaevi Rzazade in Izv.
Akad. Nauk AzSSR, 3 (1955) 23.

Perennial. Plant 25—35 cm high, grayish in early vegetative growth
phase, densely pubescent, later sub-glabrous. Tap root, thick, woody,
short, perennial, woody, ascending, nonflowering shoots together with
young, leafy shoots forming low, rather compact tussock. Flowering
shoots numerous, slender, up to | mm thick, stiff, mostly erect,
subglabrous by end of vegetative growth, brown, branched above, with
more or less short, weakly divergent branches. Lower cauline leaves
petiolate, withering, early lamina oval, twice pinnately cut, terminal
lobes linear, 2-4 mm long, subacuminate, middle cauline leaves short-
petiolate or sessile, once or twice pinnate; uppermost leaves sessile,
linear. Capitula sessile or pedunculate, upright, almost linear-cylindrical,
3-4 mm long, in narrow, compact, spicate-paniculate inflorescence.
Involucral bracts tuberculate-glandular and densely whitish-pubescent,
with scarious border, outer bracts small, ovate, inner larger, linear-
oblong. Florets 4—5.

Dry stony slopes.—Caucasus: Southern Transcaucasia. General
distribution: Eastern Anatolia. Described from the former Kars Region.
Type in Berlin.

Note. A. issaevi Rzazade is a rather stunted form of A. spicigera
C. Koch with undeveloped capitula.

166. A. sieberi Bess. in Bull. Soc. Nat. Mosc. IX (1836) 80.—A.
herba alba var. laxiflora Boiss. FI. or. III (1875) 365, p. min. p.—A.
herba alba ssp. saxicola Krasch. in herb.—A. dumosa Poljak. in Bot.
Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954) 425.

Perennial. Low semishrub 20-35 cm high. Root thick, woody,
vertical with numerous, strongly branched, perennial, woody shoots.
Flowering shoots erect, grayish-tomentose, virgate, stiff, 7-15 cm long,

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592

repeatedly branched, with almost horizontal or upwardly directed ter-
minal branches, forming rather compact, stiff, hemispherical crown. —
Leaves of nonflowering shoots petiolate, withering early, 0.9-1.5 cm
long, oval, twice pinnately cut, lobes usually 2 on each side, terminal
lobes linear, 1.5-3.0 mm long, subacute. Leaves of fertile branches
smaller, subsessile, simple pinnate, with pinnate auricles at base; upper-
most leaves very small, linear. Capitula sessile, ovoid, 3-4 mm long,
remote or in clusters of few on terminal branches. Involucre many-
rowed, imbricate; outer involucral bracts small, oval, convex, densely
pubescent, inner much larger, oblong, subglabrous, with wide scarious
border. Florets 3-4; corolla pink or yellow; anthers linear, apical
appendages of anthers narrowly linear, almost subulate, basal
appendages shorter, acuminate; style as long as stamens; stigma lobes
linear, short-ciliate, divergent.

Stony mountain slopes.—Soviet Central Asia: mountainous
Turkmenia (Kopetdag), Kara-Kum (southwest). General distribution:
Northern Iran. Described from Krasnovodsk District. Type in Leningrad.

Note. The report by B.A. Fedtschenko (Perech. Rast. Turk., 1V
(1911) 192) on the distribution of Artemisia herba-alba var. laxiflora
Boiss. and var. densiflora Boiss. in the Chu, Sary-Su, Balkhash, and
Khiya districts and the Trans-caucasian Region, in fact, pertains not to
one taxon, but to a group of four: A. herba-alba Asso, A. oliveriana
Bess., A. sieberi Bess., and A. billardierana Bess. Of these the last
two were united by Boissier as one variety (A. herba-alba var. laxiflora
Boiss.), while A. oliveriana Bess. was designated as var. densiflora
(Fl. or. III, 365). A. billardierana Bess. (from the vicinity of Dam-
ascus) is very close to A. herba-alba Asso (from western Africa and
Spain), and A. oliveriana Bess. (from northern Iran) apparently is similar
to the Mediterranean A. sogdiana Bge. However, this question cannot
be decided finally because of the lack of herbarium specimens. Thus,
from the above-mentioned species, only A. sieberi Bess. is reliably
found in the USSR.

Series 9. Lehmannianea Poljak.—Stems simple or branched above,
with short, less often long branches; capitula remote or in compact
spikes or racemose-spicate or racemose-paniculate, or even spicate
inflorescence; lower cauline leaves twice, thrice or even four times
pinnately cut.

167. A. lehmanniana Bge. in Mém. Acad. Sc. Pétersb. VII (1854)
340; Boiss. Fl. or. III, 368; Fedtsch. Perech. Rast. Turk. [V, 195.—A.
skorniakowi Winkl. in Tr. Peterb. Bot. Sada, XI, 2 (1892) 331; O.
Fedtsch. Fl. Pam. 116; Fedtsch. Perech. Rast. Turk. IV, 199; Poljak.

623

593

in Tr. Inst. Khim. Akad. Nauk Kaz. SSR, IV, 26.—A. korshinskyi Krasch.
ex Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954)
415.—A. lehmanniana ssp. leucotricha Krasch. nom. (in herb.).

Perennial semishrub, 20-45 cm high. Root thick, sometimes up to
2 cm thick, woody, vertical, with short, almost from base strongly
branched, perennial, woody, ascending, nonflowering shoots covered
with brownish bark. Flowering shoots numerous or few, 15—25(50) cm
high, arachnoid-hairy, later subglabrous, strong, sulcate, straight or
arcuate, simple or with short branches. Leaves grayish, densely
pubescent, later subglabrous or with remnants of pubescence (f.
tomentosa Krasch.); lower leaves long-petiolate, 2—3(5) cm long, with
truncate auricles; lamina roundish or prolate, thrice pinnately cut;
terminal lobes linear to linear-subulate; middle cauline leaves short-
petiolate, apical leaves sessile, mostly pinnate. Capitula sessile, ovoid,
2.5-3.0 mm long, in groups of few in compact globose or ovoid spikes
(heads), latter on short, remote branches, usually not exceeding spikes,
forming narrowly spicate-paniculate inflorescence. Involucral bracts
oval, glabrous, with wide, brown, scarious border, apically emarginate-
toothed, dorsally punctate-glandular. Florets 7-8; corolla purple;
stamens scarcely exserted. Receptacle small, smooth.

High-mountain rubbly and clayey-rubbly slopes.—Soviet Central
Asia: Pamiro-Alai, Tien Shan. General distribution: Kashgaria.
Described from Zeravshan Range (Karatau). Type in Leningrad.

Note. The following varieties should be recognized:

1) Var. skorniakovii Winkl.— distinguished by short stem, 12-30
cm high, grayish, arachnoid-hairs persisting up to the end of vegetative
period, and short-petiolate, very closely borne leaves. Eastern Pamir,
on rubbly slopes and moraines. This variety is usually confused with
A. rhodantha Rupr., also found in the Pamir, although the latter is well
distinguished, being an herbaceous perennial.

2) Var. korshinskyi Krasch. ex Poljak.—distinguished by branched
stems with rather long upright branches.

Economic Importance. In the early stages of its vegetative growth,
it is eaten readily by sheep and goats and is relished by yaks; in the
summer it is not eaten by live-stock. According to the studies of
Pekhachek (1948), 100 g of green mass contains 2 to 8 mg of vitamin
C and 3.2—6.2 mg of vitamin A. Experiments on digestibility, conducted
by Pekhachek (1944) at the Pamir Biological Station, have established
that 100 kg of the herb contains 3.0 kg of digestible protein and 22.7
fodder units, corresponding to 7.5 and 56.8 respectively, when converted
to absolute dry weight. Fodder with such a nutritive value, in Larin’s
opinion, is in no way inferior to the best hay from leguminous herbs.
It must be noted that Pekhachek’s data pertain not to var. skorniakovii

594

(C. Winkl.) Poljak., as considered by the author of the report, but
probably to A. rhodantha Rupr., which is found in the vicinity of the
Pamir Biological Station.

According to the data of M.I. Goryaev and R.N. Sazonova (1958),
the plant contains 0.07% essential oil; the oil contains a small quantity
of phenols, azulene-forming sesquiterpenes, and ketones.

168. A. vachanica Krasch. ex Poljak. in Bot. Mat. Gerb. Bot.
Inst. Akad. Nauk SSSR, XVI (1954) 413.

Perennial, glaucescent green plant, 25-30 cm high. Tap root thick,
woody, with numerous perennial, strongly reduced, ascending, woody,
nonflowering shoots forming tussock. Flowering shoots numerous,
slender, more or less erect, sulcate, greenish, weakly pubescent or
subglabrous, simple or with short, appressed branches in upper part.
Leaves of nonflowering shoots and lower cauline leaves usually densely
grayish-lanate, short-petiolate, 1.0-1.5 cm long, twice pinnately cut,
terminal lobes 1.0—1.5 mm long, oblong-linear, obtuse; middle cauline
leaves sessile, weakly pubescent or subglabrous, apical leaves
undivided, lanceolate, keeled, short-acuminate. Capitula in simple,
sparse, spicate or racemose-spicate inflorescence, remote, solitary, less
often 2-3, sessile, ovate, up to 2 mm long. Involucral bracts punctate-
glandular; outer bracts small, oblong, more or less pubescent, inner
larger, oval, glabrous or subglabrous, with wide scarious border. Florets
3-4; corolla purple-pink.

Stony mountain slopes and in high-altitude semi-desert zone.—
Soviet Central Asia: Pamiro-Alai Region (mountainous Badakhshan).
Endemic. Described from vicinity of Lyangar. Type in Leningrad.

169. A. baldshuanica Krasch. and Zapr. in Tr. Bot. Inst. Akad.
Nauk SSSR, Ser. 1, No. 3 (1936) 351.

Perennial semishrub, 80-100 cm high, with long, perennial,
ascending or erect, woody, nonflowering shoots covered with gray
cracking bark. Flowering shoots ascending or erect, branched in upper
half, with long (up to 20-35 cm), strongly divergent branches. Leaves
grayish from white tomentose, caducous hairs; leaves of nonflowering
shoots and cauline leaves subsessile, 3.5—5.0 cm long and 1.5—3.0 cm
wide, oval or oblong, thrice or four times pinnately cut, their terminal
lobes linear, 2-4 mm long, scarcely acuminate. Capitula sessile, ovoid,
mature capitula narrowly goblet-shaped, 3.0-2.5 mm long, 2—3 or more
at apices of branches in spicate, rather broad, paniculate inflorescence.
Involucral bracts convex, with scarious border; outer bracts small,
round-oval, inner larger, oblong. Florets usually 5, with purple-red
corolla; anthers on comparatively short filaments, apical appendages

625

595

of anthers narrow, subulate, basal appendages much shorter, subobtuse;
style almost as long as stamens, its lobes short-linear, sulcate, erect,
with thick margin, apically truncate, with short, upright cilia. Achenes
small, 0.5 mm long, ovoid, somewhat flat, brownish.

Clayey and clayey-rubbly mountain slopes.—Soviet Central Asia:
Pamiro-Alai Region. Endemic. Described from Lyangar District. Type
in Leningrad.

According to the studies of I.P. Tsukervanik and L.S. Grach,
the essential oil of A. baldshuanica Krasch. and Zapr. contains about
60% a- and f-thujone, 16% thujyl alcohol, a small quantity of
isothujylvalerianate and aldehydes (semicarbazone, with melting point
at 199-200°).

Series 10. Santolinae Poljak.—All leaves simple, linear or ob-
long-linear, short pinnately lobed; involucral bracts withering before
maturity of achenes.

170. A. santolina Schrenk in Bull. Phys. Math. Acad. Sc. Pétersb.
III, 7 (1845) 106; Fedtsch. Perech. Rast. Turk. IV, 193.—Exs.: GRF
No. 3198.

Perennial. Root rather thick, woody, vertical, with short, more or
less erect, perennial, woody nonflowering shoots covered with brown
peeling bark. Flowering shoots in large number, straight or flexuous,
25-35(45) cm high, whitish-hairy, branched above, with upwardly
directed branches. Leaves of nonflowering shoots and lower cauline
leaves short-petiolate, strongly variable in size, 1-7 cm long, oblong-
linear, pinnately divided, lateral segments undivided, roundish or oblong
or in turn even shallowly divided into 2-3 roundish lobes; sometimes
leaves linear, entire (f. simplicifolia Poljak.) or broadly toothed; upper
leaves sessile, small, linear. Capitula sessile or pedunculate, ovoid, up
to 5 mm long, with 3-4 florets, remote in lax, oblong, paniculate
inflorescence. Involucral bracts withering before achenes maturity, outer
bracts small, oval, grayish-pubescent, inner larger, oblong, scarious,
glabrous, dorsally with narrow, brownish stripe. Flowering August.

Desert sands.—Soviet Central Asia: Aralo-Caspian Region, Lake
Balkhash Region, Kara-Kum, Kyzyl-Kum. General distribution:
Northern Iran, Kuldzha. Described from Ili River sands. Type in
Leningrad.

171. A. lobulifolia Boiss. Flora Orientalis, III (1875) 364.

Perennial. Whole plant more or less grayish-tomentose; root rather
thick, woody, vertical, with perennial, woody, nonflowering shoots
covered with brownish-gray peeling bark, strongly reduced, erect.

626

596

Flowering shoots rather numerous, almost straight, erect, 25-35 cm
high, branched almost from base, with more or less divergent branches.
Lower and middle cauline leaves linear, shortly pinnately lobed, their
lateral segments small, deltoid-oval or oval, short-decurrent on rachis,
usually 7-9 on each side, simple or with 2—3-parted lobes; upper leaves
small, undivided, scaly, broadly linear. Capitula sessile, oblong, up to
4 mm long, with 5-6 florets, remote or crowded in lax paniculate
inflorescence. Involucral bracts densely hairy, outer bracts small, round
or oval, convex, almost herbaceous, inner larger, linear-oblong, scarious.

Takyrs in depressions and limestones outcrops. — Soviet Central Asia:
Kara-Kum. General distribution: Iran-Afghanistan (western Afghanistan).
Described from western Afghanistan (between Herat and Tabas).

Note. This species is very similar to A. santolina Schrenk but
differs from it by having hairy deltoid-oval or oval, lateral segments
of the leaves that are very short-decurrent on the rachis.

Series 11. Tridentatae Poljak.—All leaves simple, linear, entire
or with three to five apical teeth.

172. A. deserti Krasch. in Sched. ad Herb. Fl. URSS, X (1936)
106.—Exs.: GRF No. 3199a, b.

Perennial semishrub, 50-60 cm high. Root thick, woody, vertical,
perennial, woody, branched, upward-spreading, nonflowering shoots
covered with whitish-gray, peeling bark. Flowering shoots numerous,
more or less grayish from hairs, later subglabrous, erect, relatively
slender, virgate, branched above, with straight, distinctly divergent,
slender branches. Leaves densely grayish-pubescent, undivided, linear,
with narrowly carinate base, 1-2 cm long, 2-4 mm wide, apically with
3-5 obtuse, broadly linear teeth, 1.5 mm long; upper leaves entire,
shorter and narrower. Inflorescence more or less broadly paniculate;
capitula sessile or on short peduncles, upright, remote or approximate,
2-3, ovoid-oblong, 4—5 mm long. Involucre imbricate; outer involucral
bracts small, oval, carinate, grayish-pubescent, inner larger, oblong or
oblong-oval, smooth, with broadly scarious border, greenish on convex
dorsal side. Florets 3-5; corolla purple; anthers linear, apical
appendages of anthers narrowly linear, obtuse, basal appendages shorter,
scarcely acuminate; style at achen maturity as long as stamens or
somewhat longer, its lobes linear, short-ciliate, after flowering divergent.
Achenes up to 2 mm long, ovoid, somewhat flat, brown.

Dry sandy-gravelly beds of mountain streams.—Soviet Central Asia:
mountainous Turkmenia (Kyurendag). Endemic. Described from vicinity
of Kazandzhik. Type and cotype published.

627

597

Plate XXX.

1 — Turaniphytum eranthemum (Bge.) Poljak., habit, lower and middle leaves,

involucral bract, achene; 2 — Artemisia capillaris Thunb., leaf, involucral bract,

achenes, peripheral pistillate floret, rudiment of pistil of disk floret, stamen;

3—A. saposhnikovii Krasch. ex Poljak., leaf, involucral bract, peripheral

pistillate floret, rudiment of pistil of disk floret, achene; 4—A. desertorum

Spreng., leaf, involucral bract, rudiment of pistil of disk floret, upper part of
peripheral pistillate floret.

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598

Section 2. Junceum Poljak.—Lower cauline leaves mostly 3-lobed;
hairs bifid, straight, appressed or long and erect; receptacle small,
hemispherical.

Series 12. Appressopilosae Poljak.—Perennials; hairs straight,
bifid, appressed.

173. A. juncea Kar. and Kir. in Bull. Soc. Nat. Mosc. XV (1842)
383; Ldb. Fl. Ross. II, 2, 576; Fedtsch. Perech. Rast. Turk. IV, 195;
Kryl. Fl. Zap. Sib. XI, 2788; Poljak. in Tr. Inst. Khim. Akad. Nauk
Kaz. SSR, IV, 21, 26, 96.—A. multifida Kar. and Kir. nomen in herb.—
A. macrosciadia Poljak. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVI (1954) 423; Poljak. op. cit. 93.—Ic.: Poljak. op. cit. 93.—
Exs.: Gerb. Fl. SSSR, No. 3190.

Perennial. Whole plant grayish from dense, short, compactly
appressed hairs. Root thick, woody, vertical, with strongly reduced,
perennial, woody, nonflowering shoots covered with yellowish-brownish
fibrously split bark. Flowering shoots straight, sulcate, numerous, 20—
40 cm high, branched above, with short branches. Leaves except
uppermost, petiolate, petioles as long as or shorter than lamina, lamina
cut up to base into 3 lobes; middle and upper leaves undivided, linear,
apically slightly wider, with obtuse cartilaginous apex; lower leaves
divided into 2-3 linear lobes; uppermost leaves undivided, linear.
Capitula sessile, lax, terminal on stems, in loosely arranged dense
spikes on short branches, oblong-ovoid, 4.0—4.5 mm long and about 2
mm in dia or 5—6 mm long and up to 2.5 mm in dia. Outer involucral
bracts broadly oval, grayish from compactly appressed hairs, 1/5 to
1/4 as long as inner bracts, later oblong-oval or oblong, almost smooth,
with wide, light brownish, scarious border. Florets 4—7; corolla yellow;
anthers linear, apical appendage of anthers subulate, basal appendage
scarcely visible, round; stigma lobes linear, short, short-ciliate.
Flowering August.

Rubbly slopes of [mud] volcanic cones, also rarely on sandy-
gravelly alluvial soils of river valleys——Soviet Central Asia: Aralo-
Caspian Region (Mugodzhary), Kyzyl-Kum, Lake Balkhash Region,
Syr-Darya, Dzhungaria-Tarbagatai, Tien Shan, Pamiro-Alai. General
distribution: Dzhungaria-Kashgaria. Described from Ayaguz in
Dzhungaria. Cotype in Leningrad.

Note. Var. macrosciadia Poljak. should be recognized. It is dis-
tinguished by its slightly shorter lower cauline leaves (up to 1.5-—2.0
cm) and their 4-6 mm-long terminal lobe, as also by the larger (5-6
mm long) and densely pubescent capitula. It contains 0.76% santonin.

630

599

According to the data of Z.M. Manulkin A. juncea Kar. and Kir.
contains 1.05% essential oil. Qualitative tests for aldehydes with fuchsin
sulfurous acid, as well as for cineole with resorcin, were negative.
Carbonyl compounds are present in the oil, but the reaction to camphor
and borneol was negative.

According to the data of M.I. Goryaev and R.N. Sazonova (1959),
the plants contain 0.28-0.67% santonin. The yield of essential oil varies
from 0.28 to 1.02%; the oil contains a large percentage of camphor.
According to the data of M.I. Goryaev, A.T. Petushkova and N.A.
Sviridova, approximately 0.1% alkaloids were found.

According to the studies of M.I. Goryaev, I.M. Shabanov and L.I.
Ignatova (1954), the yield of essential oil from var. macrosciadia
Poljak. was 0.48% on the basis of air-dried matter and 0.60% on the
basis of absolute-dry matter. The oil contains 46% cineole, 35%
camphor, a small quantity of various organic acids, among which the
presence of acetic acid has been determined with some degree of
reliability. In higher fractions, there are small quantities of azulene-
forming sesquiteperenes. Moreover, the oil contains a coloring substance
which imparts a green color. However, the properties of this substance
are not yet determined. There is a certain amount of santonin. According
to I.V. Larin, due to its unfavorable chemical composition, it is eaten
little or none at all by livestock.

Series 13. Leucodes Poljak.—Annuals or biennials; hairs long,
erect.

174. A. leucodes Schrenk in Bull. Phys. Mathem. Acad. Sc. Pétersb.
III (1845) 106, 107; Fedtsch. Perech. Rast. Turk. IV, 193; Poljak. in
Tr. Inst. Khim. Akad. Nauk Kaz. SSR, IV, 96.—Exs.: GRF No. 3200.

Biennial or annual. Densely covered with white, long, semi-erect
hairs up to end of vegetative period. Root slender, vertical. Stems
solitary or few, 30-60(70) cm high, 2-5 mm thick, erect, branched
almost from base or in lower half, with more or less straight, upwardly
directed branches. Lower cauline leaves petiolate, 3-7 cm long and
2.54.0 cm wide, with petioles as long as or longer than lamina, divided
up to base into 3 palmately or pinnately parted lobes; terminal lobes
lanceolate-linear, acute, 5-10 mm long and 1.0-1.5 mm wide; middle
cauline leaves sessile, like auricles divided into ternate, simple, linear
lobes; uppermost leaves undivided, sessile, linear-lanceolate. Capitula
in lax spreading panicle, remote, less often approximate, 2-3, sessile,
upright, ovoid-oblong, 5-7 mm long. Involucral bracts, excluding
innermost, pubescent, outer bracts small, broadly deltoid or oval, inner
larger, broadly lanceolate or oblong, with narrow, brown scarious
border.

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600

Desert sands, as well as outcrops of sandstones and different gypsif-
erous deposits.—Soviet Central Asia: Lake Balkhash Region, Syr-Darya,
Kyzyl-Kum, Pamiro-Alai. Endemic. Described from sands along Ili
River. Type in Leningrad.

Note. According to the data of M.I. Kurbatov, the yield of essen-
tial oil of A. leucodes Schrenk is 0.13%. According to the studies of
E.I. Satdarova (1954), the presence of the following in the essential oil
has been established: 1) carbonyl compounds—camphor comprising
93.26% by weight of the essential oil; 2) terpenic oxides—cin-
eole=0.35% by weight of the oil; 3) alcohols and complex esters—a)
Cis-pinocampheol comprising 1.02% by weight of the oil, b) bornyl
ester of isovaleric acid comprising 1.35% by weight of the oil; 4)
sesquiterpenic compounds—a) azulene-forming sesquiterpenes compris-
ing 2.74% by weight of the oil, b) azulene comprising 0.008% by
weight of the oil; 5) phenols—cresol 0.009% by weight of the oil; and
6) acids—0.068% by weight of the oil.

According to the data of M.I. Goryaev and R.N. Sazonova (1959),
up to 1.1% santonin was found in the plant.

GENUS 1551. Mausolea Bge.':?

Bge. in Boiss. Fl. or. III (1875) 361, nom. seminudum.

Capitula hemispherical; involucre broadly goblet-shaped of two
rows of involucral bracts withering by end of vegetative period;
receptacle convex, hemispherical, glabrous. Peripheral florets few,
pistillate, fertile, corolla tube absent, less often present and pubescent,
style slightly longer than ovary, its lobes lanceolate, with somewhat
uneven margin, toothed, straight, erect or divergent, central florets
rather numerous, staminate, but with abortive pistil lacking ovary,
corolla tubular-conical, with 5 teeth, slightly hairy above, anthers linear,
apical appendages of anthers long-acuminate, basal appendages short,
round, style of abortive pistil cylindrical, strongly narrowed above and
below, stigma broadly and obtusely conical (funnel-shaped), not lobed,
with dense, upright, straight, long cilia. Achenes elliptical, densely
covered with long, stiff, appressed hairs.

A monotypic genus.

Note. The genus Mausolea Bge. usually is well distinguished by
the absence of the corolla, and by the lanceolate stigma lobes of the

'Treatment by P.P. Poljakov.
*From the Greek word Mausolos—Mausolus, the king of Caria (about 360 B.C.),
in whose memory the lavish tomb, the “Mausoleum,” was erected.

601

peripheral pistillate florets, hairy achenes, form of the style and stigma
of the abortive pistil, and also by monocarpy. As regards the pubescence
of the achenes, it resembles somewhat the North American genus
Picrothamnus Nutt., but differs from it by having straight (not curly)
hairs and also all the other characters already mentioned.

1. M. eriocarpa (Bge.) Poljak. comb. nov.—Artemisia eriocarpa
Bge. in Mem. Acad. Sc. Pétersb. VII (1854) 336; Boiss. Fl. or. III,
361; Fedtsch. Perech. Rast. Turk. IV, 190; Krasch. in Spisok Rast.
Gerb. Fl. SSSR, X (1936) 98, (1949) 48.—Exs.: Gerb. Fl. SSSR, Nos.
3189, 3290.

Perennial semishrub, 45-65(70) cm high. Tap root woody, rather
thick, with solitary or usually few, erect, woody, branched shoots
covered with smooth, whitish, cracking bark; young terminal and lateral
flowering branches light brownish, velutinous from short dense hairs.
Leaves green, pubescent; lower leaves with petioles thickened at base,
(2)3-5 cm long, pinnately cut, their lobes usually 5, linear-lanceolate,
3-7(10) mm long and 1.0-1.5 mm wide, scarcely acuminate, entire or
with 1-2 short teeth; middle and upper leaves sessile, divided, into 3—
5 lobes or simple; uppermost bracteal leaves simple, linear-lanceolate.
Capitula on short or long peduncles, globose, 4-5 mm in dia, remote,
sometimes approximate, divergent or more often drooping, in lax,
racemose-paniculate inflorescence. Outer involucral bracts roundish-
oblong, convex, succulent, later coriaceous, appressed-hairy, inner
narrower, oval-elliptical, with scarious border, subglabrous. Peripheral
florets 3-7, pistillate, usually lacking corolla tube; central florets 3-
17, staminate, corolla tubular-conical, with 5 teeth, hairy in upper part.
Achenes densely covered with stiff hairs. Flowering April.

Desert-ridged hummocky or dissected, shifting sand [dunes].—So-
viet Central Asia: Aralo-Caspian Region, Lake Balkhash Region, Kara-
Kum, Kylzyl-Kum. General distribution: Iran. Described from Kyzyl-
Kum from Lehmann’s collections. Type in Leningrad.

GENUS 1552. Turaniphytum Poljak. Gen. Nov.!:?

Poljak. in Addenda, XXV, 880 and in Fl. Turkm. VII (1960) 132,
description in the Russian.

Capitula heterogamous, with few (2-10) flowers, partly pistillate
and partly staminate, with abortive pistil; borne in clusters of 6-18 in

‘Treatment by P.P. Poljakov.
*From the distribution of the genus in the Turanian lowland and the Greek word
phyton — plant.

602

SG

Wa

2

633 Plate XXXI.
1 — Neopallasia pectinata (Pall.) Poljak., habit, peripheral pistillate floret,
style, bisexual disk floret, anther, longitudinal section of the involucre and
receptacle, placement of achenes in the capitulum;, 2 — Ajania pallasiana (Bess.)
Poljak., habit, peripheral pistillate floret, pistil of the bisexual disk floret, anther,

pistil, achene, outer and inner involucral bracts.

63

wn

603

dense, more or less globose glomerules, 3—8 mm in dia, in turn aggre-
gated in long, spicate, sometimes more or less branched at base of
inflorescence, less often capitula solitary and then with a large number
of florets (up to 27). Involucre cupulate, 1.0-1.5 mm in dia, less often
2.0-2.5 mm, with almost equal involucral bracts, some
bracts adjacent to pistillate florets carinate. Receptacle usually weakly
convex, glabrous. Peripheral pistillate florets in axils of carinate
involucral bracts; their corollas tubular, small, strongly reduced in
length in comparison to width, apically with short, irregular, 2—3 teeth;
corollas of staminate florets (having only inconspicuous rudiment or
ovary) much larger, also tubular, with 5 deltoid teeth, tube long
gradually narrowed toward throat, with long and erect hairs at apex,
more or less pinkish; anthers lacking basal appendages, apical
appendages oblong-lanceolate, acute; style in pistillate florets bifid,
their branches linear, divergent; stigma of abortive pistil in staminate
florets not divided, conical. Achenes oblong-obovoid, more gradually
narrowed toward base, somewhat flattened from sides, with
inconspicuous veins; pappus absent. Perennial, herbs, usually with many
stems from base; leaves more or less pinnatipartite.

Two species of the genus are found in Soviet Central Asia and
Kazakhstan; probably they also extend to the territories of China and
Iran.

Type of genus: Turaniphytum eranthemum (Bge.) Poljak.

1. Capitula 1.0-1.5 mm in dia, with 3-10 flowers, 6-18, compactly
clustered in more or less globose glomerules .............cccssseeseeereees
BCS Oe OE GAT, BUG, 1. T. eranthemum (Bge.) Poljak.

+ Capitula 2.0-2.5 mm in dia, not forming globose glomerules .............
CUES SER a APA eee 2. T. kopetdaghense Poljak.

1. T. eranthemum (Bge.) Poljak. comb. nova.—Artemisia
eranthema Bge. in Mém. Acad. Sc. Pétersb. VII (1854) 339; Boiss. FI.
or. III, 361; O. and B. Fedtsch. Perech. Rast. Turk. IV, 192.

Perennial. Plant 15-35 cm high, grayish from rather sparse long
hairs. Root thick and usually many-headed, with flowering and short
nonflowering shoots. Stems usually rather numerous, erect or basally
ascending, simple (less often with few lateral branches), relatively
sparsely leafy. Basal and lower cauline leaves long-petiolate, up to 8
cm long and 3 cm wide, their laminas oblong to almost round, usually
twice pinnately cut, with 1-3 pairs of lobes, terminal lobes linear or
lanceolate, subacute; upper cauline leaves with shorter petioles
(uppermost leaves often sessile) and less divided (to undivided and
entire, linear). General inflorescence spicate, usually simple, consisting

636

604

of mostly globose glomerules 3—8 mm in dia, in axils of apical leaves;
glomerules of 6-18 compactly clustered capitula with 3-10 flowers
1.0-1.5 mm in dia; corolla of pistillate florets in axils of carinate
involucral bracts, 0.5—0.8 mm long; corolla of staminate florets 2-3
mm long, with long hairs at apex and more or less pink. Achenes 1.0—
1.4 mm long. Flowering May to June. (Plate XXX, Fig. 1).

Sandy steppes and deserts.—Soviet Central Asia: Aralo-Caspian
Region (southeastern part), Lake Balkhash Region (southern part), Syr-
Darya (northern part). General distribution: Possibly Dzhungaria-
Kashgaria. Described from southern Kazakhstan. Type in Leningrad.

2. T. kopetdaghense Poljak. sp. nova in Addenda, XXV, 881 and
in Fl. Turkm. VII (1960) 134, description in the Russian.—Ic.: Poljak.
Ibid. (1960) Plate XXII.

Perennial. Plant 15-25 cm high, grayish from quite long soft hairs,
with thick, woody, many-headed root. Stems usually very numerous,
erect or basally ascending, simple, sparsely leafy. Basal and lower
cauline leaves long-petiolate, 3.5-6.0 cm long and 1.2—1.5 cm wide,
their laminas oblong, twice pinnately cut, with 2-3 pairs of lobes,
terminal lobes 3-5 mm long and 0.4—0.6 mm wide, linear, subacute;
upper cauline leaves smaller and less divided, often sessile, transitional
to undivided, linear, apical leaves. General inflorescence spicate;
capitula sessile, ovoid to subglobose, 2.0—2.5 mm in dia, more or less
crowded or partly remote. Involucre of fewer bracts. Peripheral pistillate
florets 8-9, in axils of carinate involucral bracts, their corollas small,
with 2-3 irregular teeth; staminate florets 16-18, their corollas conically
tubular, with 5 teeth bearing long hairs at apex, more or less pinkish.
Achenes 1.2-1.4 mm long. Flowering June to September.

Stony slopes at 2,400—3,000 m.—Soviet Central Asia: mountainous
Turkmenia. General distribution: Probably Iran. Described from
Kopetdag. Type (Kopetdag Mountains, 2 km east of the village of
Sibir, collected on 24.VIII. 1947 by V. Nikitin) in Leningrad.

GENUS. 1553. Neopallasia Poljak.' ?

In Bot., Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVII (1955) 429.
Capitula ovoid, in almost spicate or narrowly paniculate
inflorescences; receptacle narrowly conical, glabrous. Florets
heterogeneous; peripheral florets 3-4, pistillate, fertile, their corollas

‘Treatment by P.P. Poljakov.
2Named in honor of the naturalist P.S. Pallas.

605

637 narrowly tubular, not lobed (lacking teeth), stigma lobes narrowly linear,
weakly divergent, short-ciliate; disk florets bisexual, usually 9-16, of
which 4-8 in lower part of receptacle [outer], fertile, others at apex of
receptacle sterile, their corollas tubular-conical, bluish or pinkish, with
5 teeth, glabrous, anthers linear-lanceolate, apical appendages of anthers
round, apically attenuate, acute; stigma lobes linear, short-ciliate, after
flowering divergent. Achenes somewhat flat, narrow, deltoid, finely
sulcate, at base of receptacle in single row. Annuals with pinnately
pectinate leaves.

A monotypic genus.

1. N. pectinata (Pall.) Poljak. Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVII (1955) 428.—Artemisia pectinata Pall. Reise, III,
Anhang. (1776) 755, tab. H, h, f, 2; Willd. Sp. pl. III, 1824; Bess. in
Nouv. Mém. Soc. Nat. Mosc. III, 84; DC. Prodr. VI, 120; Ldb. FI.
Ross. II, 593; Turcz. Fl. baic.-dahur. II, 2, 66; Maxim. in Bull. Acad.
Pétersb. VIII, 529; Fedtsch. Perech. Rast. Turk. IV, 200; Pampan. in
Nuov. Giron. bot. Ital. n. s. XXXIV, 684; Kryl. Fl. Zap. Sib. XI,
2816.—Exs.: GRF Nos. 3154, 3299, (sub Artemisia pectinata).

Annual. Root slender, vertical. Stems most often few or solitary,
erect, ascending only at base, 12-40 cm high, violet or brown, more
or less densely covered with bifid flexuous hairs, with strongly reduced
or sometimes elongate branches from leaf axils, less often almost simple.
Leaves green, usually glabrous, smooth, sessile, with flat, gradually
expanded base, lamina oblong, pinnately pectinate, their lobes stiff,
filiform-subulate, with short, spine-like, cartilaginous, acuminate apex,
undivided or with 1—2 similarly narrow lobes; lower and middle cauline
leaves 1.5-3.0 cm long and 0.5—1.0 cm wide; upper bracteal leaves
somewhat shorter and narrower. Capitula sessile or subsessile, ovate,
3-—4(5) mm long, in axils of upper leaves, in groups of few or solitary
on branches, forming more or less dense, almost spicate or narrowly
paniculate inflorescence. Involucral bracts oval, smooth; herbaceous,
with wide, whitish, scarious border; outer bracts somewhat shorter and
narrower than inner. Receptacle narrowly conical, glabrous. Peripheral
florets pistillate, fertile, usually 3—4, their corollas narrowly tubular,
not lobed, white membranous, other florets bisexual, of which 4—8 at
base of receptacle fertile, remaining florets at apex of receptacle, sterile,
their corollas tubular, with 5 teeth, bluish or pinkish. Achenes 1.2-1.5
mm long, ellipsoid, somewhat flat, narrow, triangular, dark brown,
finely sulcate, in lower part of receptacle in a single closed row.
Flowering August. (Plate XXXI, Fig. 1).

Loamy and loamy-sand plains in desert-steppe zone.—Eastern
Siberia: Dauria; Soviet Central Asia: Dzhungaria-Tarbagatai (Chilikta

638

606

valley in Tarbagatai), Tien Shan (Issykkul basin). General distribu-
tion: Dzhungaria-Kashgaria, Mongolia. Described from southern Trans-
Baikal. Type in London.

Note |. It is a very distinct genus differing from all other genera
of the subtribe by having two groups of distinctly different flowers in
the capitulum. The first group, consisting of vestiges of bisexual flo-
rets situated in the upper part of the narrowly conical receptacle and
closely adjoining each other, forms a unique screen that covers the
capitulum from above. Because of this, the fertile pistillate and bi-
sexual florets (second group) situated below seem to be self-pollinat-
ing. Beside this, the form and disposition of the achenes, anthers, and
corolla of the pistillate florets further emphasize the morphological
and biological uniqueness of the genus Neopallasia, which is adapted
to the severe climatic conditions of Soviet Central Asia.

Note 2. Pampanini (Pampan. op. cit. 684) segregates A. pectinata
var. yunnanensis Pamp.—a perennial growing in Yunnan Province. On
the basis of this character, Krascheninnikov (Mat. po Istor. Fl. i Rast.
SSSR, Ill (1958) 126) even proposed to treat it as a new species—A.
yunnanensis (Pamp.) Krasch. However, the absence of collections
and the brief diagnosis force me to refrain from commenting on this
taxon.

Economic Importance. According to the data of M.I. Goryaev
and R.N. Sazonova (1959), the yield of essential oil is 0.06%; the oil
contains aldehydes, phenols, and ketones.

Tribe 8. SENECIONEAE Cass. in Bull. Soc. Philom. Paris (1815)
173 and in Dict. Sc. Nat. XX (1821) 377, XLVIII (1827) 446, LX
(1830) 582; O. Hoffm. in Pflanzenfam. IV, 5 (1890-1894) 283.—Trib.
Senecionidae Less. Subtrib. Senecioneae Less. Synops. Comp. (1832)
387; DC. Prodr. VI (1837) 292 p. p.—Trib. Senecionideae Benth. in
Benth. and Hook. fil. Gen. Pl. II (1873) 206.—Fam. Senecionidaceae
Bessey in Ann. Miss. Gard. 2 (1915) 164.—Capitula heterogamous or
less often homogamous, with distinct ray of peripheral pseudoligulate
florets or lacking ray. Receptacle usually glabrous, less often hairy.
Anthers basally obtuse, sagittate or auriculate. Style branches of bisexual
florets mostly thickened, usually truncate, less often acuminate, filiform-
clavate, without or with more or less distinct appendage; style apices
to variable length (at apex, or only below appendage or finally, up to
branching of styles) covered with papillae or hairs or tuft of hairs or
with terminal corona of upright hairs; corolla yellow, less often lilac,
reddish, white or of other shades; capitula mostly homochromous, less
often heterochromous. Leaves alternate, only very rarely (in our genera!)
opposite.

639

607

Herbs, semishrubs and shrubs, widely distributed over the world.

There are fewer arborescent forms of Senecioneae in the tropics.

KEYS TO THE IDENTIFICATION OF GENERA OF THE TRIBE

+

Senecioneae!

Flowering shoots lacking normal green leaves, with only scaly
lewvory-somietiiies Collared #i:.4:8iaguloex. alow. biG). Z22i1........ 2.
Flowering shoots with normal green leaves ..............::ccs:ceeeeeseees 4.
Plants monoecious. Flowering shoots terminating in single
comparatively large capitulum, 2.0-2.5 cm in dia, drooping at achene
maturity. Peripheral pistillate florets distinctly ligulate, many
whorled, bright- or golden-yellow ................4. 1554. Tussilago L.
Plants dioecious or almost dioecious. Flowering shoot most often
terminating in several or many capitula aggregated into spicate,
paniculate, corymbose or umbellate inflorescence; only very rarely
flowering shoots terminating in single capitulum, but then capitulum
peinindiles Yoxehan...inouwle..ciaandalowado xi. Laiska 3:
Corolla of peripheral pistillate florets distinctly ligulate. Capitula
mostly aggregated in corymbose inflorescence (very rarely capitula
solitary). Plants dioecious with small basal leaves.............
ETAL, Cl ARO, LA Aedes isan 1556. Nardosmia Cass.
Corolla of peripheral pistillate florets filiform-tubular or oblique
along margin, lacking distinct ligules. Capitula mostly aggregated
in spicate or spicate-paniculate overall inflorescence. Plants not
completely dioecious, with large basal leaves ...........ccseseeseeeeeees
$U. ETO TENS. HORE Gls a Wh ea tisete. tcl 1559. Petasites Mill.
Receptacle pubescent; leaves opposite, only in very rare cases
uppermost leaves obliquely opposite or even alternate; capitula most

often solitary, less often MaNy ...........cceseeeeeeeeeee 1558. Arnica L.
Receptacle glabrous or if hairy (less often!), and then leaves alternate
UReesedate... dos 24. a Ride.aacuies..with. deeguledate S:

Leaves evergreen (wintering), coriaceous; basal leaves (particularly
along veins beneath), as also stems (particularly, younger ones),
densely covered with thick, rusty hairs. One species; in USSR found

only mh-Carpathiane....2/.2200. acura 1557. Homogyne Cass.
Leaves not evergreen (withering before winter), or plants with other
types of hairs es ....inuisioon lea.miznilovel\ helleewas, pacts 6.

‘Prepared by M.E. Kirpicznikov.
2Usually modified petioles in origin.

64

—_

608

So +

10.

Peripheral achenes lacking pappus, central with pappus or if all
achenes with pappus, then peripheral achenes with a single row of
bristles, but with several rows in central achenes ...............:008
Sn. desacuba dl ME ERTIES, SRL 1559. Doronicum L.

All achesies*with:identical pappits (72282261. aagsne ae 7.
Capitula homogamous; all florets tubular «0.00.0... ccecceeseeseeeeees 10.
Capitula heterogamous; peripheral florets mostly ligulate or less
often filiform-tubular, different from central florets ................... 8.

Peripheral florets filiform-tubular. Exclusively introduced plants; in
USSR found only in Carpathians and Western Transcaucasia .............
niall ee tibth.. teh ieee im aes lee eds 1560. Erachtites Raf.
Peripheral:florets ligulate .3 4202.18). Ace ee ee 9.
Involucral bracts dimorphic, distinctly or indistinctly two-rowed,
inner bracts broader, with wide scarious margin, outer bracts
narrower, without scarious border or very narrow border; 1-3
additional bracts may be present at base of involucre. Style branches
acuminate or obtuse and hairy only on outside, sometimes hairy the
whole length down to their point of branching .................008
US THE Le a ee 1564. Ligularia Cass.
All involucral bracts identical, one-rowed; on outside of this
involucre at its base is found a complete row of bracts, most often
much shorter than involucre, but sometimes as long. Style branches
truncate, with crown of hairs at their very tip ...... 1563. Senecio L.
Basal leaf one, lamina dorsally pubescent until its unfolding and
involute around petiole as in compound unbel; cotyledon only one,

longitudinally revolute «0.0.0.0... 1562. Syneilesis Maxim.
Basal leaves absent, or if present then of different type; cotyledons
2 wdasededacondiveas CR IMYS IRAE OR BME «wel ee is bate eae 1.

Florets whitish or more rarely reddish; capitula with few, mostly 5—
10(12) flowers; style branches acuminate, only on outside covered

With haits 1ihe 2cva.wea a onee ae ee | 1561. Cacalia L.
Florets yellow, orange, purple, or violet; capitula many-flowered;
style branches truncate, with crown of hairs ........ 1563. Senecio L.

Subtribe Senecioninae O. Hoffm. in Pflanzenfam. IV, 5(1890-

1894) 286.—Subtrib. Tussilagineae Benth. and Eusenecioneae Benth.
in Benth. and Hook. fil. Gen. Pl. II (1873) 207, 208.—Involucre mostly
one-rowed, mostly with identical bracts, free or fused at base; often
with one more row of reduced, very short bracts at base of involucre
(forming so-called “involucrum calyculatum’’). Pappus mostly of simple,
setaceous hairs.

Note. Following basically Hoffman’s system, I retain the genus

Tussilago in the subtribe Senecioneae, although it is hardly justified.

642

609

A. Cassini with full justification erected a separate group Tussilagineae
with the rank of tribe. The same Tussilagineae, but already with the
rank of subtribe, was recognized by Bentham, and also after him by
Boissier. A separate subtribe, but by the name Tussilaginanae, is also
recognized in the work of the renowned contemporary American
specialist on the Compositae, P.A. Rydberg (Carduaceae in North
American Flora, Vol. 34, 4, 1927, p. 309).

GENUS 1554. Tussilago L.':?

L. Sp. pl. (1753) 865.—Farfara Gilib. Fl. Lith. III (1781) 177.

Capitula solitary, small, cylindrical; involucral bracts one-rowed,
lanceolate, acuminate. Receptacle glabrous, flat; central florets sterile,
bisexual; corolla tubular-campanulate, with 5 teeth; stigma undivided,
clavate; peripheral florets pistillate, ligulate, in several rows, fertile,
with bilobate stigma; achenes with 5-10 ribs, cylindrical; pappus of
numerous simple hairs. Herbaceous perennials, with long creeping
rhizome; flowering shoots covered with scale leaves. Basal leaves
appear only after flowering, roundish-cordate, angular, coriaceous,
white-pubescent beneath.

The genus is monotypic, occurring in the temperate zone of Eurasia;
introduced in America.

1. T. farfara L. Sp. pl. (1753) 865; DC. Prodr. V, 208; Ldb. FI.
Ross. II, 470; Boiss. Fl. or. Ill, 377; Schmalh. Fl. Yugo-Zap. Ross.
305; Kryl. Fl. Zap. Sib. XI, 2822; Grossh. Fl. Kavk. IV, 143; FI.
Turkm. VII, 138; Grigorjev, Opred. Rast. Okr. Stalinabada, 277; Gilib.
Fl. Lith. III, 177.—Ic.: Fedtsch. and Fler. Fl. fig. 985; Hegi, Ill. Fl. VI,
fig. 383-385, tab. 256, fig. 1, la—Exs.: Fl. pol. exs. No. 540.

Perennial. Rhizome long, creeping; flowering shoots 10-25 cm
high, covered with scaly, appressed, ovate-lanceolate, acute leaves,
mostly purple-violet. Basal leaves appear after anthesis, long-petiolate,
roundish-cordate, 10-25 cm wide, angular, with irregular teeth,
coriaceous, floccose on both sides, but later glabrous above, whitish-
tomentose with soft hairs. Capitula solitary, 2.0—-2.5 cm in dia, drooping.
Florets golden yellow; pappus 4—5 times as long as achenes. Flowering
April to May.

'Treatment by L.A. Kuprianova.
2 From the Latin words tussis—cough, and agere—to bring to motion, to enable;
named for the medicinal use of the plant.

610

Widely distributed; found on young alluvial deposits, both clayey
and sandy, or leached soils, on slopes along bottoms of ravines and on
railroad embankments; also distributed in middle mountain zone on
rock outcrops and along banks of mountain streams and rivers.—
European part: All regions; Caucasus: Ciscaucasia, Dagestan Eastern
and Western Transcaucasia; Western Siberia: Upper Tobol, Ob River
Area, Irtysh, Altai; Eastern Siberia: Lena-Kolyma, Angara-Sayans,
Dauria; Far East: Ussuri (vicinity of Vladivostok); Soviet Central Asia:
Lake Balkhash Region, Dzhungaria-Tarbagatai, Amu-Darya, Syr-Darya,
Pamiro-Alai Region, Tien Shan, mountainous Turkmenia. General
distribution: Western Europe, northern Africa, Balkans-Asia Minor,
Iran, Indo-Himalayas; introduced into North America. Described from
western Europe. Type in London.

GENUS 1555. Petasites Mill.'?

Miller, Gard. Diet. ed. 4 (1754); Gaertn. De fruct. et sem. II (1791)
406, t. 166; Cass. in Dict. Sc. Nat. XXXIV (1825) 199; DC. Prodr. V
(1836) 206.

Capitula short-cylindrical or campanulate, about 1 cm in dia, with
one-rowed involucre, aggregated into spicate-paniculate, ovoid, less
often in almost corymbose inflorescence, not fully dioecious; staminate
capitula with few peripheral, pistillate, tubular florets and numerous
central bisexual, tubular-campanulate florets with 5 teeth; pistillate
capitula with numerous peripheral, filiform, obliquely cut or weakly
ligulate, pistillate, fertile florets and few central sterile, bisexual florets.
Receptacle glabrous; style bilobed. Achenes terete, with longitudinal
ribs; pappus consisting of a few rows of simple hairs, 3—3.5 times as
long as achenes. Perennials with branched, creeping rhizomes, large
basal leaves, and flowering shoots bearing only scale-leaves (modified
petioles).

The genus Petasites is distributed in western Europe, Asia Minor,
Caucasus, the European part of the USSR, Soviet Central Asia, in the
north and south of Western Siberia, on plains and in hilly regions, not
higher than the timberline. Its species composition is insufficiently
studied; the Caucasian material especially needs a monographic
treatment.

'Treatment by L.A. Kuprianova.
*From the Greek word petasos—sun-hat; so named because of the size of the
leaves.

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611

1. Basal leaves deltoid-cordate, with 2—3 lobes at base; cauline leaves
long, up to 15 cm, lanceolate-linear, with broadened acute apex;

corolla: yellowish... iiie2 is chanathe. 1. P. spurius (Retz.) Rchb.
+ Basal leaves orbicular-cordate; cauline leaves ovate, 5-8 cm long;
corolla purplish or yellowish-White .............ccccssccsscesseesseeeereenneeens 2.
2. Rhizome in upper part tuberculate; corolla purplish, stigma lobes
VAI tak Rh tdi ee Dead 2. P. hybridus (L.) Gaertn.
+ Rhizome uniformly thickened; corolla yellowish-white; stigma lobes
linear-lanceolate, acuminate .................... 3. P. albus (L.) Gaertn.

1. P. spurius (Retz.) Rchb. Fl. exc. (1830-1832) 279; Ldb. FI.
Ross. II, 469; Hegi, Ill. Fl. VI, 691, fig. 402; Fl. Yugo-Vost. VI, 364;
Kryl. Fl. Zap. Sib. IX, 2824; Vizn. Rosl. 550; Majevski Fl. 588.—P.
niveus Ldb. Fl. Ross. II (1846) 469, non Baumg.—P. tomentosus (Ehth.)
DC. Prodr. V (1836) 207; Schmalh. Fl. I, 80.—Tujssilago spuria
Retz. Obs. I (1779) 29, t. 2.—T. tomentosa Ehrh. Beitr. III (1788)
65.—Nardosmia spuria Less. in Linnaea, IX (1834) 196.—Ic.: FI.
Yugo-Vost. VI, 364, fig. 688.—Exs.: GRF No. 219; Fl. Polon. exs.
No. 539.

Perennial. Rhizome long, creeping, 5-10 mm thick, with filiform
roots at nodes. Basal leaves coriaceous, with long, white-tomentose
petioles, 6-12 cm long, deltoid-cordate, with 2-3 lobes at base,
irregularly and finely sinuate-toothed, young leaves white-tomentose
on both sides, older leaves subglabrous or floccose above, white-
tomentose beneath. Stems up to 70 cm high. Cauline leaves amplexicaul,
glabrous, broadly lanceolate; upper leaves lanceolate-linear, usually
with long, thin, attenuate apex, 8-15 cm long and 2-3 cm wide.
Inflorescence corymbose-spicate with 12—15(30) florets [capitula?].
Capitula 5-8 mm long, 1.0-1.5 cm in dia; pistillate florets white or
whitish-yellow; stigma of bisexual florets ovate. Achenes a fourth to
a third as long as pappus, terete, somewhat ribbed, 1.5—2.0 mm long;
pappus hairs in many rows, with several cilia. Flowering and fruiting
May.

Along sandy banks of rivers and lakes, sometimes forming
continuous stands, often in forest-steppe and steppe regions.—Euro-
pean part: Dvina-Pechora, Baltic Region, Ladoga-IImen, Upper Volga,
Volga-Kama, Upper Dnieper, Middle Dnieper, Volga-Don, Trans-
Volga, Upper Dniester, Bessarabia, Black Sea Region, Lower Don;
Western Siberia: Ob River Area, Upper Tobol (west), Irtysh; Soviet
Central Asia: Aralo-Caspian Region (northern part), Tien Shan (north),
Lake Balkhash Region (east), Syr-Darya. General distribution: Central
and northern Europe. Described from western Europe.

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612

2. P. hybridus (L.) Gaertn. Mey. and Scherb. Fl. Wett. III
(1801) 184; Hegi, Ill. Fl. VI, 683; Szaf. Kulcz. Pawl. Rosl. Polskie,
631; Grossh. Fl. Kavk. IV, 144.—P. officinalis Moench. Meth. (1794)
568; Ldb. Fl. Ross. II, 468; Schmalh. Fl. 79; Vizn. Rosl. 550
.—Tussilago hybrida L. Sp. pl. (1753) 866.—T. petasites MB. FI.
taur.—cauc. II (1808) 305.—P. georgicus Mand. in Zam. po Sist. i
Georg. Rast. Tbil., 13 (1947) 76; Fl. Gruzii, VIII, 389.—Ic.: Hegi, Ill.
Fl. VI, taf. 266, fig. 2.—Exs.: Pl. Finl. exs. No. 392; Fl. Polon, exs.
No. 357.

Perennial. Rhizome long, thick, in upper part nodulose, tuberculate,
up to 3-5 cm. Basal leaves long-petiolate, larger, up to 60 cm and
more wide, roundish-deltoid, cordate, with 2—3 lobes on each side,
with irregular, small, deltoid teeth, subglabrous, scabrous from small
spiny hairs, grayish-white, tomentose beneath. Stems up to 60 cm high,
10-20 mm thick, white-tomentose. Cauline leaves (leafy petioles)
sometimes with considerable lamina, basally violet, white-tomentose
in upper part, lower leaves broadly ovate, 5-10 cm long and 1.5-—2.0
cm wide, upper lanceolate. Inflorescence dense, spicate, with many
florets [capitula?], 6-10 cm long, up to 20 cm at fruiting; capitula S—
8 mm long, about 6-8 mm in dia. Florets dirty purple, with violet
tinge, aromatic, style lobes in bisexual florets short and obtuse.
Flowering April to May.

Wet forest meadows, along banks of rivers and streams.—Euro-
pean part: Ladoga-IImen, Upper Volga, Upper Dnieper, Middle
Dnieper, Bessarabia, Black Sea Region, Crimea, Lower Don; Caucasus:
Ciscaucasia, Dagestan, Eastern and Western Transcaucasia (Baku),
Talysh (Lenkoran). General distribution: central Europe, southern
Scandinavia. Described from western Europe. Type in London.

Note. P. georgicus Mand., described from the vicinity of
Tbilisi, differs from P. hybridus (L.) Gaertn. by having a yellowish-
white corolla and, as I.P. Mandenova reports, a thin acuminate stigma
in the staminate florets; all other characters (size and form of the
capitula, structure of the inflorescence, leaves, and rhizome) are the
same as in P. hybridus. Final resolution of the question whether P.
georgicus is a separate species must be deferred until more complete
collections are available.

3. P. albus (L.) Gaertn. De fruct. et sem. II (1791) 406; Ldb. FI.
Ross. II, 469; Szaf. Kulcz. Pawl. Rosl. Polskie, 632; Schmalh. FI. II,
79; Vizn. Rosl. 550; Grossh. Fl. Kavk. IV, 144.—Tussilago alba L.
Sp. pl. (1753) 866.—Ic.: Hegi, Ill. Fl. IV, 693, fig. 405.

Perennial. Rhizome long, not thick, nodulose, with filiform roots.
Basal leaves orbicular-cordate, with angular, acute and coarse teeth,

645

613

cordate, subglabrous above, whitish-pubescent, floccose beneath. Stems
slender, 30-60 cm high, white-tomentose, arachnoid-hairy. Cauline
leaves (petioles of leaves) lacking lamina, oblong-ovate, along margin
and in upper part arachnoid-white-tomentose, attenuate-acuminate, 2—
3 cm long and 5-8 mm wide. Inflorescence lax, almost corymbose at
flowering, later spicate-paniculate, 15-30 cm long, with 10—15(25)
flowers. Capitula 15 mm long and 5-8 mm in dia. Involucral bracts 6-
8 mm long. Florets greenish-yellowish-white; style lobes of bisexual
florets linear-lanceolate, acute. Flowering April to May.

Banks of rivers and mountain streams.—European Part: Baltic
Region, Upper Dniester; Caucasus: Ciscaucasia, Eastern and Western
Transcaucasia. General distribution: Northern and Central Europe,
Atlantic Europe. Described from western Europe. Type in London.

Note. P. kablikianus Tausch. is similar to P. albus (L.) Gaertn.; it
is distinguished by leaves that have smooth margins and lack coarse
teeth, the absence of conspicuous white hairs, and a yellowish-white
corolla. This species does not occur in the USSR; it is distributed in
the Sudety and Tatra mountains and was mistakenly cited in the Vizn.
Rosl. URSR for the Soviet Carpathians.

GENUS 1556. Nardosmia Cass.!:?

Cass. in Dict. Sc. Nat. XXXIV (1825) 186; Ldb. Fl. Ross. II
(1844-1846) 466; DC. Prodr. V (1836) 205.—Tussilago L. Sp. pl.
(1753) 865, pro parte.—Petasites Kom. in Tr. Peterb. Bot. Sada, XX V
(1907) 683, non Mill.

Capitula with one-rowed involucre, dioecious; staminate capitula
with small, numerous, pistillate, distinctly ligulate florets and numerous,
sterile, bisexual florets with 5 teeth; pistillate capitula with numerous
peripheral ligulate pistillate fertile florets and few, central, bisexual
florets. Receptacle glabrous. Style of tubular-infundibuliform florets
not thickened below stigma (except N. japonica S. and Z.); stigma
bifid, its lobes lanceolate, oblong-lanceolate, or linear, less often short,
oval or short-lanceolate. Achenes terete, with longitudinal ribs; pappus
many-rowed, consisting of simple hairs, 2.5-3.5 times as long as
achenes. Perennial, with creeping, branched rhizome; basal leaves
smaller than those of Petasites, flowering shoots bearing scale leaves
(modified petioles).

‘Treatment by L.A. Kuprianova.
*From the Latin words nardus—the ancient name of an aromatic balsam, and
osme—aroma; named for the aroma of the plant.

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614

Species of this genus are distributed in Eurasia, North America,
Japan, Sakhalin, Kuril Islands, China and Mongolia. They grow in
mountainous regions on stony slopes and balds, in alpine meadows,
tundra zone, peat bogs and along banks of rivers and streams in the
north of the forest zone.

Type of genus: N. denticulata Cass.

Note. The genus Nardosmia is similar to Petasites. However, it
usually has been kept separate by the authors of Russian floras. The
main differences between these genera are as follows: the genus
Nardosmia has a distinctly ligulate corolla in the peripheral pistillate
florets, a corymbose (not spicate) inflorescence, and small basal leaves.
The range of the genus Nardosmia mainly extends over Eurasia and
North America, whereas that of the genus Petasites is mainly Central
Europe. Ecologically, these genera, also are different. Species of the
genus Petasites are mainly found in the forest or mountain-forest zones,
often in association with the broad-leaved forests, while species of the
genus Nardosmia, with few exceptions, are the members of tundra
zone or high-altitude regions.

1. Plants short, with solitary capitulum or, less often only in pistillate
forms, with 2-3 capitula. Rhizome slender, cordlike, up to 2 mm

thick Ai:..ik. es. stapes bes 4. GRRE RNA bcos ee Zn
+ Plants tall, with numerous capitula aggregated in corymbose
inflorescence; rhizome thicker, up to 2-3 CM ........eseeseeesseeeeteeees 3.

2. Leaves oval, cuneate at base, glabrous above, white-tomentose
beneath; capitula in pistillate forms sometimes 2-3 ...........s:cceceeeeeees
JSD be, LEA: AEE eee Ce 8. N. gmelini (Turcz.) DC.

+ Leaves deltoid, with cordate base, glabrous on both sides, capitula
always ‘solitairyoisi til bition eae 9. N. glacialis Ldb.

3. | Leaves palmately incised or palmately dissected almost halfway

into 7-8 wide lobes, each in turn cut into 3 acute lobes; florets light

violet.or Whites: late ine..Aaee ey 6. N. palmata (Ait.) Hook.

Leaves entire, ' finely: or coarsely toothed sus. vsis.$ecsiisdsiiedaens aedeacbans 4.

Leaves 60 cm, up to 1.5 cm wide, finely and acutely toothed,

cauline leaves of staminate individuals broadly ovate, up to 5 cm

wide, oblong in fertile pistillate forms, 1.5—2 cm wide .............0+4+

Ait. inns. boddeesd . niciesouwe gil 7. N. japonica S. and Z.

Leaves: smallertiiive gia hens ii divind dle Rib Made gaeaes Si.

Florets yellow, whitish-yellow or white; leaves deltoid-reniform or

reniformi: ‘finely toothed! ...a.ic i Nn chnte Rg a 6.

+ Florets reddish, pink, or white; leaves deltoid, coarsely toothed
sevo\desandznsdawtergh vies oifcbom anna itddce <A iat Rs Hise cteaiel in ecee: WORE tees eens ena 8.

> +

M+

615

6. Leaves deltoid-reniform, (10)20—35 cm wide, glabrous on both sides,
coriaceous; florets yellow; capitula 7-15, in corymbose inflorescence
bode ee. RUS BREA OR Aa 5. N. laevigata (Willd.) DC.

+ Leaves reniform, with round apex, 2—7 cm wide and 2-5 cm long;
florets white; capitula 3-10, in corymbose inflorescence .......... TN,

7. Petioles of basal leaves somewhat expanded, amplexicaul; pistillate
florets 3.0-3.5 mm long, with 1 mm-long ligule; staminate florets
O=boninflengins i ticde i Ani Aine 1. N. saxatilis Turcz.

+ Petioles of basal leaves hardly expanded, pistillate florets ligulate,
limb 5—6 mm long, 2.0—2.5 mm wide; staminate florets 8.0-9.5 mm
Lom pele etiaihds Sena cise eee 2. N. fomini Bordz.

8. Leaves broadly deltoid, cordate, 15-20 cm long and up to 20 cm
wide, coarsely toothed, teeth 3-4 cm long and as wide, deltoid and
acuminate, in turn divided into unequal, acuminate, small teeth,
grayish-tomentose beneath, glabrous above. Florets white or pink
BLT Gi KL AOA LE IN, cts 3. N. angulosa Cass.

+ Leaves deltoid or oblong-deltoid, 3-6 cm long and 3—4(6) cm wide,
with 7-8 coarse, obtuse teeth. Florets reddish. ................scssescssssssenes
Mix tatinn ell uti Aaaea Minn tis: 4.N. frigida (L.) Hook.

Subgenus 1. Nordosmia.—Inflorescence corymbose, with numer-
ous capitula. Style of central tubular-infundibuliform florets thickened
or not below stigma. Stigma bilobed, with oblong-lanceolate lobes.

Section 1. Nardosmia.—Peripheral florets ligulate; style below
stigma lacking a ring but thickened.

Series 1. Saxatiles Kuprian.—Leaves small, reniform, with round
apex, sinuate-toothed with acute teeth; florets white. High-mountain
plants. Besides our species, we should also include N. fragrans (Vill.)
Rchb. and N. dérfleri (Hay) Kuprian. comb. n. from southern Europe.

1. N. saxatilis Turcz. in Bull. Soc. Nat. Mosc. I (1838) 94 nomen;
ibid., XIX, 11 (1846) 138; FI. baic.-dahur. II, 4.—N. laevigata
subfoemina DC. Prodr. V (1836) 205, pro parte; Ldb. Fl. Ross. II,
466; Kryl. Fl. Zap. Sib. XI, 2826.—Petasites saxatilis Kom. in Tr.
Peterb. Bot. Sada, XXV (1907) 684.—Ic.: Gmel. Fl. sib. II (1749) tab.
68, fig. 2.

Perennial. Rhizome slender, 2-3 mm thick, filiform. Basal leaves
small, petiolate, 3-10 cm long, reniform, with round apex, cordate, 2—
7 cm wide, 2—5 cm long, sinuate sharp-toothed, petiole and particularly
along veins covered with short thick hairs. Stems 5—15(30) cm high,
white-tomentose above. Cauline leaves amplexicaul, ovate-lanceolate,

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616

1.5-2.0 cm long, 0.5—1.0 cm wide, glabrous, along margin with short,
white-tomentose, fimbria. Capitula 9-10, conical, 9-12 mm long, about
10 mm wide, in corymbose inflorescence. Florets white; sterile capitula
with only staminate florets having infundibuliform corolla, 9-11 mm
long; styles of staminate florets thickened below stigma, scarcely
exceeding corolla; stigma shortly bilobed, pubescent; pistillate florets
3.0-3.5 mm long, with ligule about 1 mm long, with 2-3 teeth at tip,
styles of pistillate florets not thickened below stigma. Achenes smooth,
1/4 as long as pappus. Flowering May to June; Fruiting June.

Alpine regions and adjoining forests, stony mountain slopes, high
balds, on banks of streams.—Western Siberia: Altai; Eastern Siberia:
Angara-Sayans, Dauria; Far East: Zeya-Bureya, Uda River Area. Gen-
eral distribution: northern Mongolia, northeastern China. Described
from the collection of Turczaninow from Kultuk. Type in Leningrad.

Note. N. saxatilis Turcz. greatly resembles the alpine Caucasian
species N. fominii Bordz., the only member of the genus Nardosmia in
the Caucasian flora.

2. N. fomini Bordz. in Zap. Kievsk. Obshch. Estestv. XXV (1915)
123; Grossh. Fl. Kavk. IV, 144.—Ic.: Bordz. Ibid., Fig. 5.

Perennial. Rhizome branched, 1-2 cm thick, filiform. Basal leaves
long-petiolate, cordate-reniform, 3-6 cm wide, 2—4 cm long, apically
roundish, sinuate-sharp-toothed, glabrous; petioles glabrous, at base
slightly broadened and somewhat amplexicaul; cauline leaves
amplexicaul, ovate or ovate-lanceolate. Stem 12-25 cm high, in upper
part arachnoid-white-hairy. Capitula large, 1.5 cm in dia, 5—7 in
corymbose inflorescence. Involucral bracts oblong-lanceolate,
aeuminate, ventrally glabrous, with short, papilliform hairs at base.
Florets white, peripheral florets pistillate, ligulate, ligule (4.5)—5-6
mm long and 2.0-2.5 mm wide corolla of staminate florets
infundibuliform, 8.0—9.5 mm long, their styles slightly exserted from
corolla, bifid, weakly thickened. Achenes terete; pappus bristles simple,
2.5-3 times as long as achene. Flowering May.

Alpine meadows.—Caucasus: Main Range. Endemic. Described
from cultivated specimens brought from Gudaur District. Type in Tbilisi.

Series 2. Frigidae Kuprian.—Plants with deltoid leaves, with
coarse, undivided, deltoid teeth or teeth in turn divided into unequal,
acute small teeth; florets reddish, pink, or bright white. Plants of tundra
plains and peat marshes of the north of the forest zone.

3. N. angulosa Cass. Dict. Sc. Nat. XXXIV (1825) 188, excl. syn.
Linn.; Less. in Linnaea, VI (1831) 107.—N. frigida Hook. Fl. bor. am.

649

617

I (1833) 307; DC. Prodr. V, 205; Ldb. Fl. Ross. II, 467; Kryl. Fl. Zap.
Sib. XI, 2823, pro parte.—Petasites frigidus Fries Summa Veg. Scand.
(1846) 182, 4.—Ic.: Perf. Fl. Sev. Kr. 107, Gmel. FI. sib. IV, t. 70.—
Exs.: GRF No. 1222.

Perennial. Rhizome creeping. Basal leaves long-petiolate, broadly
deltoid, cordate, 15—20 cm long and up to 20 cm wide, deeply sinuate,
with coarse acute teeth up to 3-4 cm long and broadly deltoid, in turn
divided into unequal, acute small teeth. Leaves grayish-tomentose
beneath, glabrous above, younger leaves arachnoid-hairy. Stems 10-
40 cm high, up to 70 cm at fruiting, with amplexicaul leafy petioles,
lower leaves ovate-lanceolate, upper lanceolate, acuminate, sometimes
with vestigeal lamina. Inflorescence long, spicate; capitula 1.5 cm in
dia, on long peduncles elongating after flowering. Florets bright white
or pink; staminate capitula with one whorl of peripheral ligulate florets,
with 4-6 mm long ligule, projecting above involucre, much shorter in
pistillate capitula. Style much longer than corolla, not thickened below
stigma, stigma bilobed with oblong, pubescent lobes. Achenes smooth,
a fourth to a third as long as pappus, pappus consisting of many simple
hairs. Flowering May to June, beginning July.

Peat bogs of north of forest region and mountains of Siberia.—
European part: Karelia-Lapland, Dvina-Pechora, Volga-Kama; West-
ern Siberia: Ob River Area, Irtysh (east), Altai; Eastern Siberia:
Angara-Sayans, Dauria, Lena-Kolyma, Kamchatka (occasionally). Gen-
eral distribution: Scandinavia, Mongolia (north), North America. De-
scribed from Siberia from Gmelin’s material. Type in Leningrad.

Note. Ledebour (op. cit.) treated the Kola tundra plants under the
name N. frigida B., distinguishing them from the usual N. frigida that
grows in the marshes of the forest zone. The material in our hands,
from the arctic and tundra regions of the Soviet Union, is, in fact,
somewhat different from the forest and montane Siberian material.

Tussilago frigida, described by Linnaeaus (Sp. pl. 1753, 865), was
characterized already earlier in the flora of Lapland (Fl. Lapp. 1737,
303) as the plants growing in the lowlands of the Lapland alpine re-
gion and in the bordering forests with deltoid leaves and acute apices
but with undulate-acuminate margins with 7-8 teeth.

Thus, the Linnaean name T. frigida must be applied to the arctic
and tundra circumpolar species; the plants from the forest zone with
toothed leaves having coarse and acute teeth should be called N.
angulosa Cass.

4. N. frigida (L.) Hook. Fl. bor. I (1833) 307 quoad nom.; DC.
Prodr. V, 205.—Nardosmia frigida B. Ldb. Fl. Ross. II (1844-1846)
467.—N. frigida auct.: Perf. Fl. Sev. Kr. 360, pro parte; Kryl. Fl. Zap.

650

618

Sib. XI, 2823, pro parte; Popov, Fl. Sr. Sib. II, 740, pro parte.—
Tussilago frigida L. Sp. pl. (1753) 865.—T. corymbosa R. Br. in Suppl.
to Parry’s first Voyage (1824) CCLXXIX.—Exs.: Herb. norm. Nos.
5286, 5285.—Ic.: Hultén, Fl. Al. and Jukon, X, 1580, fig. 3a, b, c, f.

Perennial. Rhizome creeping. Basal leaves petiolate, with 3-4 cm
long petiole, deltoid or oblong deltoid, 3-6 cm long and 3—4(6) cm
wide, cordate, with acute teeth, in typical with 7-8 coarse, obtuse,
deltoid teeth, glabrous above, white-tomentose beneath. Stems 10—20
cm long, with leafy petioles, lower leaves ovate-lanceolate, upper
lanceolate, 1-2 cm and 2—4 cm long; acuminate, often with vestigeal
lamina. Inflorescence corymbose (and not spicate), somewhat lax at
fruiting; capitula on longer peduncles; florets reddish, 1.5 cm wide;
structure of florets and achenes as in previous species. Flowering end
of July, August.

Marshy tundras, along banks of rivers and streams.—Arctic: Arctic
Europe, Novaya Zemlya, Arctic Siberia, Chukotka, Anadyr; Far East:
Kamchatka, Okhotsk. General distribution: Arctic, Scandinavia
(northern part), North America (arctic region). Described from high
altitudes of Lapland. Type in London.

Note. Typical N. frigida grows on the Kola Peninsula along the
coast of the Barents Sea, on the islands of the Arctic Ocean and on
Novaya Zemlya, the Polar Urals and the Yamal Peninsula. Plants found
to the east of the Yamal relegated by me to this species are somewhat
different with respect to the cutting of the lamina, but have a similar
corymbose inflorescence. Similar plants from North America were
named N. corymbosa by Ledebour (FI. Ross. II, 467, 468). These
plants are hybrids between the amphi-Pacific species N. palmata,
occurring in America and Asia, and the circumpolar species N. frigida.
Typical N. palmata is not found in the range of the hybrids. Thus,
these hybrid forms could be considered a separate species of hybrid
origin. It appears, however, that these plants are extremely polymorphic
to the extent that Rydberg (Rydb. in North. Am. Fl., 34, 1927, 311-
115{sic.]) has described several species from the North American
material alone. In the Flora of Alaska and Yukon Hultén ignores
Rydberg’s species, listing some of them as synonyms of the hybrid
forms, and accepts only one species, Petasites hyperboreus Rydb.,
which, apparently, is also one of the extreme hybrid forms between N.
palmata and N. frigida.

Series 3. Laevigatae Kuprian.—Leaves deltoid- reniform, glabrous
on both sides; florets yellow. A monotypic series.

65

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619

5. N. laevigata (Willd.) DC. Prodr. V (1836) 205, submascula,
pro parte; Ldb. Fl. Ross. II, 467; Perf. Fl. Sev. Kr. 360; Krascheninn.
Fl. Yugo-Vost. Evrop. Ch. SSSR, VI, 364; Kryl. Fl. Zap. Sib. XI, 282;
Bobrov in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX, 18.—
Tussilago laevigata Willd. Sp. pl. III, 3 (1804) 1969.—Petasites
laevigatus (Willd.) Rchb. Fl. Germ. exs. (1830) 279, quoad nomen. pl.
germ. excl.—N. straminea Cass. in Dict. Sc. Nat. XXXIV (1825) 189.—
Ic.: Ldb. Ic. Fl. Ross. IV, tab. 341; Geml. FI. sib. II, t. 69.—Exs.:
GRF No. 2024.

Perennial. Rhizome thick, 0.5—1.0 cm thick, long. Basal leaves
long-petiolate, with bluish-violet, 20—35(55) cm-long petioles, lamina
deltoid-reniform, short-acuminate, broadly sinuate-toothed, cordate,
glabrous on both sides, coriaceous, 10-25(35) cm wide and 5-20 cm
long. Stems 15-60 cm high, arachnoid-hairy above. Cauline leaves
ovate-lanceolate, 3-8 cm long, glabrous. Capitula in corymbose
inflorescence, 7—10(15). Florets yellow or yellowish. Peripheral
pistillate florets ligulate, one-whorled; corollas of staminate florets
infundibuliform, 5-6 mm long, their styles almost as long as corolla,
pubescent below stigma, with inconspicuous thickening, also pubescent;
stigma bilobed, lobes oblong. Involucral bracts lanceolate 6 mm long
and 2 mm wide in middle. Achenes smooth, about 3 mm long. Flowering
April to June.

Gravelly banks of rivers and mountain streambeds, on shoals and
abundant in the water.—Arctic: Arctic Europe, Arctic Siberia (Polar
Urals); European part: Karelia-Lapland, Dvina-Pechora, Volga-Kama
(middle and southern Urals); Western Siberia: Ob River Area, Altai;
Eastern Siberia: Yenisei, Angara-Sayans, Dauria. Endemic. Described
from Lake Baikal Region. Type specimen—Plate 69 in the Flora Sibiri
[Flora of Siberia] by Gmelin.

Section 2. Brevilobae Kuprian.—Style of tubular-infundibuliform
florets thickened below stigma and with a ring; stigma lobes short,
lanceolate or ovate, 0.3-0.5 mm long.

Type of section: N. palmata (Ait.) Hook.

Series 4. Palmatae Kuprian.—Leaves divided into 7-8 broadly
cuneate lobes, in turn divided into acute lobules in turn with unequal,
coarse and sharp teeth; florets violet or white. Several North American
species, somewhat resembling N. palmata, are included in this series:
N. arctica (Porsild) Kuprian. comb. n. h. 1., N. speciosa Nutt. and
others.

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6. N. palmata (Ait.) Hook. Fl. Bor. am. I (1835) 308; DC. Prodr.
V, 206.—Tusillago palmata Ait. Hort. kew. III (1789) 188, t. 11.—
Petasites palmata A. Gray in Bot. Calif. I (1876) 407; Kom. and Allis.
Opred. Rast. Dalnevost. Kr. II, 1043.—Petasites tatwakianus Kitamura
in Acta Phyt. et Geobot. IX (1940) 64.

Perennial. Rhizome thick, creeping, about | cm thick. Basal leaves
with petioles 15-40 cm long, almost round or roundish-reniform,
cordate, 15-60 cm wide, palmately dissected almost up to half, 7-8
broadly cuneate lobes, usually trifid into acute lobes in turn with
unequal, coarse and large sharp teeth, leaves glabrous above, finely
white-pubescent beneath. Stems thick, (15)60—100 cm wide at fruiting,
with amplexicaul, leafy, ovate-lanceolate petioles, 4-7 cm long and
1.5—4.0 cm wide, with finely tomentose fimbria. Inflorescence more or
less corymbose, at fruiting somewhat lax, with elongated peduncles.
Florets light violet or white, with pleasant aroma; style of tubular-
infundibuliform florets slightly exceeding corolla, with ring below
stigma, conically thickened; stigma lobes short, lanceolate, papillate,
staminate capitula homogamous or heterogamous, with pistillate ligulate
florets. Flowering May to June.

Sandy and stony banks of forest rivers and streams, in groups
forming large stands.—Far East: Zeya-Bureya, Uda River Area,
Sakhalin, Ussuri. General distribution: northeastern China, North
America from southern Alaska to California. Described from cultivated
specimens. Type in London.

Note. Kitamura segregates the Far Eastern plants relegated by the
Russian flora specialists to N. palmata (Ait.) Hook. from the North
American plants, describing them under the name N. tatewakianus.
The main differences being that in N. tatewakianus the leaves are
palmately lobed, the ligules of the florets are 1 mm long, and the
achenes 3.0-3.5 mm long, whereas in N. palmata the leaves are deeply
palmately dissected, the corolla is distinctly ligulate, the ligules are up
to 5 mm long, and the achenes 2 mm long. At present it is not possible
to resolve the question of the separate status of the Far Eastern race
because of the great diversity of the North American material and
nonavailability to me of the type specimen of the species.

Hybrids of N. palmata and N. japonica are found.

Series 5. Japonicae Kuprian.—Ligulate florets absent; pistillate
florets with infundibuliform, obliquely divided corolla.

7. N. japonica S. and Z. Fl. Japon. (1843) 181.—Petasites
giganteus Fr. Schm. ex Trautv. in Tr. Peterb. Bot. Sada, VIII (1883)
433, non Fuss. 1866.—P. japonicus Kitam. Compost. Japon. III (1940)

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162.—P. japonicus var. giganteus Makino in Bot. Mag. Tokyo, 23
(1909) 18; Kitam. Compos. Japon. III, 164.—P. japonicus F. Schm.
ex Kom. and Alis. Opred. Rast. Dalnevost. Kr. II (1932) 1043.—P.
amplus Kitamura in Acta Phyt. et Geobot. I (1932) 115.

Perennial. Rhizome 1.0-1.5 cm thick, in upper part tuberculate, up
to 2.0-2.5 cm thick. Basal leaves long-petiolate, petioles up to 2 cm
long, very large, (30)60 cm—1.5 m wide, reniform, with small teeth,
dark green and almost glabrous above, light colored beneath,
particularly younger leaves covered on both sides with short, white,
arachnoid-hairy pubescence. Stems (15)60 cm high, strongly elongated
at fruiting, covered with short, thick hairs. Cauline leaves amplexicaul,
oblong-lanceolate to broadly ovate, (2)5-8 cm long and 1.5-5.0 cm
wide, acuminate, covered with very short, thickish hairs, narrower in
fertile individuals than in sterile. Capitula conical, 1.0—-1.5 cm in dia,
20-30, in corymbose inflorescence; florets white, tubular,
hermaphrodite; sterile capitula with only staminate florets having
infundibuliform corolla; pistillate capitula heterogamous; corolla of
pistillate florets obliquely cut, 10 mm long, style much exceeding
corolla, cylindrically thickened below stigma, with ring, stigma short-
bilobed, lobes ovate, acuminate, papillate. Involucral bracts broadly
lanceolate, membranous, with prominent veins. Achenes smooth, a
fourth to a third as long as pappus. Flowering May; fruiting June.

North facing slopes of conical hills, on riverbanks, gravel beds.—
Far East: Sakhalin (and Kuril Islands). General distribution: Northern
Japan. Described from Japan. Type in Florence.

Note. Petasites amplus Kitamura apparently is very similar to N.
japonica §. and Z. and is distinguished by larger basal leaves, with the
lamina up to 1.5 m wide and having an almost smooth margin with
only fine acuminate teeth.

The leaves of N. japonica S. and Z. are extraordinarily similar to
the European Petasites albus. There is no need for the name P. amplus
Kitamura since the epithet “gigantea” was not used in the genus
Nardosmia.

If it were proved that this plant should be recognized, then it
would have to be called N. gigantea (Fr. Schmidt).

Kitamura described Petasites himalaicus Kitam. (Acta Phyt. et
Geobot., XV, 4, 1954) from Nepal, where this species is found in
birch groves at 3,300 m. This species should be referred to the genus
Nardosmia and called N. himalaica (Kitam.).

Subgenus 2. Endocellion (Turcz. ex Herd.) Kuprian. comb. n.—
Endocellion Turcz. ex Herb. in Bull. Soc. Nat. Mosc. XXXVIII (1865)
375.—Small plants with solitary capitula or less often, in pistillate

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specimens, with 2-3 capitula; stigma of central tubular-infundibuli-
form florets hairy, stigma lobes linear. Plants of mountain tundra and
tundra plains.

Type of subgenus: Nardosmia gmelini Turcz. ex DC.

8. N. gmelini Turcz. ex DC. Prodr. VII (1838) 271; Ldb. Fl. Ross.
II, 2, 466.—Tussilago gmelini Turcz. in Herb. (1835).—Hardosmia
populifolia (Smith) Schischk. in Fl. Zap. Sib. XI (1949) 2826, non
Perdicium populifolium Smith in Rees, Cycl. XXVI (1819) No. 4.—
Endocellion boreale Turcz. ex Herd. in Bull. Soc. Nat. Mosc. XXX VIII
(1865) 375.—Petasites gmelini (Turcz. ex DC.) Polunin in Rhodora,
LIII (1951) 288.—Ic.: Gmel. Fl. sib. II (1749) 141, t. 67, fig. 1.

Perennial. Rhizome slender, 2—3 mm thick. Basal leaves with 2-
5 cm long petioles, oval, cuneate at base, 2-4 cm long and 0.5—1.5 cm
wide, weakly sinuate-toothed or entire, glabrous above; white tomentose
beneath. Stems 10-30 cm high. Cauline leaves 0.7—3 cm long, brownish-
violet. Capitula solitary or 2-3 about 2 cm in dia. Involucral bracts
oblong, violet, acuminate, white-arachnoid-hairy. Pistillate capitula 2—
3, fertile, with numerous florets, 2-3 peripheral whorls with 8.0—8.5
mm-long ligules; style of ligulate florets with short, bilobed stigma,
stigma lobes ovate; florets of inner whorls with shorter ligules; florets
in center of capitulum campanulate, small, style of central florets with
oblong pubescent-hairy and almost up to base bifid stigma. Achenes
3—4 mm long, smooth, cylindrical, with indistinct ribs; pappus 2.5
times as long as achene. Flowering June to July.

Rubbly tundra in montane regions and on plains.—Arctic: Arctic Siberia
(Polar Urals, Taimyr, Vitim Plateau); Chukotka, Anadyr; Eastern Siberia:
Angara-Sayans, Dauria; Far East: Okhotsk. General distribution:
Mongolia (north). Described from Gmelin’s material. Type—Plate 67,
Fig. 1 in the “Flora Sibiri” [Flora of Siberia] by Gmelin.

9. N. glacialis Ldb. Fl. Ross. II, 2 (1844-1846) 466; Herd. in
Bull. Soc. Nat. Mosc. XXX VIII, 370.—N. billingsiana Fisch. ex Herd.
I. c. nomen.—Petasites glaciales (Ldb.) Poiunin in Rhodora, LII (1951)
228.

Perennial. Rhizome slender, 2 mm thick, long. Basal leaves with
1-5 cm long petiole, deltoid, 2-3 cm wide, glabrous on both sides,
somewhat cordate, with coarse and unequal teeth, teeth broadly deltoid,
acuminate, 0.5-1.0 cm at base, 0.5—1.0 cm long. Stems slender, 10-20
cm high, younger stems white-arachnoid hairy above. Cauline leaves
lanceolate, upper smaller, linear-lanceolate, with narrow, white-
tomentose, fimbria. Capitula solitary terminal, 10-12 mm in dia.
Involucral bracts lanceolate, acuminate, at base densely crisped hairy.

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623

Central florets tubular-infundibuliform, with oblong, hairy stigma 1
mm long, bifid almost to base. Peripheral florets pistillate, ligulate,
ligule ovate, apically slightly sinuate, 3-4 mm long, in one whorl;
stigma of pistillate florets undivided, slightly thickened, short-papillate.
Pappus 2.5 times as long as achene; achenes terete, smooth, about 2
mm long. Flowering July to August.

Dry stony slopes and in tundra.—Arctic: Arctic Siberia (to east of
Taimyr), Chukotka, Okhotsk. Endemic. Described from Arctic Siberia.
Type specimen apparently lost.

Note. In restoring Endocellion Turcz. ex Herd. to the rank of
genus, the name Endocellion glaciale (Edb.) should be accepted for
this species.

GENUS 1557. Homogyne Cass.!:?

Cass. in Bull. Soc. Philom. (1816) 198; DC. Prodr. V (1836) 204.

Capitula cylindrical, solitary. Involucral bracts in one row, linear-
lanceolate. Florets heterogamous; central florets bisexual, fertile, with
tubular, 5-lobed corolla; peripheral florets pistillate, with longer corolla,
their stigmas of same shape as in bisexual florets. Achenes cylindrical,
somewhat ribbed; pappus narrowed, its bristles simple. Perennial herbs
with reniform toothed or lacerate wintering leaves.

Of the three species found in Western Europe, we have one.

1. H. alpina (L.) Cass. in Bull. Soc. Philom. (1816) 198; DC.
Prodr. V, 205; Szaf. Kulcz. Pawl. Roslin. Polsk. 632; Vizn. Rosl.
USSR, 552; Hegi, Ill. Fl. VI, 696, fig. 410.—Tussilago alpina L. Sp.
pl. (1753) 865.

Perennial. Rhizome branched, creeping, slender, 1.5—2.0 mm thick.
Basal leaves petiolate, 2-4 cm long, roundish-reniform, 1.5-5.0 cm
wide, sinuate-toothed, teeth broadly deltoid, with short apex, petioles
as well as leaves, particularly along veins, covered with thick, ochreous
hairs. Stems 10-—20(40) cm high, densely hairy, particularly younger
stems, with similar hairs. Cauline leaves few, the lower amplexicaul,
broadly ovate, upper narrowly lanceolate. Capitula solitary; involucral
bracts in single row, linear-lanceolate; florets lilac. Achenes 5 mm
long, somewhat ribbed, half as long as pappus. Flowering April to
May; fruiting June.

'Treatment by L.A. Kuprianova.
7From the Greek words. omos—equal, identical, and gyne—wife, women, the
stigma of the pistillate (female) florets is of the same shape as in the bisexual florets.

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624

Alpine and subalpine mountain zones and in bordering forests.—
European Part: Upper Dniester. General distribution: mountains of
Central and Southern Europe. Described from Western Europe. Type
in London.

GENUS 1558. Arnica L.!:?

L. Sp. pl. (1753) 884; Maguire, Monogr. gen. Arn. in Brittonia,
IV, 3 (1943).

Capitula solitary or few; involucre 2-rowed, with herbaceous, more
or less equal involucral bracts, very rarely one-rowed, receptacle
convex, covered with hairs or cilia. Peripheral florets ligulate, pistillate,
with yellow or orange ligules; other florets tubular, bisexual, yellow or
orange, often pale in lower part, with 3 teeth; anthers almost as long
as filaments, mostly yellow, less often dark purple; style with slender,
stigma distinctly exserted from corolla, grooved inside, papillate outside,
apically extending almost into raceme. Pappus consisting of many
bristles or short- to long-barbate, less often almost pinnate, white,
slightly pinkish, or dull, as long as tubular corolla or slightly longer.
Achenes linear-terete, mostly slightly narrowed toward both ends, with
longitudinal ribs, basally always with a white ring, glabrous, setose-
hairy or even glandular. Plants herbaceous, with opposite leaves, less
often only upper pair obliquely opposite or alternate. In all, the genus
includes 12 species.

Type of genus: A. montana L.

1. Anthers purple or blackish-violet (when dry); pappus sordid or straw-

colored (Subgenus III. Andro-purpurea Maguire) ............::1000+00 2:
+ Anthers yellow; pappus white, somewhat pinkish, less often straw-
colored x.y sana... BRN. BS. dees ORI. Aa 4.

2. Corolla tube glabrous or only with occasional, very sparse hairs,
abruptly broadened into limb almost as long as tube .............::2000000
Nea AAR Bs thee atl peed tS ote 7. A. unalaschcensis Less.
+ Corolla gradually attenuate into tube, shorter than limb, with scattered
ordenser;‘upright halts, ¢.0\.0) cat ui eee ie i 3.
3. Plants tall, up to 100 cm high, glabrous, hairy only at stem apex
below capitula, with 12-20 pairs of somewhat thick leaves; capitula
always erect, pappus. straw-yellow x... ..2..... 00a. Aan).
ee ee ne 8. A. sachalinensis (Rgl.) A. Gray

'Treatment by M.M. Iljin.
*From the Greek word arnicos—sheepskin, or arnos—sheep.

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625

+ Plants low, with more or less hairy stems, lacking leaves or only
with 1-4 pairs of cauline leaves; capitula always drooping; pappus
Gall agate nel dheaphaticca tic hiie. ceesiscsienns 6. A. lessingii Green
4. _ Basal leaves wide, ovate, oval, or oblong-oval, mostly subobtuse,
sessile or subsessile; hairs in upper part of stem simple and glandular;
pappus straw-colored. Plants of forest regions in western European
part of USSR (Subgenus II. Montana Maguire) ...0..........::ccccsceeseeeeees
SOG ea YeRE sR OD GA plate tleiaeh ge Lists chewieene 8535 5. A. montana L.
+ Basal leaves narrower, subacute or acuminate, distinctly narrowed
into petioles; hairs in upper part of stem similar or only simple;
pappus white or with scarcely noticeable pinkish shade. Arctic and
subarctic plants of Siberia and far north of European part of USSR

(Suligeovn sl wAnction Maguire) nseiiiadanancdaamiudelencwe B:
Site Plants with onlysimaple thang singcivces istics tvccatevisthisedledentensth 6.
+ Plants with simple and glandular hairs 2.0.0.0... .ccccssceseesseeenees ts
6. Basal leaves narrow, lanceolate or linear-lanceolate, entire,

acuminate, sparsely hairy beneath. Kola Peninsula ......................
Paha der GREE. KGUK NE AWA opie. 1. A. alpina (L.) Olin and Ladau
+ Basal leaves lobed, oval or oblong-oval, obtuse or subacute, remotely
toothed in upper part, glabrous or subglabrous beneath, usually
glaucescent. Plants of Chukotka Peninsula ...........0...cceseeeseeeeees
pibpartessathies pike each batisat ses atc. ans 4. A. frigida C.A.M. ex Iljin
7. Stem densely tomentose below capitulum covering most of the
glandular hairs; corolla of ligulate florets usually dark yellow. Plant
af Okhotsk: Distniet sc .ielwssits..sis..ccsacesansds 3. A. intermedia Turcz.
+ Glandular hairs at top of stem very visible under hand lens; corolla
of ligulate florets light yellow. Plant of far northeast of European
part of USSR and Siberia .................. 2. A. iljinii (Maguire) Iljin.

Subgenus 1. Arctica Maguire. Monograph. gen. Arn. (1943) 406.—
Anthers yellow; pappus white; leaves narrow. Plants of the Arctic.

1. A. alpina (L.) Olin and Ladau, Dissert. de Arnica, Upsaliae
(1799) 11; Murr. in Sv. Vet.-akad. Nandl. 337; Iljin in Tr. Bot. Muz.
Akad. Nauk SSSR, XIX, 112.—A. montana alpina L. Sp. pl. (1753)
884; Fedtsch. and Fler. Fl. Evrop. Ross. 988 p. p.—A. alpina (L.) Olin
subsp. genuina Maguire Monogra. gen. Arn. (1943) 408.—A.
angustifolia DC. Prodr. VI (1937) 317 p. p. non Vahl.—Ic.: Iljin, op.
cit.; Maguire, loc. cit. f. 26; fig. 27; Svensk. Bot. X, t. 699.—Exs.: Pl.
Fin. exs. No. 1394.

Perennial. Plant 10—25(30) cm high, with simple, usually solitary
stem, hairy almost from base, more densely near capitula, with simple
hairs (glandular hairs absent). Cauline leaves 1-3(4) pairs, broadly to

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626

narrowly lanceolate, entire, acuminate, sessile; basal leaves mostly
smaller, sometimes only acute or even almost subobtuse, and more
tapering toward base in short winged petiole; all leaves on both sides
sparsely covered with long hairs, 5-10 cm long and 5-10 cm wide,
with 3-5 parallel veins. Capitulum solitary, campanulate, 1.5—2.0 cm
long and 2.5-3.0 cm in dia when mature. Involucre and stem apex
densely whitish-pubescent-hairy from simple hairs, with occasional,
indistinct, very short, stalked glands under hair cover; involucral bracts
12-20, lanceolate, acuminate, dark green, usually sordid purple in upper
part, 10-13 mm long and 1.5-2.5 mm wide. Florets pale-yellow,
ligulate, 10-15, with oblong-linear ligules, 4-7 mm wide, 15-30 mm
long, mostly with 7-10 dark yellow veins, with occasional sparse hairs,
often glabrous above, subglabrous on inner (ventral) side, apically
with 3—4(5), oblong-deltoid teeth; tubular florets 6—9 mm long, almost
as long as pappus, densely hairy in lower part like base of ligulate.
Achenes 4—7 mm long, oblong, densely covered with long, upright,
simple hairs. Flowering July to August; fruiting August to September.

Meadows of alpine and subalpine mountain zones.—Arctic: Arctic
Europe; European Part: Karelia-Lapland (only on Kola Peninsula).
General distribution: Scandinavia (Lapland). Type in London.

Note. In the herbarium of the Botanical Institute of the Academy
of Sciences of the USSR, there is an indisputable specimen of this
species from the Altai, from the vicinity of Riddersk. It can be pre-
sumed that there is some error in the labeling, particularly since this
report has not been confirmed. It would be more likely to expect to
find the north Siberian species A. iljinii (Maguire) Iljin in the Altai.

Economic Importance. Chirkh (1917) mentions that this species
is also sometimes referred to as A. montana L. The value of this species
as a medicinal plant is reported by Hocking (1944) also. It is possible that
both the rest of the arnica species similar to A. alpina (L.) Olin and Ladau
and of the other subgenera may have similar medicinal properties.

2. A. iljinii (Maguire) Iljin comb. nova.—A. alpina subsp. iljinii
Maguire Monogr. gen. Arn. (1943) 411.—A. angustifolia Iljin in Tr.
Bot. Muz. Akad. Nauk SSSR, XIX (1926) 110, non Vahl; Kryl. FI.
Zap. Sib. XI, 2833, non Vahl.—A. montana f. stenophylla Ldb. FI.
Ross. II (1844) 622 p. p. (excl. specim. Alach-Jun.).—A. alpina auct.
siber. non Olin; Fedtsch. and Fler. Fl. Evrop. Ross. 988 p. p.—lIc.:
Maguire, loc. cit., fig. 26, f. 27; Iljin, loc. cit. X, Fig. 2.

Perennial. Plant 10-50 cm high, with slender, creeping, 1-3 mm-
thick brownish rhizome, densely covered with appressed, flat, entire or
strongly divided scales, densely covered under scales in its upper part
with long, whitish hairs gradually decreasing downward and vanishing.

627

Stem solitary or few, erect or weakly ascending, at base often dirty
purple, more often simple, sometimes with short flowering branches or
even long branches, pubescent, with sparser simple hairs and subsessile
glands below, more densely hairy below capitulum and at base of
involucre, simple, long, somewhat entangled hairs mixed with very
numerous, multi-cellular glandular hairs that are shorter than simple
hairs. Basal leaves few pairs, lowermost leaves shorter, often withering
before flowering, others longer, oblong-oval to broadly or narrowly
lanceolate, narrowed toward base into broad petiole, like cauline leaves
entire, less often with occasional teeth and short cilia along margin,
acuminate, with 3—5(7) veins, on both sides with scattered simple hairs
and denser, subsessile glands, less numerous above, 5—20 cm long
including petiole, 0.5—3.0 cm wide; cauline leaves 1-3 pairs, reduced
upward, much narrower, lower leaves with short winged petiole, others
narrowed toward base but sessile, borne in lower part of stem. Capitula
solitary or few at apices of stem and branches, 2-6 cm in dia at
flowering, involucre campanulate, 10-15 mm long; involucral bracts
lanceolate, acuminate, usually reddish-violet in upper part or entirely,
(12)14-18(20), with simple hairs and sessile (or subsessile) glands.
Ligulate florets light yellow; ligule, 12-25 mm long and 3-10 mm
wide, 7-9 veined, with 3 teeth at apex and even 3-fid, glabrous on
both sides with short tube covered with simple upright hairs; tubular
florets (6)7—9(10) mm long, yellow, almost as long as pappus, with
limb covered with simple upright hairs, sometimes extending into base
of expanded part of tube, glabrous in remaining part with short hairy
teeth. Pappus white, serrate-barbed; achenes linear, 4-6 mm long, dark
gray, very finely and closely, longitudinally ribbed, covered with simple
upright hairs. Flowering second half of June to first half of August;
fruiting second half of July to September.

Tundra and forest-tundra as well as alpine region, northern taiga
region, on stony slopes, rocks, clay and limestone outcrops, gravelly
and sandy shoals of rivers, in grassy and lichen-moss tundras, meadows,
forest glades willow thickets, open taiga forests, etc.—Arctic: Arctic
Europe (except Kola Peninsula), Novaya Zemlya, Arctic Siberia,
Anadyr; European part: Dvina-Pechora (north); Western Siberia: Ob
River Area (north), Yenisei (north); Eastern Siberia: Lena-Kolyma
(north). Endemic. Described from Ust-Yenisei Port (69°39' N. Lat.).
Type in Leningrad.

3. A. intermedia Turcz. in Bull. Soc. Nat. Mosc. XXIV (1851)
203; Iljin in Tr. Bot. Muz. Akad. Nauk SSSR, XIX, 111.—A. montana
B. stenophylla Ldb. Fl. Ross. II (1844) 622, p. p.—A. angustifolia DC.
(non Vahl) Prodr. VI, 317 p. p., A. alpina subsp. intermedia (Turcz.)

660

628

Maguire, Monogr. gen. Arn. (1943).—Ic.: Iljin, loc. cit. Plate X, la
and b. ;

Perennial. Plant 10-30 cm high, stem mostly simple solitary or
few, erect or weakly ascending, hairy from base, particularly densely
hairy-villous below capitulum with long, divergent, simple hairs and
shorter glands, more or less sparse, concealed under densely woolly
cover at stem apex and base of involucre, also dense and long-villous.
Basal leaves few, narrowly lanceolate to oblong-lanceolate, narrowed
toward base, acuminate or acute, with 3—7 veins, on both sides with
long, sparse or dense, semiappressed hairs, sometimes with very short,
remote teeth; cauline leaves sessile, usually narrower, 1—2(3) pairs,
often with undeveloped short pedunculate capitula in their axils.
Capitula 4—5 cm in dia when open. Involucre 12—15 mm long; involucral
bracts (13)15-20(23), mostly oval-lanceolate, sub-acute, often with
sordid purple shade, with dense entangled hairs, in lower part,
concealing glands, sparsely glandular in upper half with sessile glands,
long-ciliate-hairy only along margin, but apically with a tuft of hairs,
sometimes with similar hairs, somewhat below and on dorsal side among
glands, glabrous on inner side. Ray florets usually dark yellow when
dry, ligulate, 15-18, ligule mostly 15—20 mm long and 3-6 mm wide,
with 7-11 longitudinal veins, apically with 3 teeth, less often 3-fid,
glabrous on inner side, sparsely hairy in lower part on outer side,
abruptly terminating into short tube, mostly 3-4 mm long, covered
with obliquely upright hairs; tubular florets about 6 mm long, with
erect hairs in lower half, glabrous above. Pappus serrate or scarcely
serrate-barbate, 8-9 mm long, longer than tubular florets and tube of
ligulate florets; achenes oblong, 4.5-5.0 mm long, brownish, densely
covered with semiappressed and obliquely upright, simple, long hairs.
Flowering July; fruiting August(?).

Eastern Siberia: Lena-Kolyma (Aldan, Allakh-Yun); Far East:
Okhotsk. Endemic. Described from Allakh-Yun. Type in Leningrad.

4. A. frigida C.A.M. ex Iljin in Tr. Bot. Muz. Akad. Nauk SSSR,
XIX (1926) 112; Kom. Vved. v Izuch. Rast. Yakutii, 1, 162.—A. alpina
Less. in Linnaea, VI (1831), 235 (non Olin and Ladau); Herder in
Bull. Soc. Nat. Mosc. XL, 1, 423; Ldb. Fl. Ross. II, 623.—A. lessingii
Trautv. ex Iljin, op. cit. 112, non Greene.—A. angustifolia Vahl. B.
lessingii Torr. and Gray, Fl. North Amer. II (1843) 449.—A. louseana
Farr. subsp. frigida (C.A.M. ex Ijin) Maguire in Madrono, VI (1942)
153 and in Monogr. Gen. Arn. (1943) 417.—Ic.: Iljin, loc. cit., Plate
XI, Fig. 3a and 3b.

Perennial. Plant 5-30 cm high, with short, glabrous rhizome,
densely covered with brownish remnants of leaves. Stems single or

=

jj

Plate XXXII.
f tubular floret.

Habit, tubular floret, corolla o

A. unalaschcensis Less.; 3 —A. sachalinensis

1—A. lessingii Greene; 2 —

Gray.

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630

few, simple, very rarely with 1-2 branches, usually reddish-purple at
base or in lower half, densely hairy throughout, particularly densely
below capitula, with long, simple, multi-cellular hairs (uppermost cell
filiform), lacking glandular hairs and simple glands, with 1-3 pairs of
leaves, less often even leafless. Leaves of basal rosette lobate, oblong-
oval or oval, mostly short-petiolate, less often petioles longer but always
shorter than lamina, lamina obtuse or acute, with remote teeth mainly
in upper part, like petioles ciliate-hairy, with sparse or occasional hairs
above, mostly glaucescent and glabrous beneath or sometimes with
occasional hairs, particularly on midrib; cauline leaves narrower,
lanceolate or linear, entire, sessile, usually more densely pubescent;
uppermost leaves often obliquely opposite or even alternate. Capitula
solitary, very rarely 2-3 at apices of branches, (2)3.5—5.0 cm in dia.
Involucre (10)12—14 mm long, densely pubescent in lower part, usually
very sparsely above, purple entirely or only along margin and midrib;
involucral bracts linear or lanceolate, acuminate, (2)2.5—3.0 mm wide,
usually 12-16. Ligulate florets (9)10—15, light yellow, with (5)7—10
veins, with 3 teeth or almost 3-fid, ligule (15)17-23 mm and 3—5 mm
long limb, usually densely hairy with simple, long hairs on short
unicellular base, ligule 3-5 mm wide; tubular florets 6-9 mm long,
tube shorter than limb and densely covered with erect hairs as in ligulate
florets, with teeth often barbate and short-hairy. Pappus as long as
tubular florets or slightly longer than it and limb of ligulate florets,
silvery white, usually with scarcely noticeable pinkish shade, short-
barbate; achenes linear, 4-6 mm long, blackish-gray or dark brown,
with sparse, often occasional, obliquely upright hairs, much longer in
upper part, often becoming glabrous. Flowering July to August; fruiting
July to September (?).

In dry lichen, meadow, rubbly and stony tundras as well as on
gravel beds, rocks, sand bars in river valleys, coastal bluffs and
along the coast of Arctic Ocean in tundra zone.—Arctic: Arctic
Siberia, Chukotka, Anadyr; Eastern Siberia: Lena-Kolyma. General
distribution: North America (islands in Bering Strait and Alaska).
Described from Gulf of Northern Laurentia and Gulf of Eschscholtz.
Type in Leningrad.

Note. The species was first established under this name by
Trautvetter, but he did not publish a description. Iljin, in Tr. Bot. Muz.
Akad. Nauk SSSR, XIX, 1926, pp. 112 and 116-117.

Subgenus 2. Montana Maguire Monogr. Gen. Arn. (1943) 483.—
Anthers yellow; pappus straw-yellow; leaves in rosette wide. Plants
mostly of forest and mountain-forest zones.

664

631

5. A. montana L. Sp. pl. (1753) 884; DC. Prodr. VI, 317; Maguire,
op. cit. 485; Iljin in Tr. Bot. Muz. Akad. Nauk SSSR, XIX, 110; Ldb.
Fl. Ross. II, 622 (excl. specim. Ekaterinenburgensis and B. stenophylla;
Fedtsch. and Fler. Fl. Evrop. Ross. 487.—A. plantaginisfolio Gilib. FI.
Lith. III (1782) 210.—A. alpina Willd. ex Steud. Nom. ed. 1 (1821)
70 non Olin.—A. helvetica G. Don. ex Loud hort. Brit. (1830) 351.—
Doronicum montanum Lam. Encycl. meth. II (1790) 312.—D. arnica
Desf. Cat. h. paris. ed. 1 (1804) 101.—D. oppositifolium Lam. op. cit.
p. 312.—Ic.: Maguire, loc. cit. fig. 94, No. 95; Iljin op. cit. X, 3;
Rchb. Ic. Fl. Germ. XVI, t. 958; Hegi, Ill. Fl. VI, 2, f. 414-418; Kom.
Lekarstv. Rast. SSSR, IV; Zemlinskii, Lekarstv. Rast. 57.—Exs.: FI.
Polon. exs. No. 178; Fl. exs. reipubl. Bohem.-Sloven. No. 287; FI.
Hungar. exs. No. 589; Hayek, FI. stir. exs. Nos. 1270, 1271.

Perennial. Plant 15-80 cm high, simple or branched. Rhizome
short; stem erect, with simple and glandular hairs, particularly dense
in upper part, with rosette of leaves at base and 1-2, less often 3 pairs
of cauline leaves. Leaves of basal rosette broad, oval or oblong-oval,
entire, subobtuse, with prominent lateral veins, sparsely hairy above,
mostly glabrous beneath or only along vein weakly hairy, short-petiolate
or subsessile; cauline leaves oblong or lanceolate, less often oblong-
oval (lower pair), in upper half of stem, particularly on branches, with
only few alternate, linear, acuminate leaves. Capitula solitary at apices
of stem and branches, hemispherical, 1(2)—3(5) cm in dia. Involucre of
22(26) lanceolate, subacute involucral bracts with anthocyanin color,
14(12)-17 mm long and 2-5 mm wide, with simple, longish and often
glandular hairs on out side, along margin and at apices hairy-ciliate.
Ligulate florets yellow, usually 11-15, ligules with 3 (less often 2-4)
teeth, tube usually long, pubescent, almost as long as pappus; tubular
florets paler, short-toothed, also with pubescent tube as long as pappus
or longer, anthers yellow. Achenes 6-10 mm long, narrowed toward
base, with erect, simple, serrate-barbed hairs. Flowering June to August
(at higher altitudes July to September); fruiting June to September.

Mainly in pine groves and pine-birch forests, in forest glades,
clearings and dry meadows, high-mountain meadows.—European part:
Baltic Region, Upper Dnieper, Upper Dniester. General distribution:
Scandinavia, Central Europe, Atlantic Europe, mountains of northern
Italy and northern Balkans. Described from Alps and northern Europe.
Type in London.

Economic Importance. Not much used in modern allopathic medi-
cine; however, it finds a place in the official pharmacopoeia of the
USSR. The flowering capitula and a tincture of them are used for
medicinal purposes. In other countries, only an extract of the flowers
is used, or the flowers are used as a plaster; sometimes the rhizomes

665

632

are used for obtaining tinctures and extracts. It is used internally to
stimulate cardiac activity and externally as a lotion and liniment. In
homeopathic medicine, it is used as a cardiac tonic and as a preparation
to stop bleeding in various traumas. Arnica is widely used in folk
medicine against fevers, diarrhea, paralysis, epilepsy and other nervous
disorders, internal hemorrhages, as a diaphoretic, and in the form of
poultices for contusions. The rhizome contains a significant quantity
of tannins; according to M. Rylov (1894), up to 32%. According to
V.K. Varlikh (1901) and N. Ya. Demyanov, V.I. Nilov, and V.V.
Williams (1933), moreover, it contains a bitter substance—arnicin,
resins, wax, gum, as well as 0.5 to 1.5% essential oils on a dry weight
basis. According to the latter authors, the physical properties of these
oils are as follows: 1) specific gravity, calculated in relation to the
density of water at 15°C=0.982 to 1.000; 2) refractive index of yellow
light at 20°C=1.507 to 1.508; 3) pH=4 to 10; 4) ester number=60 to
100; 5) dissolves in 7-12 volumes of 80% alcohol. This oil contains:
hexycapronate, caproic, formic and isobutyric acids, phlorol
(ethylphenol), phloraisobutyric ester, thymohydroxynonomethy! ester
and phloromethy! ester. According to V.K. Varlikh, the dry flowers
contain 0.1-0.9% essential oil, fats, wax, resins, yellow pigments
(apparently carotenoids), tannins, proteins, gums, mineral salts and
arnicin. According to the above-cited three authors, fresh flowers
contain 0.04—0.07% essential oil with the following physical constants:
1) specific gravity in relation to the density of water at 15°C=0.8905
to 0.9029; 2) melting point =20—30°C; 3) acid number=62.6 to 127; 4)
ester number=22.7 to 32.2; 5) does not dissolve well in alcohol. This
oil contains lauric and palmitic acids and palmitine. According to
Womer, the rhizome contains up to 9%, and according to Dragendorf
9.7%, inulin, but the flowers, in addition to the above substances, also
contain dextrose, crystalline phytosterol-arnidiol or arnisteral and an
amorphous substance—arnicin. According to M.M. Glukhov, this
species of arnica is a good honey-producing plant.

Subgenus 3. Andropurpurea Maguire, Monogr. Gen. Arn. (1943)
486.—Anthers purple or blackish-violet (when dry); pappus sordid or
straw-yellow. Plants of the forest and arctic zones.

6. A. lessingii Greene in Pittonia, IV (1900) 167; Iljin in Tr. Bot.
Muz. Akad. Nauk SSSR, XIX, 114; Maguire, Monogra. Gen. Arn.
(1943) 486; Kom. Fl. Kamch. III, 160; Hulten, Fl. of Kamtsch. IV,
192.—Ic.: Macquire, loc. cit. fig. 90, fig. 91; Hultén, loc. cit. t. 6, f,
e; Iljin, loc. cit. Plate XI, Fig. 4.

666

633

Perennial. Plant 6-40 cm high, with slender, reddish-brown, weakly
branched or simple, glabrous, rhizome covered with brown leafy scales
on remote nodes, crowded only at apex. Stems solitary or few, mostly
erect, in basal part more or less ascending, terete, leafless or with 1-
3(4) pairs of leaves, covered with multi-cellular, weakly reddish hairs,
somewhat longer at places at cellular septa[nodes?], in upper part mostly
long-pointed, sparse in lower part of stem, denser above. Basal leaves
oval or oblong-oval, lobate, less often only oblong, 3—7(10) cm long
and 1—2 cm wide, narrowed into very short petiole, entire or less often
with remote teeth, particularly in upper part of lamina; cauline leaves
mostly narrowed into narrowly lanceolate, often somewhat obliquely
opposite, mostly uppermost leaves toward base also shortly narrowed
and extending into short sheath common with opposite leaf; all leaves
pubescent-ciliate along margin, glabrous on both sides, less often with
occasional hairs, distinctly paler beneath, sometimes hairy along midrib.
Capitula solitary, drooping or semi-drooping, with ligulate florets 4.0—
5.5(6.0) cm in dia. Involucre broadly campanulate, reddish-purple in
lower part, green in upper; involucral bracts lanceolate or less often
oblong-ovate, (10)12—15(17), with distinct, but not prominent, midrib
and indistinct lateral veins, more or less scattered hairs on outer side
as on stem, ciliate, glabrous on inner side. Ligulate florets pale yellow
or dark colored, (8)9-13, with 5—9-veined ligules, (14)16.5—22.0 mm
long and 2.5—5.0(8.0) mm wide, glabrous and in upper part with 3
glabrous teeth, at base narrowed into 3.5—5.0 mm-long tube covered
with sparse and obliquely upright, longish hairs; tubular florets 6—8(9)
mm long, gradually narrowed into distinctly shorter tube, also with
hairs as in ligulate florets, above with teeth covered with papillae or
occasional short hairs; anthers blackish-violet, exserted from tube.
Pappus short-barbed, exceeding corolla and almost as long as stigma
of tubular florets and much longer than limb of ligulate florets,
somewhat sordid; achenes linear, 5-6 mm long, with numerous scabrous
ribs, glabrous, grayish or brownish. Flowering July; fruiting August.
(Plate XXXII, Fig. 1).

Subalpine and alpine mountain zones.—Far East: Kamchatka.
General distribution: North America (Alaska, Aleutian Islands).
Described from Alaska and adjacent islands without their exact
designation. Type in Quebec in Notre Dame University.

7. A. unalaschcensis Less. in Linnaea, VI (1831) 238; DC. Prodr.
VI, 317; Ldb. Fl. Ross. II, 623; Fedtsch. Komand. Ostr. 55; Iljin in Tr.
Bot. Muz. Akad. Nauk SSSR, XIX, 117; Hultén, Fl. of Kamtsch. IV,
193; Maguire, Monogr. Gen. Arn. 489.—A. obtusifolia Less. in Linnaea,
VI (1831) 237; Ldb. op. cit. 623.—A. langsdorffiana Fedtsch. in Herb.

667

634

Herder in Bull. Soc. Nat. Mosc. XL (1867) 424.—Ic.: Maguire, op.
cit. fig. 96, fig. 97; Iljin, op. cit. Plate XI, Fig. 1.

Perennial. Plant 8-40 cm high, with thick, brown, glabrous rhizome
tapered downward, densely covered with remnants of old leaves; stem
simple, solitary, sulcate-ribbed, at base weakly ascending, somewhat
thick, covered from very base with long, simple, articulate, often
longitudinally appressed, multicellular hairs particularly densely above.
Basal leaves oval or oblong-oval, less often oblong, tapered into broad,
flat and amplexicaul petiole, obtuse or slightly subacute, with remote
teeth; cauline leaves 2-3, less often up to 5 pairs, uppermost pair
obliquely opposite or even alternate, oblong-ovate or oblong, sessile,
narrowed toward base, in very rare cases long-petiolate (f. petiolata
Iljin) sparsely serrate; all leaves sparsely hairy on both sides, densely
ciliate-hairy on margin, with 3, less often 5 veins, with arcuate lateral
veins, 5(3)—12 cm long and 1-4 cm wide. Capitula solitary, on peduncle
3-15 cm long, with ligulate florets 3—6 cm in dia. Involucre 10-17 mm
long, of 16—27 lanceolate, often narrowly lanceolate involucral bracts
callously thickened above, or even acuminate and recurved, with sparse
or denser hairs on outside but less dense than at tip of peduncle with
same type of hairs, less often also with sessile or subsessile glands.
Ligulate florets yellow, 15-25, with glabrous ligules, with 3-4 teeth or
lobes, 12-19 mm long and 3-7 mm wide, with 7-12 darker veins,
terminating into short, entirely glabrous, 3.0-4.5 mm long tube; corolla
of tubular florets 5-7 mm long, with glabrous tube and broadly
campanulate limb, almost as long as tube, their teeth with occasional
short hairs and papillae on outside at tip, stamen tube blackish-violet
(when dry), scarcely exserted from limb; pappus sordid-yellow, barbed,
as long as tubular florets but longer than tube of ligulate florets; achenes
3.0-4.5 mm long, linear, grayish, covered with sparse, obliquely upright
hairs and sometimes also with glands, sometimes subglabrous. Flowering
July to August; fruiting September. (Plate XXXII, Fig. 2).

Grassy alpine balds, tallgrass slopes of [mud] volcanic cones,
coastal lowlands.—Far East: Kamchatka (south; Commander Islands),
Sakhalin (Kuril Islands). General distribution: Japan, North America
(Aleutian Islands, Unalaska). Described from Unalaska Islands. Type
in Leningrad.

Note. Maguire (op. cit.). reports that in the tubular florets the
corolla tube can be glabrous or pilose (p. 489), and, on this basis, he
treats A. tschonoskyi I|jin—the species described by me—as a synonym
of A. unalaschcensis Less. I cannot agree with this. While processing
material for the Flora of the USSR, 1 re-examined all the specimens of
this species from the Commander Islands, the south of Kamchatka and
the Kuril Islands, and despite a most careful examination under a

668

635

binocular microscope did not find any hairs on the tube of the tubular
[or] ligulate florets; they were always entirely glabrous or sometimes
with only rudiments of occasional hairs. Therefore, A. tschonoskyi,
with a strongly pilose corolla tube of the tubular florets is a separate
species, growing in Japan. Moreover, typical A. unalaschcensis Less.
is not found at all in Japan, even among the plants with glabrous
corolla tubes. The question of the taxonomic status of the Japanese
plants can be resolved only by studying much more material. Maguire’s
error lies in the fact that among the specimens determined by him from
our herbarium, one specimen, undoubtedly belonging to A. mollis Hook.,
was determined by him as A. unalaschcensis Less. (Tilesius’ collections
of E. Sibiria N. am.); since in A. mollis Hook. the corolla tube of the
tubular florets is strongly pilose, the source of the inaccuracy in the
description of A. unalaschcensis lies here. We must consider this
determination as an inadvertent slip of pen, because these two species
even belong to different subgenera.

The forms described by Herder (Herder in Byull. Mosk Obshch.
Prir. 40 (1867) p. 423), minor seu genuina and B£. major (pedalis) as
well as f. scaposa B. Fedtsch. (Fl. Il. Comm. 1906, 55), are not taxo-
nomically significant.

8. A. sachalinensis (Rgl.) A. Gray in Proc. Amer. Acad. XIX
(1884) 55; Kom. Opred. Rast. Dalnevost. Kr. II, 1044; Ijin in Tr. Bot.
Muz. Akad. Nauk SSSR, XIX, 117; Maguire, Monogr. Gen. Arn. 491.—
A. chamissonis F. Schmidt. Reise (1868) 191.—A. chamissonia
sachalinensis Rgl. Suppl. ind. Sachal. (1874) 151.—Ic.: Maguire, op.
cit. fig. 98, 99; Iljin, op. cit. Plate XI, Fig. 2.

Perennial. Plant up to 100 cm high, with long branched rhizomes;
stems somewhat thick, ribbed-sulcate, simple, branched only at apex,
glabrous, only below capitula with variously long, moniliform hairs,
setose at tip. Leaves numerous, usually 12—20 pairs, 5—15 cm long and
1.2-4.0 cm wide, distant in lower part of stem, rather crowded in
upper half, oblong or oblong-oval, sessile, narrowed toward base and
merged with opposite leaf into short sheath, acuminate or acute, with
remote sharp teeth, coarse, almost coriaceous, with distinct reticulate-
arcuate veins, particularly beneath, on both sides glabrous, very rarely
upper leaves with scarcely noticeable and very sparse short hairs, usually
turning black when dry, mainly above; rosette not developed; lower
cauline leaves withering before anthesis. Capitula solitary at apices of
stems and branches, mostly in corymbose inflorescence, 20-35 mm in
dia. Involucre 14-20 mm long; involucral bracts 8-15, oval or
lanceolate, leafy, 4-8 mm wide, with distinct longitudinal veins, mostly
at base with articulate, multi-cellular hairs like stem apex, in upper

669

636

part with sparse scattered hairs and glands or mostly glabrous, ligulate
florets light yellow, 20-35 mm long, 10-18, with 3 teeth, with 7-
11(13) veins, ligule 4.5-9.0 mm wide, with shorter tube having long,
multi-cellular, erect hairs, terminating into erect hairs or thin long-
pointed tip; ligule glabrous, stamen tube blackish-violet (when dry);
tubular florets numerous, 7.5-12.0 mm long, gradually elongated into
similarly hairy tube, much shorter than limb. Pappus stramineous, longer
than tubular florets and tube of ligulate florets, short-barbed; achenes
linear-terete, 6.5-9.0 mm long, sulcate-ribbed, with sparse, long,
obliquely upright hairs, often glabrescent. Flowering July to August;
fruiting August to September. (Plate XXXII, Fig. 3).

Mixed forests, shrub thickets, in mountain regions, along
riverbanks.—Far East: Uda River Area, Sakhalin. Endemic. Described
from Sakhalin Island. Type in Leningrad.

GENUS 1559. Doronicum L.!:?

L. Sp. pl. (1753) 885.—Aronicum Neck. Elem. I (1790) 27.—
Grammarthron Cass. in Bull. Soc. Philom. (1817) 32.

Capitula semiglobose or broadly campanulate, mostly solitary or
sometimes 2—6(8), in corymbose inflorescence. Involucral bracts in 2—
3 rows; outer bracts lanceolate, oblong-lanceolate, or lanceolate-linear,
herbaceous, sparsely pubescent or in lower part glandular-hairy; inner
bracts linear, linear-lanceolate, membranous, smooth, ciliate or in upper
part, glandular-hairy, all bracts long-acuminate. Receptacle more or
less convex, glabrous or hairy. Disk florets tubular, bisexual, yellow,
in several whorls. Peripheral florets pistillate, ligulate, yellow, in single
whorl; anthers at base undivided or auriculate-sagittate; stigma lobes
of central florets apically truncate, short-tufted, in peripheral florets
short, obtuse. Achenes oblong or oblong-turbinate, obtuse, with 10
equal, longitudinal ribs, mature achenes brown or dark brown, glabrous
or covered with appressed, erect, white, straight hairs; pappus of ligulate
florets (sometimes undeveloped) of numerous, one-rowed (in peripheral
achenes) or many-rowed (in central achenes), white or rusty, sparsely
barbed bristles.

Herbaceous perennials with basal and alternate cauline amplexicaul
leaves.

The genus includes about 36 species distributed in the mountains
of temperate regions of Asia and Europe, where its species are found
up to 3,500 m; one species is found in northern Africa. The large,

‘Treatment by S.G. Gorschkova.
?Name of Arabic origin.

670

637

beautiful, golden-yellow flowers in the capitula of Doronicum and the
easy cultivation are fostering wide distribution of some of the species
in gardens and parks.

Ly

+

Ovaries and achenes of one type, smooth or sparsely pilose, all
Wrath rapa GRIN, LRA Re Li lasidse. arctan ceee ER otasdlcs #5
Ovaries and achenes of different types, peripheral ones glabrous
very rarely sparsely pilose, without pappus; central ones short-
appressed pilose, sometimes glabrous, with pappus ...............0+ 6.
Capitela. solitary sometimes 251.008 6052 AR RR dda 3
Capitula 2-4, on 10-28 cm-long peduncles with 1-3 small pales
BO WIGS, ATL I ode Us cd RRO Alda Clade 3. D. bargusinense Serg.
Leaves large, basal and lower cauline leaves 5.5—12.5(16.0) cm
long, 3.4—8.5(9.0) cm wide; basal leaves ovate or obovate-elliptical,
oblong or oval, less often roundish, with petiole 6-23 cm long;
ligulate florets with yellow corolla and glabrous ligules ............. 4.
Leaves 1.7-4.5 cm long, 2-3 cm wide; basal leaves of different
shape; ligulate florets with golden-yellow or pale yellow corolla,
ticirlipules sparsely, pilose at. base «........0iciecssscsdnseosavenvedecsediees 5:
Stem leafy up to tip, basally covered with small scale-like,
amplexicaul leaves; basal leaves ovate or obovate-elliptical, less
often roundish, petiole 6-19 cm long; lower cauline leaves ovate-
oblong, narrowed into winged petiole 2 cm long, other cauline
leaves sessile. Capitula with involucre 2.2—3.0(4.0) cm in dia; outer
involucral bracts oblong-lanceolate or lanceolate, inner linear-
lanceolate or linear; ligulate florets with corolla 1.8—2.4 cm long,
their tubes slightly shorter than pappus or half as long; achenes
glabrous or sometimes sparsely pilose ............. 1. D. altaicum Pall.
Stem in upper part without leaves, without scale-like leaves at base;
basal and lower cauline leaves broadly ovate or elliptical, with
petiole 13-23 cm long; middle leaves with petiole 3-5 cm long.
Involucre 2 cm in dia; outer involucral bracts lanceolate-linear,
sometimes divided up to a half; inner bracts linear; ligulate florets
with corolla 1.4—1.5 cm long, their tubes as long as pappus; achenes
pions 2. OAKS. ak Win STE A deren eve 2. D. schischkinii Serg.
Basal leaves elliptical, elliptical-roundish or oblong, 4.5 cm long
and 2 cm wide, with winged petiole, note; 4 mm long; cauline
leaves ovate-oblong or oblong; lower leaves narrowed into 4 mm
long, winged petiole, almost as long as lamina; ligulate florets with
golden-yellow Corolla ........ccscsseceseesenesesees 4. D. clusii (All.) Tausch.
Basal leaves ovate-roundish, 1.7—2.3 cm long, 2.2-3.0 mm wide;
with petiole 4-6(14) cm long more or less thickened at base; lower
cauline leaves ly: ate or cordate, amplexicaul, sessile; ligulate florets

671

638

~

oo 4

11.

with) pale: yellow! corolla» ::;.2.c.ntonnlinnetias eho. 2 ieee,
BH Wa eee Tals 5. D. carpaticum (Griseb. and Schenk) Nym.
Capitula numerous. /..02::.)..0308.. 4 ae ee eee i.
Capitula solitary: «....A0a0ed08. ne eet ae ee 9.
Rhizome tuberous, 0.8—1.3 cm thick; upper cauline leaves fiddle-
shaped and oblong-lanceolate, sinuate-toothed ...............:.008
gas ditha eee eel, Agen eas. Be ane 9. D. pardalianches L.
Rhizome slender; upper cauline leaves ovate or ovate-cordate .... 8.
Basal leaves 3.5 cm long, 4 cm wide, ovate-cordate, obtuse, with
4 cm-long petiole; cauline leaves decurrent, with cordate,
amplexicaul auricles. Involucre 2—3(3.5) cm in dia; involucral bracts
sparsely pilose; ligulate and tubular florets with glabrous corolla
nla sda can cuaaedee veRiRO Rea eae Bi unt Sate Biles epote 6. D. austriacum Jacq.
Basal leaves 9-19 cm long, 9.5—18.0(27.0) cm wide, broadly ovate-
cordate, with 22—35-cm long petiole; cauline leaves with amplexicaul
auricles. Involucre 3—4(4.5) cm in dia; involucral bracts densely
pubescent; corolla of ligulate florets with ligules short-pilose on
outside at base, corolla of tubular florets sparsely pubescent ..........
daanndlanisonnahabntvtngactietinndaai es MEME sg WORE? 8. D. macrophyllum Fisch.
Rhizome tuberous or with nodular thickenings ...............:::eeee 10.
Rhizomeslendeén): ects ccoiasleleenhuiecnsl eee see Ll.
Plant up to 80 cm high, glandular-pilose; rhizome tuberous; basal
leaves oblong-elliptical or oblong, 4-11 cm long, 2.0—3.8 cm wide;
cauline leaves lanceolate or broadly linear; corolla of ligulate florets
bright yellow, with glabrous ligules .............eeessseseseseeeesenneees
of, es RE SR cee Se) Sins: 10. D. longifolium Griseb. and Schenk.
Plant up to 47 cm high, glabrous; rhizome with nodular thickenings;
basal leaves broadly ovate-roundish or roundish-cordate, 3—5(6)
cm long and 3.5—6.0(6.5) cm wide; cauline leaves ovate or ovate-
elliptical; corolla of ligulate florets pale yellow, with ligules crisped-
pilose at: base: dee-\eew wal. wie et 7. D. orientale Hoffm.
Basal leaves elliptical or obovate-oblong, 2.5-7.0(10) cm long and
1.7-3.0(4.3) cm wide, with up to 16—20-cm long petiole; outer
involucral bracts lanceolate, 1.5—3.5(4.0) mm wide, inner bracts
narrowly lanceolate, 1.2-3.0 mm wide; corolla of ligulate florets
with glabrous tubes and basally pilose ligules ..............:sscessesseeeeeneees
«Sad: BERG Bch: Del tae | branes 11. D. oblongifolium DC.
Basal leaves obovate-spatulate or almost round, 3—11 cm long and
4.0-8.5 cm wide, with up to 15 cm-long winged petiole; outer
involucral bracts lanceolate-linear or lanceolate, 1.8—2.0(3.0) mm
wide; inner linear, 1.0—1.2(1.5) mm wide; corolla of ligulate florets
with tubes densely glandular-hairy on outer side and glabrous ligules
eg atas degieol Airs. wtibass 12. D. turkestanicum Cavill.

672

639

Section 1. Aronicum Neck. Elementa bot. I (1790) 26, sub gen.
—Grammarthron Cass. in Bull. Soc. Philom. (1847) 32, sub gen.—
Doronicum sect. Doronicum Tausch in Flora, XI (1828) 778.—
Doronicum sect. Aronicum Hoffm. in Pflzfam. IV, 5 (1892) 294.—
Sect. Doronicastrum Cavill. in Ann. Conserv. Jard. Bot. Généve,
XIII-XIV (1909-1911) 339 p. p.—Ovaries and achenes all similar
glabrous or sparsely pilose, with pappus of one-rowed bristles in
peripheral florets, and many-rowed bristles in central florets.

Series 1. Altaica Gorschk.—Basal leaves ovate, obovate-elliptical
or broadly ovate, 6-16 cm long, 3.0—-8.5 cm wide, with (13)20-—28 cm-
long petiole; ligulate florets with yellow corolla and glabrous ligules.

1. D. altaicum Pall. in Acta Ac. Petrob. VI, 2 (1783) pro Anno
1779, 271; Ldb. Fl. Alt. IV, 113; Cavill. in Ann. Cons. Jard. bot.
Généve, X, 191, XIII—XIV, 355; B. Fedtsch. Rast. Turk. 742; Kryl. Fl.
Zap. Sib. XI, 2830.—Aronicum altaicum DC. Prodr. VI (1837) 320;
Ldb. Fl. Ross. I, 624; Turcz. Fl. baic.-dahur. II, 85.—Arnica altaica
Turcz. in Bull. Soc. Nat. Mosc. I (1838) 95.—Ic.: Pail. op. cit. tab. 16;
Pall. Enum. Pl. b. 2.—Exs.: P. Smirn. Pl. Alt. exs. No. 92.

Perennial. Plant (8.5)10-80 cm high, glabrous; rhizome 0.3-0.8
cm thick, horizontal, sometimes oblique. Stem solitary, erect, simple,
densely glandular-pilose above, green or brownish, sometimes reddish-
violet, throughout leafy. Basal leaves (often withering) ovate or obovate-
elliptical, less often roundish, 5.5—10.0(16.0) cm long and 3.4—6.0(9.0)
cm wide, with 6—19(20) cm-long petiole; lower cauline leaves ovate-
oblong, 5—6(11) cm long and 4.0—4.7(6.0) cm wide, narrowed into 2
cm-long broad-winged petiole; basal and lower cauline leaves at base
of stem often reduced to small, amplexicaul, scale leaves 1.0—2.5 cm
long and 0.8-—1.6 cm wide; other leaves broadly ovate; middle leaves
4—7(10) cm long and 2—4(7) cm wide, upper leaves 2.5-5.5 cm long
and 0.8—2.5 cm wide, broadly cordate, amplexicaul; all leaves glabrous,
subobtuse or weakly acute, sinuate-short-toothed sometimes entire,
along margin glandular-ciliate. Capitula solitary (less often 2) large.
Involucre 2.2—3.0(4.0) cm in dia; involucral bracts uniform, 1.0—1.3
cm long; outer bracts oblong-lanceolate or lanceolate, (1.5)1.8-2.0
mm wide, at base glandular-pilose; inner bracts linear-lanceolate or
linear, 0.5—1.0 mm wide, smooth, all bracts long-acuminate. Ligulate
florets 2.0-2.5 cm long; corolla 1.8—2.4 cm long, yellow, their tubes
2.0-2.5 mm long, ligules 1.6—2.2 cm long and 2.5—3.5 mm wide, one-
third longer than involucral bracts, smooth, with 3—4 veins and 3 teeth
at apex; central tubular florets 5.0-5.5(6.0) mm long, their corollas
4.0—4.5 mm long, yellow, slightly longer or as long as pappus, with 5

640

small teeth 1 mm long. Achenes dark brown, smooth or sparsely hairy,
2—4 mm long and 0.8 mm wide, all with white pappus, 3-4 mm long,
or with numerous barbed bristles. Flowering June to August.

Alpine and subalpine meadows, spruce forests, juniper thickets.—
Western Siberia: Ob River Area, (southern part), Irtysh, Altai; Eastern
Siberia: Yenisei, Angara-Sayans, Dauria; Soviet Central Asia:
Dzhungaria-Tarbagatai, Pamiro-Alai Region, Tien Shan. General
distribution: Mongolia, Dzhungaria-Kashgaria. Described from Altai.
Type in London.

2. D. schischkinii Serg. in Sistemat. Zam. Gerb. Tomsk. Univ., 1-
2 (1949) 15; Kryl. Fl. Zap. Sib. XI, 2831.

Perennial. Plant 35-60 cm high; rhizome horizontal, creeping, up
to 1 cm thick; stem solitary, erect, densely glandular pilose below
inflorescence, glabrous or sparsely short-hiary in remaining part, like
leaves. Basal and lower cauline leaves broadly ovate or elliptical, (8)11—
12.5 cm long and (4)7-8.5 cm wide, petiole (6)13—23 cm long; middle
cauline leaf with 3-5 cm-long petiole; upper leaves 4—5, ovate or
oblong-ovate, gradually reduced, 2-12 cm long and 0.6-3.5 cm wide,
sessile, semiamplexicaul all leaves with irregular small teeth. Capitula
solitary. Involucre 2 cm in dia; outer involucral bracts lanceolate-
linear, 1 cm long and 1.5 mm wide, sometimes divided up to a half,
densely glandular-pilose; inner bracts slightly shorter, linear, 0.8 cm
long and 0.8 mm wide, smooth; all bracts long- and thin-pointed.
Ligulate florets 1.5-1.6 cm long, corolla yellow, 1.4—1.5 cm long,
corolla tube 1-2 mm long, as long as pappus, ligules 1.3-1.4 cm long
and 1.8—2.0 mm wide, a third or a fourth longer than involucral bracts,
smooth, with 3-4 veins and 3 teeth 1 mm long; central florets 5.0—5.5
mm long, with yellow corolla 2 times as long as pappus, glabrous, with
5 teeth 1 mm long. Achenes glabrous, brownish, pappus 1.5—2.0 mm
long, as long as achene consisting of numerous serrate bristles.
Flowering July to August and September.

Alpine meadows, stony taluses and terraces above glaciers.—
Western Siberia: Altai. Endemic. Described from upper reaches of
Katun River. Type in Tomsk.

3. D. bargusinense Serg. in Sistemat. Zam. Gerb. Tomsk. Univ.
1-2 (1949) 14; Kryl. Fl. Zap. Sib. XI, 2831.

Perennial. Plant glabrous, 45-75 cm high; rhizome horizontal or
weakly ascending, 0.5-1.0 cm thick. Stem basally covered with scale
leaves. Basal leaves ovate, 6-15 cm long and 3.0—7.5 cm wide, glabrous,
inconspicuously toothed, petiole 20-28 cm long; cauline leaves ovate,
4—8 cm long and 2.5-6.0 cm wide, with fine and coarse teeth, sessile,

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641

amplexicaul, only lowermost leaf with short winged petiole, on both
sides glabrous, less often with short sparse hairs. Capitula 2-4
(sometimes solitary), on 10-28 cm-long peduncles with 1-3 small,
lanceolate, acute leaves 4 cm long and 1 cm wide, densely glandular-
hairy below capitulum. Involucre 2.5-3.5 cm in dia; outer involucral
bracts lanceolate, 1.5 cm long and 1.5-1.8 mm wide, inner bracts
lanceolate-linear, 1.3-1.5 cm long and 0.5—0.6 mm wide; all bracts
thin-acuminate, glandular-hairy. Ligulate florets 2 cm long, with yel-
low, glabrous, corolla 1.8 cm long, corolla tube 2 mm long and linear,
ligule 1.6 cm long and 1.5 mm wide, with 3-4 veins and 3 teeth;
tubular florets 6 mm long, with yellow, glabrous, corolla, 4.5-5.0 mm
long. Mature achenes brownish, glabrous or sparsely pilose in upper
part, 3-4 mm long and 1.2 mm wide, ribbed; pappus (4)6 mm long, of
numerous, rusty, barbed bristles. Flowering July.

Subalpine meadows, along banks of streams and rivers and in wet
lowlands.—Western Siberia: Altai; Eastern Siberia: Angara-Sayans.
Endemic. Described from Barguzin Range. Type in Tomsk.

Series 2. Carpatica Gorschk.—Doronicum Subsect. Cardyophylla
Cavill. in Ann. Conserv. Jard. Bot. Généve, XIII-XIV (1909-1911)
341 p. p.—Doronicum Subsect. Grandiflora Cavill. ibid., 354, p. p.—
Basal leaves elliptical, oblong or ovate-roundish, (1)1.7—4.5 cm long
and (1.7)2—3 cm wide, petiole 4—6(13) cm long; ligulate florets with
pale yellow corolla and ligules, pilose at base.

4. D. clusii (All.) Tausch in Flora, XI (1828) 178 excl. var. B. and
var. V.; Cavill. in Ann. Conserv. Jard. Bot. Généve, X, 1220; Ej. op.
cit. XIII-XIV, 359; Popov. Konsp. Fl. Zakarp. Obl. 246.—Arnica clusii
All. in Mel. phil. et meth. soc. Turin, V (1770-1773) 70.—Doronicum
scorpioides Lam. Encyl. meth. II (1786) 313.—D. hirsutum Lam. ibid.—
A. clusii Hausm. FI. Tirol. I (1851) 462; Klok. in Vizn. Rosl. URSR,
552.—Ic.: All. Fl. pedem. tab. XVII, fig. 1 and 2; Hegi, Ill. Fl. VI, 2,
724 and 725.—Exs.: Fl. Exs. Austro-Hung. No. 1817; Fl. Exs. Reipubl.
Bohem.-Sloven No. 953.

Perennial. Plant 9-20(40) cm high; rhizome Bbligue or horizontal;
stem solitary, straight or weakly flexuous, more or less pilose. Leaves
glabrous (var. glabratum Tausch) or sometimes covered with white,
long, sparse, multi-cellular, — hairs (var. villosum Tausch), entire or
sinuate-toothed, elliptical or oblong, 4.5 cm long and 2 cm wide, basally
narrowed into winged petiole 4 cm long; cauline leaves ovate-oblong
or oblong, lower ones 3.5—4.5 cm long and 1.6-2.0 cm wide, basally
narrowed into winged petiole, 2.5 cm long; upper semiamplexicaul, 1—
2 cm long and 0.3-0.8 cm wide, sessile, at base often sinuate-toothed

675

642

or incised; all leaves acuminate. Capitula solitary (less often 2), large.
Involucre 3 cm in dia, with 1.3 cm long, uniformly acuminate involucral
bracts; outer bracts lanceolate, 3 mm wide, covered with multi-cellular,
long, simple and glandular hairs, inner bracts linear, 1 mm wide,
sparsely pubescent; all bracts long-ciliate along margin. Ligulate florets
with golden-yellow corolla, 2.5 cm long, corolla tubes glabrous, shorter
than pappus and ligule, 2.2 cm long and 2.8 mm wide, at base sparsely
pilose; central florets with glabrous, yellow corolla 4 mm long, longer
than pappus, with 5 teeth 0.5 mm long. Achenes 2.5 mm long,
pubescent; pappus in all achenes 3.0-3.5(4.0) mm long, of numerous,
serrate bristles. Flowering July.

Stony, turfy slopes at 1,500-—3,000 m.—European part: Upper
Dniester (Chernaya gora[Black Mountain]; Pope Ivan). General
distribution: Central Europe. Described from Pemonte.

Note. It is a rare plant in our country, collected by M.G. Popov
in the Carpathians, which, as mentioned by him, represents a transition
from D. clusii to the more southern species D. longifolium.

5. D. carpaticum (Griseb. and Schenk) Nym. Syll. fl. europ. Suppl.
(1865) 1; Cavill. in Ann. Conserv. Jard. Bot. Généve, X, 227, XIII-
XIV, 146; Popov, Konsp. Fl. Zakarp. Obl. 248.—Arnica scorpioides
Baumg. Enum. stirp. Fl. Transs. III (1816) 133.—Arnicum scorpioides
var. carpaticum Griseb. and Schenk in Wiegm. Arch. (1852) 342.—A.
carpaticum (Griseb. and Schenk) Schur in Verb. Siebenb. Nat. Ver.
(1859) 137; Klok, in Vizn. Rosl. URSR, 552.—Exs. : Fl. Exs. Austro-
Hung. No. 1816.

Perennial. Plant 12-50 cm high; rhizome horizontal; stem solitary,
erect, simple, glabrous, sometimes glandular-pilose in upper part below
inflorescence. Leaves membranous, glabrous or sparsely pilose, ciliate,
at base of stem approximate; basal leaves usually numerous, ovate-
roundish, (1)1.7—-2.3 cm long and (1.7)2.2-3.0 cm wide, obtusely
toothed, with petiole (2)4—6(14) cm long, at base more or less expanded;
lower cauline leaves 4.5-6.5 cm long and 2.8-4.0 cm wide, fiddle-
shaped or mostly cordately amplexicaul, sessile, mostly acuminate;
upper leaves 2.0-2.5 cm long and 0.8-1.2 cm wide, cordately
amplexicaul, sessile, acuminate. Capitula solitary. Involucre 3 cm in
dia, with similar bracts 1.3 cm long; outer involucral bracts lanceolate,
2 mm wide, sparsely glandular-pilose in lower part, inner ones linear,
1.0-1.2 mm wide, smooth; all bracts acuminate, long-ciliate. Ligulate
florets with pale yellow corolla, 2 cm long, tubes 2.0-2.5 mm long,
shorter than pappus, glabrous, ligules 1.8 cm long and 3 mm wide, a
third longer than involucral bracts, with 3-4 veins and 2-3, mostly
irregular teeth in upper part, sparsely pilose in lower part on outer

676

643

side; central florets with yellow corolla, 3-4 mm long, as long as
pappus and glabrous, with 5 teeth. Achenes brownish, 2 mm long and
0.8 mm wide, whitish-hairy; pappus in all achenes 4(5) mm long,
consisting of numerous, white, barbed bristles. Flowering June to July.

Hollows, turfy slopes and on limestone rocks of alpine zone.—
European part: Upper Dniester (Chernaya Gora, [Black Mountain],
Goverla; Petrosh, Svidovets, Bliznetsy). General distribution: Central
Europe, Balkans-Asia Minor. Described from southern Transylvania.
Type in Leipzig.

Section 2. Pardalianches Tausch in Flora, XI (1828) 182.—
Doronicum sect. Eudoronicum DC. Prodr. VI (1837) 320; Hoffm. in
Pflanzenfam. IV, Abt. 5 (1892) 294.—Doronicum sect. Doronicastrum
Cavill. in Ann. Conserv. Jard. Bot. Généve, XIII-XIV (1911) 339
p. p.—Ovaries and achenes of different types, peripheral glabrous,
very rarely sparsely pilose, lacking pappus, central covered with short,
appressed-pilose, sometimes glabrous, with pappus.

Series 1. Austriaca Cavill. in Ann. Conserv. Jard. Bot. Généve, XIII-
XIV (1909-1911) 338 and 340 pro sub sect.—Stems densely leafy; lower
leaves ovate-cordate, 3.5 cm long, 4 cm wide, at base narrowed, with
approximate auricles and radiating principal veins; capitula numerous.

6. D. austriacum Jacq. Fl. Austr. II (1774) 18; Cavill. in Ann.
Conserv. Jard. Bot. Généve, XIII-XIV, 225, 340; Vizn. Rosl. URSR,
51.—D. longifolium Rchb. Fl. Germ. Exc. II (1830-1832) 234.—Arnica
austriaca Hoppe and Sturm. Deutschl. Fl. Heft. 38 (1814) 16.—Ic.:
Jacq. op. cit. tab. 130; Rchb. f. Ic.: Fl. Germ. tab. 66-CMLVII; Sturm
op. cit. tab. XIX, 2; Hegi, Ill. Fl. VI, 2, 713, fig. 421, 714, fig. 422.—
Exs.: Fl. exs. Reipubl. Bohem.-Sloven. No. 376; Fl. Siles. exs. No.
835; Fl. exs. Austro-Hung. No. 1818; Fl. stir. exs. No. 582.

Perennial. Plant 60-100 cm high; rhizome cylindrical, short,
horizontal or oblique, not branched. Stem erect, branched above,
glandular particularly below capitulum, and pilose on peduncles. Leaves
sparsely pilose above with whitish, multi-cellular hairs, with similar
long, white hairs and short glands beneath (more densely along veins);
lower leaves ovate-cordate, 3.5 cm long, 4 cm wide, obtuse, obtusely
toothed, with slender, petiole 4 cm long; cauline leaves 15-20 cm
long, 8-10 cm wide, ovate, acuminate, lamina decurrent, with cordate,
amplexicaul auricles, 2—4 cm long and 1.5—2.0 cm wide, upper leaves
oblong or lanceolate, 6.5—9.5(17) cm long, 3.5—4.5(6.5) cm wide, long-
pointed, sessile, amplexicaul. Capitula numerous [sic.; recte few] (3-
8), aggregated in corymbose inflorescence. Involucre 2—3(3.5) cm in

677

644

dia, with similar bracts, 8-9 mm long; outer involucral bracts lan-
ceolate, 1.8 mm wide, sparsely pilose; inner ones 1.2 mm wide, nar-
rowly lanceolate, long-glandular-pilose only in upper part; all bracts
acute and ciliate. Ligulate florets 2 cm long, with bright yellow co-
rolla, 1.8-1.9 cm long, tube 2 cm long and linear ligules, twice as long
as involucral bracts, glabrous, with 4-5 veins and 3 teeth above; tubular
florets 6 mm long, their corollas yellow, slightly longer than pappus, with
5.1 mm-long teeth. Achenes oblong, dark green or brown, 2 mm long, 0.8-
1.0 mm wide, with 10 light colored ribs; peripheral achenes glabrous,
lacking pappus; central achenes with white, appressed hairs and 4 mm
long pappus of numerous, white, finely barbed bristles. Flowering July.

Montane, spruce, beech, ash forests, and alder thickets —European
part: Upper Dniester. General distribution: Central Europe, Atlantic
Europe, Mediterranean Region (west), Balkans-Asia Minor. Described
from Austria. Type in Vienna.

Series 2. Cardiophylla Cavill. in Ann. Conserv. Jard. Bot. Généve,
XII-XIV (1909-1911) 338 and 341, pro subsect. p. p. (excl. D.
carpaticum Nym.).—Stems with few leaves, lower leaves broadly ovate-
cordate, 3-5(6) cm long, 3.5—6.0(6.5) cm wide, with remote auricles
and flabellate midrib; capitula solitary or few. Beside D. orientale
Hoffm., this series includes D. carpetanum Boiss. and Reut. from Spain
and D. cordatum from the mountains of central Europe.

7. D. orientale Hoffm. in Comment. Soc. phys. med. Moscou, I
(1808) 8; Willd. Enum. pl. Hort. Reg. Bot. Berol. 898; Cavill. in Ann.
Conserv. Jard. Bot. Généve, XIII—-XIV, 242, 342; Grossh. Fl. Kavk.
IV, 145.—D. caucasicum MB. FI. taur.-cauc. II (1808) 321, III, 577;
DC. Prodr. VI, 320; Ldb. Fl. Ross. II, 625; Boiss. FI. or. III, 380;
Schmalh. Fl. II, 81.—D. pardalianches auct. non L.: Sibth. and Sm.
Prodr. fl. graec, II (1813) 183.—Ic.: Gartenfl. XX VII, 127; Bot. Mag.
LIX, tab. 3143.—Exs.: Fl. cauc. exs. No.100.

Perennial. Plant glabrous, 47 cm high; rhizome horizontal,
nodulose, branched, with lanate scales on collar. Stem solitary, straight
or sometimes more or less flexuous, green. Basal leaves (3-4), broadly
ovate-roundish or roundish-cordate, 3—5(6) cm long, 3.5—6.0(6.5) cm
wide, broadly sinuate-toothed or obtusely toothed, sometimes entire,
with 3.5—5.5(10.0) cm-long petiole; cauline leaves (1-2) ovate or ovate-
elliptical, 3.0-5.5(8.0) cm long, 2.5—4.0(7.0) cm wide, sessile,
amplexicaul, subacute; all leaves smooth, less often light green beneath.
Capitula solitary, on long peduncles. Involucre (3.0)3.5—4.0 cm in dia;
outer involucral bracts linear-lanceolate, 1.6 cm long, 1.2—-1.5 mm wide;
inner narrowly linear-lanceolate, slightly shorter and narrower; all bracts

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long-acuminate, green, sparsely glandular with fine glands, long-ciliate
along margin. Ligulate florets 1.72.4 cm long, with pale yellow corolla
1.6—2.0 cm long, with oblong ligules slightly longer or shorter than involucral
bracts, with 4—5 veins, at base with curly hairs, and smooth corolla tube
2-3 mm long; central florets 5-6 mm long, with yellow corolla, as long as
pappus with 5.1 mm-long teeth. Achenes 1.5-1.8 mm long, 0.5—0.8
mm wide, of different types; outer achenes glabrous, without pappus;
inner covered with stiff appressed hairs; pappus white, 2 times as long as
achene, of numerous, barbed bristles. Flowering March to April.

Mountains, in middle mountain zone.—Caucasus: Ciscaucasia,
Eastern and Western Transcaucasia. General distribution: Central
Europe, Mediterranean Region, Balkans-Asia Minor. Described from
vicinity of Dzekhet.

Series 3. Macrophylla Cavill. in Ann. Conserv. Jard. Bull. Généve,
XIII-XIV (1909-1911) 338 and 344, pro subsect.—Stems with few
leaves; lower leaves broadly ovate-cordate, 9-19 cm long, 9.5-
18.0(27.0) cm wide, twice as wide at base, auricles widely spaced,
principal veins palmate; capitula numerous. Besides D. macrophyllum
Fisch., this series includes D. haussknechtii Cavill., D. maximum Boiss.
and Huet., D. macrolepis Freyn and Sint., D. balansae Cavill., and D.
cacaliaefolium Boiss. and Heldr.

8. D. macrophyllum Fisch. Catal. Jard. Gorenk. (1812) 40; DC.
Prodr. VI, 321; Ldb. Fl. Ross. II, 625; Boiss. Fl. or. II, 379; Schmalh.
Fl. II, 81; Grossh. Fl. Kavk. IV, 145—Cavill. in Ann. Conserv. Jard.
Botan. Généve, XIII-XIV, 248, 345.—D. austriacum MB. FI. taur.-
cauc. II (1808) 321, excl. syn. III, 576 non Jacq.—D. vaginatum C.
Koch in Linnaea, XXIV (1851) 356.—D. dolichotrichum Cavill. op.
cit. XIV (1909-1911) 252.—Ic.: Sturm Deutschl. Fl. tab. 1243; Larin
et al. Korm. Rast. Senokos. i Pastb. SSSR 581, Fig. 426.

Perennial. Plant 75-100 cm high, covered with long, white, multi-
cellular and short-glandular hairs (except leaves, corolla, and achenes
of ligulate florets); rhizome short, without branches. Stem erect, green,
simple or branched above and sometimes reddish-violet, sparsely leafy.
Leaves broadly ovate-cordate, with coarse, acute teeth; basal leaves 9-
19 cm long, 9.5—18.0(27) cm wide, with 22-33 cm-long petiole; cauline
leaves 6.5—16.0 cm long, 7-16 cm wide, with amplexicaul auricles 4
mm long, 4.0-5.5 mm wide, with 4-17 cm-long petiole; upper leaves
5.0-9.5 cm long, 3-7 cm wide, sessile amplexicaul; bracteal leaves a
third to a half as long. Capitula large, on 6-20(32) cm-long peduncles,
in clusters of 2—5(6) in corymbose inflorescence. Involucre (2)3—4(4.5)
cm in dia, with similar involucral bracts, 1.1—-1.3(1.8) cm long, thin-

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and long-acuminate apices, densely pubescent, ciliate; outer bracts
lanceolate or oblong-lanceolate, 1.5(1.8) mm wide; inner narrowly
lanceolate, 1.0(1.2) mm wide. Ligulate florets 2.0-4.2(4.5) cm long,
with bright yellow corolla 1.8-4.0(4.4) cm long, and glabrous tube 2—
3 mm long, ligule longer than involucral bracts, 3.0-4.5 mm wide,
with 4—5 dark veins, short-pilose at base, with 3 teeth above; central
tubular florets 5-7 mm long, with yellow corolla 4.0-5.5 mm long,
sparsely pubescent, slightly longer than pappus, with 1.2 mm-long teeth.
Achenes of peripheral ligulate florets 4 mm long, 1.5 mm wide,
glabrous, without pappus; achenes of central tubular florets 2-3 mm
long, 1.0-1.2 mm wide, covered with white appressed hairs, with
pappus; pappus as long as achenes, of numerous white barbed bristles.
Flowering June to August.

Alpine and subalpine zones, in hornbeam forests, rhododendron thickets, —
and meadows.—Caucasus: Ciscaucasia, Dagestan, Eastern, Western, and
Southern Transcaucasia. General Distribution: Armenia and Kurdistan,
Iran (north). Described from Beshtau Mountains. Type in Leningrad.

Series 4. Pardalianches Cavill. in Ann. Conserv. Jard. Bot. Généve,
XITI-XIV (1909-1911) 338 and 348, pro subsect.—Stems sparsely
leafy; lower leaves broadly cordate-ovate, 4-9 cm long, 3.5—8.0 cm
wide, at base narrow, auricles approximate and main veins divergent;
capitula numerous. Beside D. pardalianches L.., this series includes D.
roylei DC. from the western Himalayas, D. thirkei Schultz. from
Olympus Mountain in Greece. D. reticulatum Boiss. from Lydia, and
D. atlanticum Chab. from the mountains in Algeria and Tunisia.

9. D. pardalianches! L. Sp. pl. (1753) 885 excl. var.; DC. Prodr.
VI, 320; Ldb. Fl. Ross. II, 625; Cavill. in Ann. Conserv. Jard. Bot.
Généve, XIII—XIV, 278 and 350; Vizn. Ross. URSR, 551.—Ic.: Rchb.
Ic. Fl. Germ. XVI, tab. 64, 955, f. II; Sturm. Fl. Deutsch]. Ed. 2, XIII,
tab. 58; Hegi, Ill. VI, 2, tab. 719, fig. 428.

Perennial. Plant 30-120 cm high, covered with several rows of
multi-cellular, white, long and short hairs and sparse glandular short-
stalked glands (except tubular florets and peripheral achenes); rhizome
horizontal, tuberous, 0.8—1.3 cm thick; stem solitary, straight or weakly
flexuous, simple or branched in upper part. Basal leaves ovate-cordate,
4-9 cm long, 3.5-8.0 cm wide, petiole S—17 cm long; middle cauline
leaves similar, (4)8—-11 cm long, (2)7.0—8.5 cm wide, sinuate-toothed,
with auriculate petioles 5.5—-11.0 cm long; upper cauline leaves oblong-

'The Latin epithet of this species is the Greek name of some poisonous plant in
the works of Aristotle, which was used to poison snow leopards; the very name means
“snow-leopard killer.”

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lanceolate 1-8 cm long, 0.44.5 cm wide, amplexicaul, sessile, of these,
lower leaf lyrate; all leaves obtuse or upper leaves more or less subacute.
Capitula 2-5. Involucre 2.5—4.5(5.5) cm in dia; involucral bracts
uniform, long-acuminate, outer narrowly lanceolate, 1.8 cm wide, inner
linear, 0.8-1.2 mm wide. Ligulate florets 1.3-1.6 cm long, with yellow
corolla 1.2—1.4 cm long, their tubes 1.5—2.0 mm long, glabrous, ligules
linear, 1.0-1.2 cm long, mostly as long as involucral bracts, (1.5)2—3(4)
mm wide, with 4 veins and 3 teeth above, in lower part pilose; tubular
florets 4—5 mm long, their corollas yellow, 3.5—4.5 mm long, one-third
longer than pappus, with 5 teeth. Achenes of peripheral ligulate florets
glabrous, without pappus; achenes of central tubular florets appressed-
pilose, with 3 mm long white pappus. Flowering July to September.

Mountains up to 1,500—1,700 m, in forests and coastal scrubs.—
European part: Upper Dniester. General distribution: Scandinavia,
Central Europe, Atlantic Europe, Mediterranean Region (west), Balkans-
Asia Minor (west). Described from Switzerland. Type in London.

Economic Importance. A medicinal plant; its rhizomes are used
in the treatment of epilepsy and fits (Visn. Rosl. URSR, 551). Culti-
vated in Western Europe as a beautiful ornamental plant.

Series 5. Plantaginea Cavill. in Ann. Conserv. Jard. Bot. Généve,
XIII-XIV (1909-1911) 338 and 351, pro subspec.—Stems sparsely or
densely leafy; lower leaves oblong, elliptical or obovate-oblong, less
often roundish, 2.5-11.0 cm long, 1.7-8.5 cm wide, at base narrowed
into petiole, its main veins divergent; capitula solitary or few. Besides
D. longifolium Griseb, and Schrenk, D. oblongifolium DC., and D.
turkestanicum Cavill., this series includes two more species: D.
plantagineum L. from the Iberian Peninsula, southern England, and
France, and D. falconeri Clarke from high-altitudes of Kashmir and
western Tibet (Karakorum).

10. D. longifolium Griseb. and Schenk in Wiegm. Archiv. XVIII
(1852) 341, non Rchb.; Cavill. in Ann. Conserv. Jard. Bot. Généve,
XIII, 292, 353.—D. hungaricum Rchb. f. Ic. Fl. Germ. et Helv. XVI
(1854) 34; Boiss. Fl. or. III, 381; Schmalh. Fl. II, 81; Fedtsch. and
Fler. Fl. Evrop. Ross. 988; Vizn. Rosl. URSR, 551.—Ic.: Rchb. f. op.
cit. tab. 65, fig. 1—Exs.: Fl. exs. Reipubl. Bohem.-Gloven. No. 752
and No. 952; Fl. Hung. exs. No. 83; Fl. exs. Austro-Hung. No. 3441.

Perennial. Plant 25-80 cm high, more or less glandular-pilose;
rhizome tuberous, branched. Stem erect, green or in upper part
brownish-violet, at base usually densely leafy. All leaves undivided,
glabrous or sparsely pilose, entire or sinuate-toothed; basal and lower
cauline leaves oblong-elliptical or oblong, (4)5—11 cm long, 2.0-3.8

68

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648

cm wide, subobtuse, narrowed into 4—9 cm long petiole; cauline leaves
lanceolate or broadly linear-oblong, 2—5(8.5) cm long, 0.7—2.0 cm
wide, sessile, at base toothed; upper leaves thin-acuminate. Capitula
mostly solitary. Involucre 3.0—3.5 cm in dia; involucral bracts linear-
lanceolate, similar, 1.3-1.5 cm long; outer bracts 1.2—-1.8 cm wide,
inner 0.8-1.0 cm wide; all bracts glandular-pilose, long- and thin-
acuminate, ciliate along margin. Ligulate florets (1.6)2.6—3.1 cm long,
with bright yellow corolla (1.5)2.5—3.0 cm long, their tubes 2.0—-2.5(3.0)
mm long, ligules (1.3)2.3-2.7 cm long, 2 mm wide, with 4 veins, glabrous,
with 3 teeth above; central florets 5 mm long, with yellow, glabrous
corolla 4 mm long, slightly longer than pappus, with 5 teeth 1 mm
long. Achenes of different types; peripheral achenes without pappus;
central covered with white, upright hairs, with pappus; pappus of
numerous, white, barbed bristles, 2.5—3.0 mm long. Flowering April.

Forests and glades.—European part: Upper Dniester
(Transcarpathian Region; Chernaya Gora[Black Mountain], vicinity of
Vinogradov), Bessarabia, Black Sea Region. General distribution:
Central Europe, Balkans-Asia Minor (northwestern part). Described
from Hungary. Type in Vienna.

11. D. oblongifolium DC. Prodr. VI (1837) 321; Ldb. Fl. Ross. II,
625; Boiss. Fl. or. III, 381; O. and B. Fedtsch. Perech. Rast. Turk. IV,
213; Cavill. in Ann. Conserv. Jard. Bot. Généve, XIII—XIV, 296, 353;
B. Fedtsch. Rast. Turk. 742; Grossh. Fl. Kavk. IV, 145.

Perennial. Plant glabrous, 12-50 cm high; rhizome short, truncate,
at collar covered with black scale-like bristles. Stem solitary, flexuous,
green, sometimes reddish-violet above, with simple and glandular hairs
in upper part. Basal leaves elliptical or obovate-oblong, 2.5—7.0(10.0)
cm long, 1.7—3.0(4.3) cm wide, obtuse, narrowed into petioles up to
16(20) cm long; lower cauline leaves similar to basal leaves but
narrowed into broadly winged petiole, semiamplexicaul or sometimes
sessile; middle leaves linear-oblong, 4-12 cm long, 2-3 cm wide,
sessile, like upper leaves somewhat acute, semiamplexicaul, upper 3—
4 cm long, 1 cm wide; all leaves glabrous, entire or indistinctly toothed,
glandular-ciliate along margin. Capitula solitary, on long peduncle,
with involucre 3—4(4.5) cm in dia. Involucral bracts 1.2—2.0 cm long,
covered with numerous white, long, simple, capitate, stalked glands,
acuminate; outer bracts lanceolate, 1.5-3.5(4.0) mm wide, inner
narrowly lanceolate, 1.2-3.0 mm wide. Ligulate florets 2.2—2.8(4.0)
cm long, their corollas pale yellow, 2.0—-2.6 cm long, with 2 mm-long
tubes and 1.8-2.4(3.8) cm-long and 3.6—6.0 mm-wide ligules, with 4—
6 dark veins and 3 teeth above, hairy outside at base; central florets
5—6 mm long, with yellow glabrous corolla, 3.0-3.5(4.5) mm long,

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with large, 1.5 mm-long teeth. Achenes different types, peripheral
achenes 1.8 mm long, 0.5 mm wide, glabrous or sparsely appressed-
pilose, without pappus; central achenes 2.0-2.8 mm long, 0.8 mm wide,
appressed-pilose; pappus of numerous, white, barbed bristles, 2.0-
2.3(3.0) mm long and 0.8 mm wide. Flowering June to July.

Alpine zone, in ravines, passes, moraines, near glaciers, in subalpine
and alpine meadows, spruce forests, and shrub thickets.—Caucasus:
Ciscaucasia, Dagestan. Eastern, Western, and Southern Transcaucasia;
Western Siberia: Altai; Soviet Central Asia: Dzhungaria-Tarbagatai,
Pamiro-Alai, Tien Shan. Endemic. Described from eastern Caucasus.
Type in Geneva.

12. D. turkestanicum Cavill. in Ann. Conserv. Jard. Bot. Généve,
XIII-XIV (1909-1911) 301, 354; Kryl. Fl. Zap. Sib. XI, 2832.—D.
oblongifolium DC. var. leiocarpum Trautv. in Bull. Soc. Nat. Mosc.
XXXIX (1866) 361; O. and B. Fedtsch. Perech. Rast. Turk. IV, 213.—
Ic.: Cavill. op. cit. fig. 46 and 47 (p. 302).

Perennial. Plant 25-80 cm high; rhizome horizontal or oblique.
Stem solitary, erect, simple, covered with sparse, capitate glandular
hairs, sometimes glabrous below. Leaves entire or with sparse small
teeth, with short-acuminate or subobtuse, glabrous or sparsely short-
pilose beneath and along margin. Basal leaves obovate-spatulate or
almost round, 3-11 cm long, 4.0-8.5 cm wide, on winged, more or
less condensed petiole, 5-15 cm long; cauline leaves oblong-ovate or
oblong, 3—10(13) cm long, 2.0-4.5 cm wide, upper leaves 1-3 cm
long, (0.4)1.4—2.0 cm wide; less often lower 2-3, with petiole (4)7-11
cm long, leaf next to them narrowed into winged, flattened, petiole 3—
4 cm long and 2.5 cm wide, other leaves sessile. Capitula solitary,
with involucre 3—4(5) cm in dia. Outer involucral bracts lanceolate or
lanceolate-linear, 1.2-1.7 cm long, 1.8—2.0(3.0) cm wide, inner linear,
1.2-1.5 cm long, 1.0-1.2(1.5) mm wide; all bracts long-acuminate,
glandular-hairy. Ligulate florets 1.8-3.0 cm long, with light yellow
corolla 1.6—2.7 cm long, their tube 2.5 mm long, densely hairy and
glandular on outside, ligules 1.4—2.3(2.4) cm long, 2.0-2.8 mm wide,
slightly or a third longer than involucral bracts, glabrous, with 4—S
veins and 3 teeth above, central florets 5.5—7.0 mm long, their corollas
dark yellow, 4.5-5.5 mm long, 1/3 longer than pappus, with 5 teeth 1
mm long. Achenes of different types; outer achenes glabrous, 2.8(4.0)
mm long, 0.5(0.8) mm wide, without pappus; central achenes (2.0)3.0-
3.5(4.0) mm long, 1 mm wide, glabrous, covered wih sparse, appressed
hairs; their pappus 3.0-3.5 mm long, of numerous, barbed bristles.
Flowering June to August.

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Alpine region, mixed herb meadows, on stony slopes of moraines,
rocks, stony and rubbly tundras and in spruce forests and juniper
thickets.—Western Siberia: Altai; Soviet Central Asia: Dzhungaria-
Tarbagatai, Tien-Shan. General distribution: Dzhungaria-Kashgaria.
Described from Soviet Central Asia.

GENUS 1560. Erechtites Raf.':?

Raf. Fl. Ludov. (1817) 65.—Neoceis Cass. in Bull. Soc. Phil. (1820)
90.

Capitula many-flowered, heterogamous, peripheral florets pistillate,
filiform-tubular, in 1-2 rows; disk florets bisexual or partly sterile, with 4—
5-toothed, narrowly tubular corolla; anthers at base without or with very
short auriculate appendages; style branches with conical, appendages hairy
in upper part. Achenes oblong, with 10-20 fine ribs; pappus many-rowed,
consisting of weakly scabrous hairs. Erect annual, or perennial, with
alternate, undivided or pinnatipartite leaves. Capitula with cylindrical
or campanulate involucre; involucral bracts few, narrow, linear-setaceous.
Inflorescene compound, corymbose; corolla yellow or whitish.

Species of this genus are distributed in North and South America.

1. E. valerianifolia (Wolf) DC. Prodr. VI (1837) 295; Makashvili,
in Byull. Inst. Chaya i Subtrop. Kult. 3, 106; Manden. in Fl. Gruzii,
VIII, 417.—Senecio valerianifolius Wolf, Ind. sem. hort. berol. (1825)
ex Rchb. Icon. bot. exot. I (1827) 59.—Crassocephalum valerianifolium
Less. in Linnaea, V (1830) 169, in synon.—Ic.: Rchb. Ic. bot. exot. I
(1827) tab. 85 (excl. fig. stigm.).

Annual. Plants subglabrous, with erect, sulcate stem. Leaves
pinnatipartite, with 6-8 lanceolate, short-toothed lobes on each side,
sinuses between lobes narrow, with remote teeth or lobules. Inflorescence
paniculate, with erect branches, of numerous capitula, more or less erect,
later drooping; involucre cylindrical; pappus of pinkish-violet hairs.

An introduced plant, mostly a weed in tea plantations.—Caucasus:
Western Transcaucasia (Chakva, Kobulety, and others). General
distribution: Brazil and Central America. Described from Brazil? Type
unknown.

Besides this species, one other species of the genus Erechtites, E.
hieraciifolia (L.) Raf. is reported from the USSR from the Mukachev
District (M.G. Popov, Ocherk Rastitelnosti Karpat, 1949, p. 245), which
so far is found only in Western Ukraine. It is an annual, with the

'Treatment by A.I. Pojarkova.
*Erechtites is a synonym of Senecio in the work of Dioscorides.

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leaves sessile, oblong-lanceolate, irregularly coarse-toothed glabrous
below, scabrous beneath along veins. Capitula 12-15, in corymbose
inflorescence; involucre cylindrical, consisting of linear, scabrous,
involucral bracts; pappus white. This plant comes from the southern
states of North America and Mexico.

GENUS 1561. Cacalia L.':?

L. Sp. pl. (1753) 834, p. min. p. (quoad sp. Nos. 7, 8); Hoffm. in
Pflanzenfam. IV, 5 (1894) 296, p. min. p.; Kitam. Compos. Japon. III
(1942) 202, p. max. p. (excl. syn. Mesadenia).—Hasteola Raf. New
Fl. North. Am. IV (1836) 79; Pojarkova, in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XX (1960) 380.—Synosma Raf. in Loud. Gard.
Mag. VIII (1832) 247, nom. nud.; Britt. in Britt. a. Br. III, fl. N. Un.
III (1898) 474, descr.—Cacalia sect. Eucacalia DC. Prodr. VI (1837) 327
p. min. p.—Senecio Benth. and Hook. f. Gen. pl. II (1873) 446, p. min.
p. non L.—Senecio subg. Cacalia Maxim. in Mél. biol. [IX (1877) 292.

Capitula homogamous, with (1)3—20(30) flowers, most often with
5-8 flowers. Involucre narrow, mostly tubular, consisting of (2)3—12(15)
free involucral bracts, in single row, sometimes with few, very small
pales at base. Involucral bracts dimorphic; inner broader, with wide
scarious border, outer narrower, with very narrow scarious fringe.
Receptacle flat or weakly convex, finely alveolate. Flowers bisexual,
all tubular; corolla whitish, with violet or yellowish tinge, in upper
part campanulate or almost tubular, lobed up to a fourth to a third with
deltoid teeth, abruptly narrowed below into narrow tube; anthers with
narrow, deltoid apical appendage, sagitate, with sterile, acute, short,
less often long, auriculate basal appendage fused in pairs with adjacent
anthers; filaments in upper part expanded (with antherozoids). Style
branches elongate, apically with conical, acute appendage, on outer
side only below appendages or sometimes up to style fork covered
with papillate hairs, longer at base of appendage, often forming tuft.
Achenes narrowly terete, glabrous, brownish-yellowish, monochromatic,
lacking rostellum, somewhat narrowed toward base, with many thin
ribs; pappus mostly white, less often brownish, very rarely reddish,
consisting of numerous, scabrous hairs. Perennial herbs; lamina
reniform, roundish or deltoid-hastate, involute until opening or inclined
to petiole and folded like closed umbrella; lower leaves long-petiolate,
often with auricles sometimes fused along margin forming sheath.
Inflorescence often paniculate, less often corymbose or racemose.

‘Treatment by A.I. Pojarkova.
?Name of the plant in the work of Dioscorides.

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652

The genus is native mainly to eastern and southeastern Asia, where
about 50 species are concentrated; one species (C. hastata L.) extends
through Siberia into the northern part of the European territory of the
USSR, and one (C. suaveolans L.) grows in the eastern part of North
America.

Lectotype of genus: C. hastata L.

Note. Until now the question of the lectotype of the genus Cacalia
L. could not be resolved, and in this connection it is not yet clear what
generic name should be given to the Asian species of this genus, if it
is treated in a narrow sense as in the present treatment, where the
name Cacalia is limited to species of the Asian continent and one
North American species (C. suaveolens L.). Cacalia has already been
typified repeatedly. In my opinion, the question of the lectotype of the
genus Cacalia was resolved best, i.e., in conformity with the
recommendations of the International Code of Botanical Nomenclature,
1956, by Kitamura (Compos. Japon. III, 1942, p. 170), who proposed
C. hastata L. as the lectotype. Unfortunately, this lectotype does not
have priority and cannot be recognized. However, at the same time,
neither of the two lectotypes proposed earlier can be accepted
unequivocally. Cacalia alpina, proposed by Rydberg (Bull. Torr. Bot.
Club. 5, 1924, p. 369), had already been split off by Cassini (1816) as
the genus Adenostyles Cass. and is its type. The second lectotype,
proposed in the list of lectotypes of species of Linnaean genera (A.S.
Hitchcock and M.I. Green, Propos. Brit. Bot., 1929), is C. atriplicifolia
L. This species was split off by Rafinesque (New Flora of N. Amer.,
1836, p. 78) as the separate genus Mesadenia Raf. and was selected
in 1898 (Britton a. Brown, Illustr. Fl. North. Un. St., III) as the lectotype
of this genus. According to the instructions in Appendix IV of the
Code (Russian edition), the type of a aggregate genus that undergoes
splitting must be selected from among the species remaining in it after
the previous authors have or “author has already split off one or two
elements as other taxa.” These species split off earlier from an aggregate
genus take priority over the lectotype selected later from among the
remaining species of the genus only if it is proved that they satisfy
better the requirements of a lectotype. Meanwhile C. alpina and C.
atriplicifolia, as lectotypes, not only do not have priority over C. hastata
but inevitably would lead to very undesirable consequences if either of
them were to be restored as lectotype of the genus Cacalia. For example,
authors who recognize the separate status of the genus Mesadenia, will
be faced with the need to rename all the Asian species of the genus
Cacalia s. 1. (and there are about 50) and C. suaveolens, creating for
them new combinations with the new generic name. As I have shown
in my work (Pojarkova, 1960), Hasteola Raf. is such a generic name.

653

Selecting C. hastata L. as the lectotype for the genus Cacalia does not
entail the need for new combinations. In the present treatment, I have
retained the generic name Cacalia for our species with confidence that
C. hastata L. will be recognized as the lectotype of the genus Cacalia.

i.

Petioles of all leaves narrowed toward base and lacking auricles;
middle cauline leaves deltoid hastate, with attenuate middle lobe
and small, acute or acuminate, usually undivided lateral lobes;
capitula with 4—10(20) florets; involucre campanulate, consisting
Of 8—10 involucral bracts ............ccccecsceeseeseeseeseeees 1. C. hastata L.
Petioles, at least of middle cauline leaves, expanded at base into
amplexicaul auricles; lamina reniform, deltoid-reniform or broadly
deltoid hastate; capitula with 5—8(12) florets; involucre cylindrical/
or campanulate-cylindrical, consisting of 4—7(8) involucral bracts

Plants tall, up to 2-3 m high, robust; leaves to 36 cm long and 50
cm wide; capitula aggregated into large (to 60 cm long) pyramidal,
strongly branched panicle with rather numerous (to 150-350)
capitula, lateral branches of such panicles also with capitula
BROTERBIEM TMG NATMCIES -Feiiraes. Sash caisiis os 00 Shddaltl dada BOLLda cases Se
Plants not so tail, 30-100 cm high, with leaves up to 25 cm wide,
but usually much smaller; general inflorescence simple raceme or
with less numerous florets and a shorter panicle; with capitula
aggregated in raceme on its lateral branches ................::esceeeeeee 4.
Middle cauline leaves up to 35 cm long and 50 cm wide, broadly
deltoid or broadly deltoid hastate, with broad, bifid lateral lobes
and short-acuminate middle lobe; inflorescence with up to 200-350
capitula; involucre narrowly cylindrical, consisting of 4-5 involucral
bracts; florets inecapitula: (3)SA7 : t..<csciag. 0106). Ruin aeee
sesssseessessceseeeesesseseeseesesseeeeeeeesseeee Oe Co KOMaroOviana (Pojark.) Pojark.
Middle cauline leaves up to 35 cm long and 40 cm wide, deltoid
or hastate, with more or less enlarged middle lobe and small acute
or acuminate, undivided lateral lobes; panicles with up to 150-
200(300) capitula; involucre campanulate-cylindrical, consisting of
(5)6—8 involucral bracts; florets in capitula 7—10(12) ............
Cn PRES on AUR, os SAL cobl tn 8s de VE, oo cavern dere ones 2. C. robusta Tolm.
Involucral bracts (4)5—8(9) mm long, at anthesis (before development
of achenes) 1/2-2/3 as long as corolla and pappus; capitula
aggregated in simple raceme, less often as condensed raceme with
few short lateral branches in lower part; plant 30-100 cm high,
with slender, 1.5—2.0(3.0) mm thick stem and remote leaves; middle
cauline leaves 5-10 cm long and 7—14(16) cm wide .................0.
pervens MUR aM tes ap habs <b bieewxdchliors usb hy outs ates odbeanihvackt 5. C. auriculata DC.

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654

+ Involucral bracts 8—11(12)mm long, at flowering as long as corolla
and pappus or slightly shorter, approximately 3/4, rarely 2/3 as
long; capitula aggregated into panicles; plants taller, up to 120-150
cm high, with densely leafy stems, up to 7-8 mm thick; middle
cauline leaves up to 15 cm long and 25 cm wide ............:::se0e a

5. Inflorescence compact, with long, upright lateral branches often
adpressed to main axis; peduncles thick; leaves irregularly toothed,
large, with coarse, lobed teeth alternating with somewhat smaller
teeth icc tea. eRe ee 6. C. kamtschatica (Maxim.) Kudo.

+ Inflorescence lax, narrowly cylindrical, panicle with usually remote
branches; peduncles slender; leaves regularly toothed .................
UR RRS A. Te 4. C. praetermissa (Pojark.) Pojark.

Series 1. Hastatae Pojark.—General inflorescence lax pyramidal
panicle with numerous capitula; lateral branches of inflorescence mostly
in racemes, or weakly branched panicles. Involucre cylindrical-
campanulate, with 8—10(12) involucral bracts. Flowers in capitulum 8—
15(20), corolla tube two-fifths to two-thirds as long as limb, oblong-
campanulate. Pappus of achenes white. Plants 40-150 cm high, with
numerous, approximate, cauline leaves; lamina deltoid-hastate, up to
26 cm wide, with small undivided lobes and sinuate toothed margin.

C. hastata L. belongs to this series, which, judging from the
herbarium material, has a closely related race in central China.

Not C. robusta Tolm., as A.I. Tolmatchev believes (1957), but the
eastern North American species C. suaveolens L. is a close relative of
C. hastata. It is distinguished from C. hastata by a corymbose
inflorescence, more numerous-flowered capitula, shorter basal
appendages and bigger antheropoidea. C. suaveolens must be put into
a separate monotypic series Suaveolentes Pojark.

1. C. hastata L. Sp. pl. (1753) 835; DC. Prodr. VI, 327; Turcz.
Fl. baic.-dahur. II, 86; Maxim. Prim. fl. amur. 164; Kom. Fl. Manchzh.
III, 691; Fl. Kamch. III, 171; Fedtsch. and Fler. Fl. 989; Kitag. Lineam.
fl. Manshur. 441; Kitam. Compos. Japon. III, 25 (excl. ssp. and var.
var.).—Cacalia suaveolens auct., non L. (1753); Georgi, Beschr. R.
Reich. III, 5 (1800) 1229.—C. glabra Ldb. Ind. sem. hort. Dorpat.
(1820) 16.—C. sagittifolia Mertens in Linnaea, V (1830) 63, nom.
nud.—Ligularia hastata Less. Synops. Compos. (1832) 390; Turcz.
Catal. baic. (1838) 11.—Senecio sagittatus Sch. Bip. in Flora, XXVIII
(1845) 498.—C. hastata a. pubescens Ldb. FI. Alt. [TV (1833) 52; Fl.
Ross. II, 626; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1044;
Kryl. Fl. Zap. Sib. XI, 2834.—C. hastata B. glabra Ldb. FI. Alt. IV
(1833) 52; Fl. Ross. II, 626; Kryl. Fl. Zap. Sib. XI, 2834.—Senecio

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sagittatus &. pubescens Maxim. in Bull. Acad. Sc. Pétersb. XIX (1874)
483; Mél. Biol. IX, 293.—S. segittatus B. glaber Maxim. in Bull.
Acad. Sc. Petersb. XIX (1874) 483; Mél. biol. IX, 293, p. p. (excl. pl.
jap.).—C. hastata ssp. hastata Hara, Enum. Spermatoph. japon. (1952)
149.—Hasteola hastata Pojark. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XX (1960) 381.—Synosma hastata Pojark. olim in
sched.—Ic.: Pall. Fl. Ross. tab. 110; Fedtsch. and Fler. Fl. Fig. 989,
Kom. and Allis. loc. cit. plate 313; Krasch. in Fl. Yugo-Vost. Evrop.
Ch. SSSR, VI, Fig. 689; Pojarkova, loc. cit. Fig. 3.

Perennial. Plant with horizontal rhizome, densely covered with
slender, filiform, adventitious roots. Stem robust, erect, simple, 40-
150 cm high and up to 8 mm thick at base, glabrous or subglabrous
below, densely glandular hairy-floccose above below inflorescence, as
also peduncle. Cauline leaves numerous, very approximate, glabrous
on both sides (f. glabra Ldb.) or with more or less dense, appressed
hairs (f. pubescens Ldb.) beneath; lowermost leaves strongly expanded
deltoid-reniform, with hastate base, long-petiolate, withering early,
considerably smaller than leaves above; middle cauline leaves 5.5—
25.0 cm long and 6-26 cm wide, broadly deltoid-hastate, with acute or
acuminate lobes, lateral lobes small, undivided and with truncate,
cuneate or broadly sinuate base decurrent as wings in upper part of
petiole; leaves gradually reduced toward stem apex, with increasingly
fewer lobes and becoming deltoid; uppermost and bracteal leaves
lanceolate; bracteal leaves usually linear, all leaves usually sinuate-
and small-toothed. Capitula drooping, usually aggregated into lax,
narrowly pyramidal panicles, with long branches directed upward at
acute angle, sometimes in racemes only in weaker plants; peduncles
(axes of last order) 4-20 mm long; capitula 9-13 mm long and up to
8 mm in dia, with 8—15(20) florets: Involucral bracts linear to almost
lanceolate, greenish, glandular-floccose on outer side, sometimes only
in lower part, with usually 2—4 small subulate pales at base. Corolla
whitish, 8.5-11 mm long, its tube two-fifths to a half as long as,
oblong or narrowly campanulate limb, limb incised usually up to a
third, less often almost to middle, into narrow, lanceolate teeth with
long, acute apices; basal appendages of anthers elongated. Achenes
light brown, 6-8 mm long; pappus white, as long as achene or half as
long. Flowering July to August; fruiting from September.

Thin coniferous and small-leaved forests, rarely in pine woods in
the Far East also in oak forests, shrubby thickets in river valleys, forest and
riverine meadows; in mountains along river valleys in forest zone, in places
ascending to lower part of alpine zone.—Arctic: Anadyr (at the south-
ern boundary of the region); European part: Dvina-Pechora (in
the west, barely crossing the Severnaya Dvina River), Volga-Kama,

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Trans- Volga (rarely, western part); Western Siberia: Ob Region, Irtysh
(eastern part), Altai; Eastern Siberia: Yenisei, Angara-Sayans,
Lena-Kolyma, Dauria; Far East: Kamchatka (including Commander
Islands), Okhotsk, Zeya-Bureya, Uda River Area, Ussuri, Sakhalin
(Sakhalin and southern Kuril Islands). General distribution: Mongolia
(north), China (northeast), Korea, Japan? Described from Siberia. Type
in London.

Hybrids of C. hastata L. x C. kamtschatica (Maxim.) Kudo. (=C.
auriculata Hultén (non DC.) x C. hastata L. (Hultén), Fl. Kamtsch. 1V
(1930) 195, Fig. 7c-f, excl. synon.) collected in Kamchatka on the
Koryatskaya River and in the interfluve of the Gorelaya and Bolshaya
Topolevaya rivers are intermediate forms between the above species,
inclined more toward C. kamtschatica; the middle cauline leaves are
shaped more like the leaves of C. kamtschatica but with longer, *
acuminate, and somewhat narrower lobes; the upper ones are more like
the corresponding leaves of C. hastata. The leaves in both the cases
are more or less uniformly toothed or with irregular teeth, but not as
sharply as in C. kamtschatica; the petioles have very weakly developed
auricles or are without them; the inflorescence is dense, paniculate of
several crowded, long branches, the capitula are large as in C. hastata
but with fewer involucral bracts (5-7) and flowers; the shape of the
corolla is like that of C. kamtschatica; the achenes are undeveloped.

Note 1. Hultén (op. cit.) placed Calcalia auriculata a. ochotensis
Maxim. in the synonymy of this hybrid form. Meanwhile, Maximowicz
considered his own combination Senecio davuricus a. ochotensis
Maxim. as the type form of Calcalia auriculata DC. (Senecio davuricus
Sch. Bip.), a species broadly interpreted by him. At the same time,
Maximowicz made the combination Senecio davuricus B. kamtschatica
Maxim. for the species later called Cacalica kamtschatica (Maxim.)
Kudo. The name C. auriculata DC. was used invalidly by Hultén for
C. kamtschatica (Maxim.) Kudo, whereas it is clearly indicated in the
work of De Candolle that C. auriculata was described by him from the
plant collected between “Yakutia and Okhotsk,” which he received from
Turczaninow who determined it as Ligulatia [sic.; recte Ligularia]
auriculata Turcz. In the herbarium of the Botanical Institute of the
Academy of Sciences of the USSR, there are four sheets from I.
Kuznetsov’s collections in 1835 from the Urak River (Okhotsk District),
all of which bear the annotation “Ligularia auriculata Turcz.” written
in Turczaninow’s own hand. There is hardly any reason to doubt that these
specimens are duplicates of the type of C. auriculata DC. They all conform
fully to the description given by De Candolle for C. auriculata.

Note 2. Given that C. hastata is not always distinguished from the
Japanese geographic races of the series Robustae, the question of the

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occurrence of this species in Japan still remains unresolved. Kitamura
(1. c.) does not report this species for Japan, which he considers as the
type subgenus of the aggregate species Cacalia hastata s. |. However,
in the herbarium of the Botanical Institute of the Academy of Sciences
of the USSR, there is a wholly typical specimen of C. hastata (glabrous
form) from the herbarium of Sakurai, on whose label it is mentioned
that it was collected from the north of Honshu Island (Mutsu Prefecture,
city of Ivakisan).

Series 2. Robustae Pojark.—Capitula numerous, up to 200-350,
aggregated in rather large, lax, strongly branched, pyramidal panicles,
60 cm long, with branches in form of many-flowered panicles; involucre
narrow, cylindrical or campanulate-cylindrical with (4)5—7(8) involucral
bracts, at base without involucral bracts. Flowers in capitulum 5—7(10);
corolla tube narrow 1/4 —1/3 as long as limb; limb narrow, campanulate-
tubular, cut a fourth to a third of the length into 5 narrowly deltoid
teeth. Tall plants, up to 2-3 m high, with large, hastate leaves; cauline
leaves few, up to 60 cm wide, their lateral lobes broad, mostly bifid.

The series Robustae includes the following four Far Eastern species,
of which two, C. robusta Tolm. and C. komaroviana (Pojark.) Pojark.,
penetrate the USSR boundaries, but C. aidzuensis Koidz. and Cacalia
tschonoskii Koidz. are endemic to Japan. Species of this series, still
not studied, are also found in western and central China.

2. C. robusta Tolm. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVIII (1957) 237.—Cacalia hastata auct. non L. (1753): Fr.
Schmidt. Fl. Sachal. (1868) 151.—Senecio sagittatus B. glaber Maxim.
in Bull. Acad. Sc. Pétersb. XIX (1874) 483 p. p.; Mél. biol. IX, 293
p. p. (quoad pl. jap.).—Cacalia hastata B. glabra auct. non Ldb. (1833):
Maxim. in sched. Matsum. Ind. pl. jap. II, 2. Dicotyl. 633, p. max. p.
Kudo in Bot. Mag. Tokyo, XXIX, 229 (excl. area Honsiu); Hara in
Bot. Mag. Tokyo, LII, 66 p. p. (quoad pl. Jap. et excl. synon.).—C.
hastata ssp. orientalis Kitam. in Acta Phyt. Geobot. VII (1938) 244
p. p. (excl synon. praeter 1, 6); Compos. Japon. III, 216 p. p.; Sugaw
Ill. fl. Saghal. IV, 1843; Hara, Enum. Spermatoph. Japon. 149 p. p.
(quoad syn. 1, 6); ?7Tatewaki in Acta hort. Gotoburg. XXI (1952) 112.—
C. hastata var. orientalis Ohwi, Fl. Japon. (1953) 1177.—Hasteola
robusta Pojark. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX
(1960) 381.—Synosma robusta Pojark. olim in Sched.—Ic.: Sugaw.
op. cit. tab. 845; Tolm. op. cit. Fig. 1; Pojark. op. cit. Fig. 4.

Perennial. Plant tall, up to 2.5—3.0 m high, with creeping rhizome.
Stem in lower part thick, straight, in upper part slender, more or less
bent, glabrous or hairy below inflorescence, sometimes densely hairy

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with short thickish hairs. Leaves glabrous above, with flexuous hairs
along veins beneath, sometimes entirely pubescent; lower leaves del-
toid reniform, withering early, small; middle leaves up to 35 cm long
and 40 cm wide, deltoid or almost hastate, with broad base, and at-
tenuate, short-acuminate middle lobe, and small, oblong-deltoid, undi-
vided acuminate, lateral lobes; leaf margin usually convex, sinuate-
toothed, with sharp-pointed teeth; petioles broadly winged, auriculate
with auricles a third to two-thirds as long as lamina; leaves in upper
part of stem sharply reduced in size, deltoid or deltoid-hastate, usually
with small, acute lobes or oblong-ovate, without lobes. Panicle up to
50 cm long, with up to 200(300) capitula, and long, slender, mostly
floccose, acutely divergent branches bearing capitula usually only above
middle; peduncles of last order 3-12 mm long, mostly floccose, pales
narrowly linear to broadly lanceolate. Involucre campanulate-cylindri-
cal, 9-10(12) mm long, consisting of (5)6—-8 linear and lanceolate-
linear, green, glabrous or weakly floccose, involucral bracts. Florets in
capitulum 7—10(12); corolla violet-grayish, 6.0-7.5 mm long, with its
tube about a third as long as campanulate limb; limb up to a third cut
into narrowly deltoid teeth. Achenes 6—8 mm long; pappus brownish.
Flowering July to August; fruiting from middle of September.

Tall-grass thickets in valleys, alder thickets and willow stands,
and also wet slopes of lower mountain zone.—Far East: Sakhalin
(southern part of Sakhalin Island); according to reports of Japanese
botanists also found on Kuril Islands. General distribution: Japan
(Hokkaido Island, and, according to Japanese sources, in northern part
of Honshu Island). Described from vicinity of the town of Dolinsk
from “Kholodnaya Dolina” [Cold Valley]. Type in Vladivostok;
topotype in Leningrad.

Note. A.I. Tolmatchev (op. cit. 244) reports that he observed forms
on Sakhalin Island in which the characters of H. hastata and H. ro-
busta are incorporated in varying combinations. Given the absence of
direct phyletic links between these two species, such forms, if, of
course, their existence were to be proved by actual collections, could
be considered only as interspecific hybrids. I did not find such forms
in the herbarium material of the Botanical Institute of the Academy of
Sciences of the USSR.

3. C. komaroviana (Pojark.) Pojark. comb. nova.—Cacalia
farfaraefolia &. ramosa auct. non Maxim. (1874): Kom. Fl. Manchzh.
III, 689 (excl. synon. and area Japon.); Nakai Fl. Korean, II, 35; Fl.
sylv. Korean. XIV, 106.—C. farfaraefolia auct. non Sieb. and Zucc.
(1846): Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1053.—C.
hastata ssp. orientalis Kitam. in Acta Phyt. et Geobot. VII (1938) 244,

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p. min. p. (quoad synon. Kom.); Compos. Japon. III, 216 p. p. (quoad
areas Korea and Mandsch.).—Hasteola komaroviana Poljark. in Bot.
Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX (1960) 381.—Synosma
komaroviana Pojark. olim in sched.—Ic.: Pojark. loc. cit. Fig. 5, 6.

Perennial. Plant robust with creeping rhizome covered with
numerous, thickish adventitious roots. Stem (0.7)1—2 m high, 8-12
mm thick at base, slender above, bent, sulcate, glabrous or with very
sparse hairs, but floccose below capitulum, with crisped glandular hairs.
Lower leaves withering much before anthesis, middle cauline leaves
20-35 cm long, 25-50 cm wide, triangular, higher ones deltoid-hastate,
apex rather shortly narrowed but often caudate, lateral lobes usually
bilobed, with long, acuminate secondary lobes and broadly sinuate
base decurrent on petiole, sinuate, sometimes irregularly toothed, teeth
abruptly narrowed into thin, acute apices, bright green above, somewhat
lighter, dull beneath, sparsely hairy on both sides with very short hairs;
petiole a third to a half as long as lamina, broadly winged(up to 1.5—
2.0 cm wide), at base expanded into large amplexicaul auricles; leaves
gradually reduced above, lower of the upper cauline leaves sometimes
of same form as middle cauline leaves but more often narrower (more
or less as long as wide), deltoid-hastate, with shallow-two-lobed or
undivided lateral lobes and toothed margin. Capitula numerous, up to
250-300, aggregated in large, 20-50 cm long, strongly branched
pyramidal paniculate inflorescence, lower branches long, up to 22 cm,
ascending, upper upward-spreading, all branched, with spreading
branchlets; lower bracteal leaves mostly remote, narrowly deltoid-
hastate or oblong, more or less large, upper leaves narrowly linear,
small, 2-5 mm long and 0.3—1.5 mm wide; peduncles of capitula usually
slender and long, 8—12 mm long or some shorter (4—6 mm long), borne
in axils of almost filiform bracteal leaves; 3.5 mm long, bearing 1-2
smaller leaves; main axis and peduncles covered with usually very
dense, short, curly, later partly arachnoid, glandular hairs. Involucre
narrowly cylindrical, of 5 involucral bracts; involucral bracts 9-12
mm long, at flowering three-fourths as long as pappus, linear-oblong,
green, sparsely glandular-pubescent on outside. Flowers in capitulum
(3)5-7; corolla 7-8 mm long, its tube 1.5—2.0 mm long, (1/4)2/7-1/3
as long as upper, narrow, almost tubular limb; anthers about 3 mm
long, with deltoid, 0.4—0.5 mm long apical appendages and 0.3-0.5 mm
long basal appendages; style branches long, hairy on outer side up to
upper one-third. Achenes 6-8 mm long, pappus white, about 7 mm long.

Dense mixed and coniferous, chiefly montane forests on slopes
and stony places in valleys of rivers and streams.—Far East: Ussuri
(southern region). General distribution: China (northeastern part),
Korea. Described from Changhua Tsailin Range between towns of
Omoso and Girin. Type in Leningrad.

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Note. Morphologically, C. komaroviana is closest to the Japanese
species C. tschonoskii Koidz. and is distinguished from the latter by
the following characters: broader leaves with a less attenuate middle
lobe and broad, rather deeply bifid lateral lobes, broadly winged petioles
expanded at the base into large auricles (rather than narrowed toward
the base and lacking auricles, or, in rare cases, with small auricles),
corolla with a longer tube and wider limb and a slightly yellowish
rather than snow-white pappus of the achenes.

Series 3. Auriculatae Pojark.—Inflorescence simple raceme or
panicle, on lateral branches capitula aggregated into simple raceme;
involucre narrowly cylindrical, consisting of (4)5 involucral bracts;
florets in capitulum (4)5—7(8); corolla tube as long or up to a half to
two thirds as long as campanulate limb; pappus of achenes white.
Plants 30-150 cm high, with reniform or deltoid reniform, up to 16(25)
cm wide leaves; petioles at base with auricles.

The following three eastern Asian species belong to this series.

4. C. praetermissa (Pojark.) Pojark. comb. nova.—Senecio
davuricus &. ochotensis Maxim. in Bull. Acad. Sc. Pétersb. XIX (1874)
485; Mél. biol. IX, 296, p. min. p.—Cacalia auriculata «. ochotensis
Kom. Fl. Manchzh. III (1907) 688 p. p.; Nakai, Fl. Korean. II, 35; in
Bull. Nat. Sc. Mus. XXXI, 114; Kom. and Alis. Opred. Rast. Dalnevost.
Kr. II, 1044 p. p.—C. auriculata var. alata Nakai, Veget. m. Waigalbon
(1916) 73; Fl. sylv. Korean. XIV (1923) 106.—C. auriculata auct. non
DC. (1837); Kitam. Compos. Japon. III (1942) 209 p. p. (quoad pl.
korean. and excl. synom.); Hara, Enum. Spermatoph. Japon. II (1952)
147 p. p. (quoad pl. korean.); Tolm. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XVIII, 240, p. min. p. (quoad pl. austro-ussur.).—
C. auriculata var. kamtschatica auct. non Matsum. (1895): Kitam. op.
cit. 210 p. p. (quoad pl. korean.) cum f. alata Kitam.; Hara, op. cit.
147 p. p.—Hasteola praetermissa Pojark. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XX (1960) 386.—Ic.: Pojarkova, op. cit. Figs. 7, 8.

Perennial. Rhizome oblique or horizontal, short, 5-15 mm thick,
densely covered with slender, filiform adventitious roots; stem 50-150
cm high, about 5(8) mm thick at base, gradually becoming thinner
above and 1-2 mm thick below inflorescence, sulcate-ribbed, stiff,
bent at nodes only upper part and often with very short, 1-3 cm long
internodes (middle internodes 8-15 cm long), densely leafy, particularly
in upper part, green or with red stripes, glabrous or with occasional
short hairs. Lower 1-2 leaves much smaller than middle cauline leaves,
reniform, with long petioles up to twice as long as lamina, withering
early; middle leaves 3-4, 7-15 cm long and (12)15-24 cm wide,

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strongly expanded, reniform or less often deltoid-reniform, usually with
deeply sinuate or almost hastate base, sinuate-toothed, often irregularly
toothed, apically usually abruptly narrowed into large acute tooth and
with 2(3) pairs of such teeth below, but sometimes with few teeth in
upper part as well (and in that case similar to leaves of C. kamtschatica);
bright- or dark-green above, lighter and glaucous beneath, on both
sides glabrous, matte; petioles mostly a fourth to a half as long as
lamina, less often 2/3 or up to 1/8 as long and along margin more or
less broadened, often broadly winged, up to 1.0-1.5(2.0) cm wide,
always with amplexicaul auricles, usually large; 1-2 upper cauline
leaves sharply reduced in size, 1/3—2/3 as long as middle leaves, deltoid
or reniform-deltoid, with truncate or cuneate base, usually with 3 large
teeth or almost 3-lobed. Capitula drooping, numerous, aggregated into
lax, less often compact, oblong, often almost cylindrical, paniculate
(only in weak plants almost racemose) inflorescence, 18-32 cm long
and 5-9 cm wide, branched almost up to apex, less often only in lower
half; branches (1.5)2—7 cm long, lower branches up to 10 cm long,
always shorter than main axis, obliquely upright, bracteal leaves (3)8-
20 mm long, lanceolate-linear, upper setaceous, only lowermost leaves
sometimes larger, resembling upper cauline leaves; peduncles of capitula
slender, 3—8(11) mm long, with 2—4 setaceous leaves 1.5—2.5 mm long.
Capitula with 5-8 florets. Involucre narrowly cylindrical; involucral
bracts 5, narrowly linear, green, less often reddish, (6)8—11 mm long
and 1.0-1.5(2.0) mm wide. Corolla at flowering time projects from
involucre by half the length of its limb, 8.5—9.5 mm long; corolla tube
as long as limb or up to 2/3 as long. Achenes (immature) 6 mm long,
brownish, pappus white, 7-8 mm long. Flowering August.

Mixed and coniferous forests in moist places, along banks of rivers
and streams, solitary or in groups, frequent.—Far East: Ussuri (southern
district). General distribution: China (Girin Province and Korea).
Described from the south of Primorsk [Pacific Coastal] Territory, from
Suchan River basin. Type in Leningrad.

Note. Besides the larger size of the plant and all its parts, C.
praetermissa differs from C. auriculata also by the following charac-
ters: a densely leafy stem, leaves that in shape and size are quite
similar to the leaves of C. kamtschatica, sometimes even developing
(like this species) several large, lobate, acute teeth on the upper margin,
almost always broadly winged petioles, involucral bracts that are longer
in absolute terms as well as in relation to the length of the corolla and
the pappus, and also by the paniculate inflorescence with many lateral
branches. Between the ranges of C. praetermissa and C. auriculata,
there is a discontinuity; our species is known in the north only as far
as Voroshilov and in the Primorsk [Pacific Coastal] Territory—as far
as Valentine Bay.

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5. C. auriculata DC. Prodr: VI (1837) 329; Ldb. Fl. Ross. II, 626
p. p. (excl. pl. kamtsch); Trautv. and Mey. Fl. ochot. 56; Maxim. Prim.
fl. amur. 165; Fr. Schmidt in Mém. Acad. Sc. Pétersb. VII, XII, 2, 51,
151 p. p.; Kitag. Lineam. fl. Manshur. 441 p. p.; Sugaw, IIl. fl. Saghal.
IV, 1845; Kitam. Compos. Japon. III, 209 p. p. (excl. area Korea et
var. kamtschatica); Tolmatch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVIII, 239 p. max. p.—Ligularia auriculata Turcz. in sched.;
DC. Prodr. VI (1837) 329, pro syn.—Senecio dahuricus Sch. Bip. in
Flora (1845) 499, non Fisch.—Senecio davuricus &. ochotensis Maxim.
in Bull. Acad. Pétersb. XIX (1874) 485 p. max. p.; Mél. biol. IX, 296,
p. max. p.—Cacalia auriculata a. ochotensis Kom. Fl. Manchzh. III
(1907) 688 p. p.; Kudo in Bot. Mag. Tokyo, XXIX, 224 (excl. area
Korea); Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1044 p. p.— _
Hasteola auriculata Pojark. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XX (1960) 391.—Ic.: Sugaw. Ill. fl. Saghal. IV, tab. 1844a;
Pojarkova, loc. cit. Fig. 10.—Exs.: Karo, Fl. amur. et zeaéns. No. 126.

Perennial. Rhizome creeping, oblique, short, densely covered with
slender, filiform adventitious roots; stem 30—100 cm (less often slightly
more) high, slender, 1.5—2.5(4.0) mm thick at base, ascending, often
paniculate, tapered above, very slender below inflorescence, sulcate,
glabrous, sparsely leafy because leaves up to a half as long as internodes
or as long sometimes slightly longer; cauline leaves 4-6, thin
chartaceous in herbarium, bright green above, lighter beneath, glaucous,
lustrous, glabrous or less often with sparse, short, thickish hairs; lower
1-2 leaves usually much smaller than upper, reniform, strongly
expanded, 2-4 cm long and 4.0—9.5 cm wide, abruptly narrowed into
long acuminate tip or sometimes emarginate, along margin sinuate and
often irregularly toothed, with long, slender, petioles up to 1.5—2(3)
times as long as lamina, expanded only at base and usually not forming
auricles; middle cauline leaves 5-10 cm long and 7—14(16) cm wide
reniform and up to twice as long as wide, to deltoid or roundish-
reniform, with deeply divided base or partly deltoid or roundish-deltoid,
with weakly notched or truncate base, often angular and then besides
terminal long acuminate large teeth, both sides with 1-2 similarly long-
lobular teeth in lower part of lamina or near its base, along margin
coarsely sinuate-toothed to indistinctly remotely toothed or partly entire
with slender petiole almost as long as lamina or a fourth to a half as
long, usually broadened only at base into small auricles (less often
auricles up to 1.5 cm long); upper leaves with short petiole, (1/8)1/6—
1/3 as long as lamina, sometimes resembling middle leaves but often
greatly reduced and different in shape, deltoid and oblong-ovate;
uppermost leaves often lanceolate, sometimes with undeveloped branch
of inflorescence or capitula in axils. Inflorescence cylindrical, very

696

663

lax, narrow, (3.5)4.5-15(23) cm long and 2-5 cm in dia, usually a
simple raceme, less often with few, short, 2.5—4.0 cm long branches in
lower part with raceme of few capitula. Capitula with 4—7 florets,
drooping, on slender peduncles, 1.5—5.0 mm, lower ones 7-9 mm long,
covered with very small capitate glands and very short, simple, straight
hairs. Bracteal leaves usually small (2 mm long), subulate or narrowly
lanceolate, only lowermost sometimes 1-3 cm long. Involucre narrowly
tubular, purple, or less often greenish-purple to green, at anthesis almost
a half as long as florets; involucral bracts 5, less often 4, oblong (4)5-—
8(9) mm long. Corolla 6-8 mm long, its tube sometimes as long as
limb but, most often, slightly shorter (less often up to half). Achenes
light-colored, yellowish, 3.5-5.0 mm long; pappus white, about 1.5
times as long as achene. Flowering second half of May to July; fruiting
September.

Dark coniferous, mixed and birch forests, mainly in valleys, coastal
scrub thickets, quite wet clearings, grassy areas, solitary or in groups.—
Far East: Okhotsk, Zeya-Bureya, Uda River Area, Ussuri (only more
northern part of Primorsk [Pacific Coastal] Territory, in the south up
to Olga Bay), Sakhalin. General distribution: China (northeastern part).
Described from Okhotsk Region. Type in Paris; duplicates of type in
Leningrad.

Note. C. auriculata should be dropped from the flora of Kamchatka
Peninsula. The specimens from Korf Bay, which Tolmatchev (op. cit.
240) was inclined to refer to this species, must be referred to C.
kamtschatica on the basis of their characters.

Cf. also Note 1 to C. hastata.

6. C. kamtschatica (Maxim.) Kudo. in Journ. Coll. agric. Hokkaido
univ. XII (1923) 60; Kom. Fl. Kamch. III,175; Sugaw. Ill. fl. Saghal.
IV, 1485; Vorob. in Tr. Dalnevost. Fil. Akad. Nauk SSSR, III(V),
75.—C. suaveolens auct. non L. (1753): Georgi, Beschr. Rs. Reichs.
Ill, 5 (1800) 1229.—C. auriculata auct. non DC. (1837): Ldb. FI.
Ross. II (1844-1846) 627 p. p. (quoad pl. kamtsch.); Fr. Schmidt. in
Mém. Acad. Sc. Pétersb. VII, XII, 2 (1868) 151 p. p.—Senecio
davuricus B. kamtschaticus Maxim. in Bull. Acad. Sc. Pétersb. XIX
(1874) 486; Mél. biol. IX, 196; Miyabe, Fl. Kuril. (1890) 244.—C.
auriculata var. kamtschatica Matsum. Shokubutsu Mei-i (1895) 56;
Kom. Fl. Manchzh. III, 688; Koidz. in Journ. Coll. Sc. univ. Tokyo,
XXVII, 121; Kudo, Fl. Param. 170; Kitam. Compos. Japon. III, 210 p.
p. (excl. pl. korean.); Hara, Enum. Spermatoph Japon. 147; Tatewaki
in Acta Hort. Gotoburg. XXI, 112.—C. auriculata Hultén, Fl. Kamtch.
IV (1930) 194, non DC. (1837); Fl. Aleut. 332.—Hasteola kamtschatica
Pojark. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX (1960)

697

664

391.—Ic.: Hultén, Fl. Kamtch. IV, Fig. 7a, b; Sugaw. loc. cit. tab.
844A; Pojarkova, loc. cit. Fig. 9.

Perennial. Rhizome up to | cm thick, creeping, short, sometimes
with 1-2 branches, densely covered with rather slender filiform,
adventitious roots; stem 60-120 cm high, at base 2-7 mm thick, stiff,
bent at nodes, ribbed-sulcate, up to inflorescence glabrous or at base
sparsely arachnoid-hairy, green or with violet-red stripes, usually
sparsely leafy because leaves roughly as long as internodes, less often
upper leaves up to 1.5-2 times as long; basal leaves scaly; lower
cauline leaf usually 1 (rarely 2), smaller than middle leaves, withering
long before flowering, with long petiole slightly broadened toward
base and lacking auricles; middle cauline leaves 3—4, up to 15 cm long
and 25 cm wide, reniform, deeply notched to almost truncate at base,
apically usually round (rarely emarginate), mostly strongly attenuate »
into large acute tooth, along margin usually with a few (5-8), similar,
large, sometimes lobed teeth separated from each other by several
smaller, acute unequal teeth, dark green and glabrous above, lighter
and glabrous beneath or along veins weakly crisped-hairy, matte,
chartaceous or white when dry, dense; petiole 10/13-1/2 as long as
lamina, slender, not winged very rarely expanded in upper part from
decurrent base of lamina, usually with small, about 0.5 mm long au-
ricles, less often auricles larger, up to 1 cm long; upper cauline leaves
very gradually reduced in size, often almost as long as middle leaf,
deltoid-reniform, deltoid or reniform lower 1-3 bracteal leaves similar
to upper cauline leaves but sometimes smaller. Capitula drooping,
numerous, aggregated in compact, branched panicle (only in weaker
plants almost racemose), oblong-ovoid, less often short-cylindrical, up
to 16-25 cm long, but mostly shorter, 8-12 cm long; inflorescence
branches ascending, almost parallel to main axis, not less than a half
as long, often almost reaching same level as of inflorescence with its
tip; peduncles of capitula somewhat thick, short, 1.5-3.0(5.0) mm long
(less often lower ones 7-10 mm long), 1/5—2/5 as long as involucre,
in axils of narrowly linear or almost filiform leaves exceeding them,
and with 1-2, less often 3-4, similar or smaller leaves; axil, and
peduncles usually densely covered with flexuous short or somewhat
long and then arachnoid hairs and very small glands. Involucre narrowly
cylindrical; involucral bracts (4)5, narrowly oblong to linear, lustrous,
pale green, (6)8—-11(12) mm long, up to flowering as long as corolla
and pappus, later about 2/3 as long. Capitulum 4—7 flowered, corolla
7-9 mm long, its tube 2/3 as long as limb; limb lobed up to 1/3-1/2
into narrowly deltoid, long-acuminate teeth. Achenes about 6 mm long,
finely sulcate; pappus almost as long as achenes. Flowering August to
first half of October; fruiting October.

69

oo

665

In thickets of alder (Alnaster fruticosus) and forests, mainly dark
coniferous forests in the valleys, in shady wet sites, in places frequent
to abundant.—Far East: Kamchatka, Commander Islands, Okhotsk(?),
Sakhalin, south and west and Kuril Islands. General distribution: Japan
(Hokkaido Island); western Aleutian Islands. Described from
Kamchatka. Lectotype in Leningrad.

Note. The report of C. kamtschatica on the continent south of
Kamchatka needs to be confirmed by new collections. In the Her-
barium of the Botanical Institute of the Academy of Sciences of the
USSR, there are only two old specimens from the Okhotsk Region:
one collected along the Urak River by I. Kuznetzov in 1835 and an-
other from the herbarium of Stubendorf.

GENUS 1562. Syneilesis Maxim.' ?

Maxim. Prim. fl. amur. (1859) 165, tab. VIII, fig. 9-18.—Senecio
sect. Cacalia Benth. in Benth. and Hook. f. Gen. pl. II (1873) 449 p.
p.—Cacalia auct. non L. (1753): Hoffman in Pflanzenfam. IV, 5 (1894)
296 p. p.

All flowers in capitula tubular, fertile, with regular white-reddish
corolla bearing 5 teeth; involucre narrow, usually with 5 unequal
involucral bracts, inner[outer] bracts broader and covering with their
edges much narrower outer[inner]; common receptacle flat, glabrous,
alveolate; anthers sagittate with short, acute, auriculate, fertile
appendage; antheropodium somewhat broader than filaments; style
branches long, at apex with deltoid acute appendage, depressed on
inner side, hairs outside in upper part. Achenes terete, monochromatic,
glabrous, with several fine ribs, with one longitudinally twisted, cordate,
slightly lobate cotyledon; pappus of numerous, fine, scabrous hairs.
Perennial herbs, with one basal, palmatisect, shield-shaped, long-
petiolate leaf, with lamina until opening inclined downward and folded
like closed umbrella. Capitula small, aggregated in corymbose or
paniculate inflorescence.

Types of genus: S. aconitifolia Maxim.

The genus includes five species distributed in the countries of
Southeast Asia: Korea, northeastern China, Japan, and Taiwan.

1. S. aconitifolia Maxim. Prim. fl. amur. (1859) 165; in Mél. biol.
IX, 299; Kitam. Compos. Japon. III, 170.—Cacalia aconitifolia Bge.

'Treatment by A.I. Pojarkova.
2From the Greek word syneileo—to roll up, because of the convolute cotyledons.

699

666

Enum. pl. Chin. bor. (1831) 37; Kom. Fl. Manchzh. III, 165; Nakai,
Fl. Korean. II, 34; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II,
1044.—Senecio aconitifolius Turcz. in Bull. Soc. Nat. Mosc. X (1837)
155.—Ic.: Maxim. Prim. fl. amur. tab. VIII, figs. 8-13; Kom. and
Alis. op. cit. Plates 312 and 313 (leaf-hand figure).—Exs.: Karo, PI.
amur. et Zecens. No. 68. -

Perennial. Rhizome short, woody, creeping, with thick adventitious
roots; stem 70-120 cm high, glabrous, finely sulcate, purple. Basal
leaf 1, withering before flowering; cauline leaves 2, remote, lower
with long, up to-30—50 cm long, glabrous, petiole, with short, expanded
base clasping stem, lamina roundish, coriaceous, glabrous but initially
floccose, 20-30 cm wide, divided up to base into 7-9 lobes deeply
incised into linear secondary lobes, with remote long, acute, teeth,
other leaf with shorter petiole and smaller lamina, with 4—5 lobes,
higher leaves sessile, lanceolate or linear, gradually reduced upward.
General inflorescence large, compressed, compound corymb, with lateral
branches bearing capitula only near apex; capitula numerous, crowded,
on 5-20 cm long peduncles, with subulate, short bracteal leaves at
base. Bracts at base of involucre, 3-4, narrow, linear, 3-5 mm long;
involucre cylindrical, later campanulate; involucral bracts 5, 9-12 mm
long, green or usually purple, glabrous, acute. Corolla about 10 mm
long, white-reddish, with narrowly campanulate limb, slightly longer
than tube; anthers usually purple. Achenes 5—6 mm long, narrow, terete,
sulcate; pappus twice as long as achene, whitish-rusty. Flowering second
half of July to August; fruiting first half of September.

Dry slopes, meadows with sandy soil, forest edges, pine groves.—
Far East: Zeya-Bureya (southeastern part), Ussuri. General distribution:
China (northeastern part), Korea. Described from Amur. Lectotype in
Leningrad.

GENUS 1563. Senecio L.!:?

L. Sp. pl. (1753) 866.

Capitula with single-rowed involucre having involucral bracts of
equal length or with one more row of usually much smaller bracts at
base, less often as long as inner bracts. Capitula solitary, few, or several
on stem forming corymbose, umbellate, paniculate or (less often)
racemose inflorescence. Receptacle glabrous, flat. Peripheral florets
pistillate, ligulate, one-rowed, yellow, orange, purple, or violet; often

‘Treatment by B. Schischkin.

*From the Latin word senex—old; the floral heads in many species are without
peripheral ligulate florets; therefore, they appear “bald.” As the flowers wither, the
receptacle becomes glabrous or “bald.”

700

667

ligulate florets absent, central florets (disk florets) bisexual, tubular,
their anthers obtuse at base; style branches truncate and bearing corona
of erect hairs near apex. Achenes terete, glabrous or pubescent, with
5-10 longitudinal fine ribs, apically obtuse, with many-rowed pappus
of white serrate hairs considerably longer than achene, very rarely
pappus lacking. Perennial, annual or biennial herbs (in the tropics
often as shrubs or trees) with simple pinnate or less often bipinnate or
ternate; or undivided, usually with toothed, alternate leaves.

The genus includes about two thousand species distributed
throughout the world, particularly in South Africa, the Mediterranean
floristic region, and in temperate Asia and America.

L..... Plante ‘annual very rarely biennial s:scs:5...civadale. cite ne. Pe
++). Plants peter lose. skin nits oleh ee see iM:
2. Plants glandular-pilose, sticky; achenes glabrous, less often weakly
pabesceatbendeenmiie sh.: isis. acca aiheosl.aeuesshenske.. 3:
+ Plants arachnoid-hairy or subglabrous, not glandular pilose; achenes
ieibads Ripiemeaiy cee iii lel ii ie, setae toed hd. 4.
3. Plants usually strongly branched; leaves often simple pinnate;
capitula 6-8 mm long; achenes glabrous .......... 92. S. viscosus L.

+ Plants usually weakly branched; leaves often twice pinnate; capitula
8-10 mm long, peduncles longer; achenes weakly pubescent and

glandular-pilose between ribs ................2645 93. S. calvertii Boiss.
4% ashipulete-fidtetsfentirely absent issiii.......scvsceiiisisstsethals.ieereaneeees 5:
+ Ligulate florets always present, numerous or few (two to three)
j aR Delis ape tebaS Mb Satin. Shan. tes phen ND. sot seen 6.

5. Leaves pinnately lobed, with broadly toothed lobes, 3—10 cm long
and 1.5-4.0 cm wide; achenes 2.0-2.5 mm long; plant 15-50 cm
high; capitula on short peduncles in compact corymbose panicle
Coil BUMS OOINUA BOE, Ey RWS cis eds eS Le 94. S. vulgaris L.

+ Leaves sinuate-toothed, less often pinnately lobed, 2-5 cm long
and 0.3-1.0 cm wide, with narrow (1-3 mm wide), mostly
undivided lobes; achenes 3.0—3.5 mm long; plants 3-20 cm high;
capitula on long peduncles in depanuperate inflorescence .........
SRL eR Mais ienies! ct iat. creel leet 95. S. dubius Ldb.

6. Ligules of peripheral florets very small, scarcely longer than
invelwene, an volute ya ai.9.. isc scsel i. CUI ON ad a ERd che saccade 7.

+ Ligules of peripheral florets flat, not involute, horizontally
spreading or decurved, considerably longer than involucre

7. Stem 25-70 cm high, branched from middle or only Beeies
involucre 6-8 mm long; ligulate florets 8-14 (European part of
Healt: 2. 08.0 LER Ei 96. S. sylvaticus L.

668

12.

13;

14.

15:

16.

Lilt

Stem 8-25 cm high, usually branched from middle; involucre4—
6 mm long; ligulate florets 2-7 (Pamiro-Alai Region and Tien
Shari). 62: AR. A 97. S. krascheninnicovii Schischk.
Short, 3-15 cm high, high-mountain plants, usually strongly arach-
noid-hairy, with numerous stems (Caucasus) ..........::s0000008
sR SERGI. 2D. LEONG. A 99. S. sosnovskyi Sof.
Taller, (S)10—60 cm-high plants, distributed in lower zones, usually
weakly pubescent, sometimes almost glabrous; stems 1-5 .......... 91
Outer involucral bracts turning black in upper part, or almost
entire; plants usually arachnoid-hairy; biennial ..............:cce08
Sieh. AEA LR BA 98. S. vernalis Waldst. and Kit.
Outer involucral bracts greenish at apex, lacking blackish spot;
plants-glabrous; annuals 5.2.05... G00 Re ST 19.
Leaves pinnate; plants growing on saline soils .............::c008
CR ee ETRE RI OI ee wh SO 102. S. noeanus Rupr.
Leaves broadly linear, undivided, with few teeth, sometimes al-

MOSPORT Ain wa BR 101. S. subdentatus Ldb.
Stem with single capitulum; usually short (5-25 cm high), alpine
OF UTChicrplaMts! eek. csc asckkk AeA. ee 12.
Steni with.3.to many capitulass:is. gol cum kd algae eae 2h:
Plants wholly:glabrousia:....00 ean ae aaa 13.
Plants more orless*pubescent:. ance ug Oh aa ee 14.
Lower leaves twice pinnate or ternate; outer involucral bracts 2—
5, linear; achenes 2.5 mm long .............. 52. S. carpathicus Herb.

Lower leaves lyrately pinnately divided, terminal lobe oblong with
obtusely toothed apices and smaller, entire, lateral lobes; involucre
one-rowed; achenes 3.5—4.0 mm long ...... 52. S. resedifolius Less.
Ligulate florets 8-10; basal leaves densely white-tomentose be-

PALA yo2i.2. SARTRE, en, Tig as, 91. S. jacuticus Schischk.
Ligulate florets 12-25; leaves more or less pubescent beneath, but
riot White-tomientosed..200s. Ae... LAMAR SOL 15.

Petioles of basal leaves usually 2-7 times as long as lamina;
upper half of stem often densely covered with long, soft, white

arachnoid hairs or dark purple hairs ...........:cccccsseeeesseeseeeeseeees 16.
Petioles of basal leaves shorter than lamina, less often as long
SBE, A LaITROTR os LATER Boas EL A eee 18.

Stem slender; leaves almost glabrous or sparsely pubescent
Loads iAH R LOA ast iL ee a ae See ae 87. S. subfrigidus Kom.
Stem thick, in upper half covered with thin felt of long, tomen-
tose, white or, sordid, brown Waits ceivsous ots aeaeaghce ye esas AV anon Li;
Ligulate florets dirty-purple; involucre 15 mm long and 25 mm
wide, upper half of stem covered with dense tomentum of sordid
hairs (Chukotka Peninsula) ......... 83. S. tichomirovii Schischk.

+

21(11).

702

24.

25s

26.

27.

669

Ligulate florets yellow; upper half of stem covered with dense

tomentum of soft white hairs .............cceeeeeee 84. S. pricei Simps.
Ligulate florets dark purple or dark orange (mountains of central
SSDs nde ss Mapes Ghai ltt. deca easdctn ach 85. S. turczaninovii DC.
hiselate Maretasy cllow:Or OfANge: souis.nsssonsess-.c-nsnsssnnandianss--cnevld 19,

Upper half of plant densely pubescent with violet hairs (Arctic)
snpalala ob tadlvebtadcalh heresy. 82. S. atropurpureus* (Ldb.) B. Fedtsch.
PRAMAS DV UNE TINRES CED: inn nnninicsnnennn-snenrnndnengctnivanns obleguesnthianbodesnsiexe 20.
Ligulate florets orange-yellow; achenes pubescent; plants 20-35
cm high (mountains of Kamchatka and Okhotsk) ..............:0e08+
Sr bereeradti nab tadwedscthy laces. 88. S. pseudoaurantiacus Kom.
Ligulate florets yellow or slightly orange; achenes glabrous; plant
19-20 cm high: bawwilissbividaauninns 85. S. turczaninovii DC.

Involucre two-rowed; outer involucral bracts few or numerous;
leaves often coarsely toothed, often lobate as well as simple pinnate
Gttwite pinnate. or temate walle. ake. samisctnslieleged...- 22:
Involucre one-rowed; leaves always undivided, along margin
coarsely or indistinctly toothed OF eNtire ...........ccscceseesseeeseeenee 76.
higulate florets wholly absent i:...snccssicvcissi..nssrebsaciligsbitgibest. adie 23.
Ligulate florets numerous; rarely 1-2 OF 3-5 0... cecseeeeeeees 26.

Leaves pinnate or lyrate; involucre 7 mm long and 2 mm wide;
inner involucral bracts much larger; capitula with numerous tubular

- florets; plants covered with thin, appressed arachnoid hairs

Dbicirtes 3S 05,513 pres this cabaeceanaseleatnct 38. S. paucilobus DC.
Leaves undivided, lanceolate or reniform-cordate, toothed;
involucre 4.5—-6.0 mm long and 2.0-2.5 mm in dia .............. 24.
Leaves lanceolate, 6-24 cm long and 1-5 cm wide; outer involucral
bracts narrowly linear, one-third to half as long as inner, latter 4—

GS swsnsesiei hr itso loan enc te 35. S. pojarkovae Schischk.
Leaves reniform-cordate, 14-15 cm long and 20-25 cm wide
sasubdlsnAaadishasnassiasianddonnnssesansannsisendbintibg> <ebeh Mit aB0sD se aeeai anes Mums eaCme, yo 25.

Capitula with 10-15 florets; involucre 8 mm long and 3 mm in
dia, inner involucral bracts 7-8; achenes terete, 4-5 mm long
hecal: eemeites Pea sanmeisbiatssiuhonss 37. S. platyphylloides Somm. and Lev.
Capitula with 5 florets; involucre 5 mm long and 2-3 mm in dia,
inner involucral bracts 4—5; achenes about 3 mm long
jasstan.aag Amaratasin actigmmaatls. tacit 36. S. rhombifolius (Willd.) Sch. Bip.
Ksigulate-flocetsyin capitulumy, 1--3(5): a rsidicseesssannadasiarssseneonenises Bis
Ligulate flonets Ge25 wusssscc...Q.oh. ates. hevebibes Wasa dee i.
Ligulate florets in capitulum 1-2; leaves quite large, 40-50 cm
long and 12—15(25) cm wide, simple pinnate or bipinnate; achenes
Ra ra Pe A ns sa snnncnrvnionnnannsncneris 39. S. othonnae MB.

. atripurpureus in the original.—General Editor.

670

29(26).

+
30.

33;

703
34.

35.

36.

a7

+

38(31).

Ligulate florets 3-5; leaves 5-10 cm long and 1-3 cm wide;
achenés glabrous visere.ageess RRR A PAR. ad 28.
Leaves deeply pinnately divided; inner involucral bracts numerous,
with dark violet, almost black, spot at apex (Carpathians)
steuvenivetoenes tan wpenidbeveshReeUN IE AR ER 49. S. carniolicus Willd.
Leaves ovate, almost entire; inner involucral bracts 8, lacking
dark violet spot at apex (Soviet Central Asia) .........ccccseeseeeseees
saves dnd dana heA ten RIOR yahtennas wanna an 54. S. karelinioides Wink.
Leaves undivided, usually toothed, less often entire ............... 30.
Leaves pinnate, sometimes bipinnate, rarely lyrate ............... 56.
Leaves with oblong or suborbicular auricles, irregularly toothed
with acute, upright teeth «0.0.0.0... 48. S. litvinovii Schischk.
Leaves without or with weakly developed auricles, usually glabrous
beneath; withperect teeth .ui.08ud REG e St.
Eero ate Morets SGG) (00. OR, NR RE See 32%
Ligulate florets,not,less. than S)04.22, Wane 38.
Capitula with 15-20 florets; involucre obconical, 8 mm long and
4.mm in:dia(Caticasts) «2006.04. Baas 43. S. lapsanoides DC.
Capitula with numerous florets; involucre broadly campanulate,
5—6 mm long and 3.5—6.0 mm in dia ...... eee eeeeeeeeceeeeneeees 33)
Root filiform, stem about 50 cm high; leaves 3-10 cm long and
l-Svemnwide Ae WRAL. SR ee 34.
Rhizome slender, creeping; stem 60-100 cm high; leaves 15-30
emlongand)4—S—8:5..6my Wide -s.csss0sdndoeeme.s 35:
Ligulate florets only slightly longer than involucre; cauline leaves
lanceolate; upito.2.d Gm: Wide 2.2.28. 2e ects sihe ah RA Lee ae
eben etek. 7. S. saposhnikovii Krasch. and Schipcz.
Ligulate florets 2—2.5 times as long as involucre; cauline leaves
ovate ‘oreblong;, up to. 5..cm. Wide. ......0.0..1.Rsaetentel Se
La abyadabte cat RRA UEE OR etl Reinet 44. S. quincueligulatus Winkl.
Leaves glabrous: on: both Sides icssisesisseesssvewevereneorencasnecabinienceasan 36.
Leaves short-pubescent along veins beneath or all over .......... 37.
Stem more or less flexuous; outer involucral bracts 56, scarcely
shorter or longer than inner bracts; achenes 5 mm long ............
JM bets vs BRR ERL LO i DEAL, 45. S. fuchsii K. Gmel.
Stem not flexuous; outer involucral bracts a fourth to a third as
long as inner; achenes 3-4 mm ONG .........ceescseesseeeseceseeereeees
ovens appre psporishagunnnantesbahnst deat eanea ee eR 30. S. schvetzovii Korsh.
Outer involucral bracts 4—6, longer than inner; achenes 4.0—-4.5
mm long (Caucasus) ..........:.scceseeeeeees 42. S. propinquus Schischk.
Outer involucral bracts 1-3, a third to half as long as inner; achenes
Simtel he). anaijeni wena 26. S. macrophyllus MB.
Ovaries, and.achenés pubescent .......3.:...:4.-s84dshsGateetioboeeenannans 39.

704

ao:

43.

44.

45.

48(44).

Qvaribs:and achenes glabrous’ s...2535:i)..i82055 001. 1.00b ea 41.
Outer involucral bracts linear, 12-15, a third to half as long as
inner; achenes about 5 mm long, hairy only along ribs ..............
eR eG sstcstiasisccsniven osvnsisdovensvoovves 29. S. paucifolius Gmel.
Outer involucral bracts 4-5; achenes about 7 mm long, entirely
PUMACRERNNE). Cae es titebsers itis. ethnic ie OBO 40.
Leaf lamina at base truncate or cordate, often pinnately divided at
base, with remote lobes ............... 23. S. olgae Rgl. and Schmalh.
Leaf lamina at base cuneately narrowed, never pinnately divided
Davee nck Bheaenl.ceee. bias. aaie..2 24. S. franchetii Winkl.
Achenes:6=Jimmilong, lustrous .i0ci.se.cbiiiee 28d laid essences 42.
Achenes\2:0-4.5 maviong, dull. :..22.:.2scc00..0cs0l asad. Abe 43.
Stem fistulose; leaves green above, densely arachnoid-hairy below,
whitish, at base with semiorbicular, indistinctly toothed auricles;
involucre broadly campanulate, 10-12 mm long and 20-25 mm in
diai(Far East) ss.caw.!...casis 15. S. pseudoarnica Less.

Stem solid; leaves without arachnoid-tomentum below, lacking
auricles, involucre 15 mm long and 20 mm in dia; ligulate florets
BROT seesecrribatats obs LE caiesccassovivenseeee 25. S. paulsenii O. Hoffm.
High-mountain plants, with stems 5-20 cm high, arachnoid-hairy;
leaves small with petioles 1.5—4.0 cm long and 0.6-1.2 cm thick,
several, times:longessthan) lamina »....).2)..0:0) Setaalauilil..as......
atch Serle. Materia c 2 50. S. tianschanicus Rgl. and Schmalh.
Piants 25-150 cm high, with much larger leaves; petioles shorter
than lamina, less often as long or somewhat longer ............... 44.
Capitula small; involucre 5-8 mm long and 4—7 mm in dia ..... 45.
Capitula larger; involucre 8-12 mm long and 10-15 mm in dia

Leaves sessile, sharply toothed, with teeth directed upward; inner
involucral bracts with blackish spot at tip .........cccccceceeeeeeee
hadplidddcl. « Sppaotemes teaedl ds dics Tegel EEE dled 46. S. fluviatilis Wallr.
Leaves, at least lower, more or less long-petiolate, with straight
teeth; inner involucral bracts lacking blackish spot at tip ......... 46.
Teeth along leaf margin almost absent; outer involucral bracts 2—
4; achenes 2—3 mm long ................. 32. S. racemulifer Pavl.

Teeth along leaf margin distinct; outer involucral bracts 4—5, as
long as inner; achenes 3.0-4.5 mm long ............:.c:ce:esseseeeeeeee 47.
Teeth along leaf margin small, acute, equal; peduncles glabrous
or weakly arachnoid-hairy ..................0000 40. S. nemorensis L.
Teeth on leaf margin large, apically obtuse or roundish, not equal;
peduncles densely white-pubescent ............ 41. S. iljinii Schischk.
GCM MISO, Boies cciccte ee oakoosscsntarscnecdovvessssenteenWheatel vine 49.
Biigpemiae TRG U7 ea ye aks enccdeceeocsadaccsshsoccceberence at

705

672

50.

Sh.

52.

59:

56(29).

Involucre 12 mm long; outer involucral bracts 8-10, linear .or
lanceolate, a half as long as inner or slightly shorter, straight;
lamina abruptly or gradually narrowed at base into petiole ......
REA VSR eR EA Te nD 20. S. buschianum Sosn.
Involucre 8 mm long; outer involucral bracts 8-10, linear, usually
decurved, a half as long as inner; marsh plants, with hollow, densely
leafy stem; leaves narrowly lanceolate, sessile, 10-15 mm long
and 1-3 mm wide, sharply toothed with upward-directed teeth

JIRA TELAT KER DN RTT, Aol 50.
Mountain plants with solid stem; leaves ovate or lanceolate, 2.5—
10.0 cm wide, toothed, teeth not directed upward ................. Sh.

Leaves dark green above, lighter beneath and sparsely pubescent
ladinialiqumeennl cvni. rere esha 33. S. paludosus L.
Leaves green above, floccose white-pubescent beneath ..........
ALR cL. LRA 34. S. tataricus Less.
Stem densely pubescent below. Leaves large; basal and lower
cauline leaves ovate, with lamina 25-30 cm long and 10 cm wide;
blackish spot at tip of involucral bracts absent ............csseee
AEROS RE isi itei lia ssusica codes 31. S. umbrosus Waldst. and Kit.

Stem glabrous below or even weakly arachnoid-hairy ............ 52.
Outer and inner involucral bracts with blackish spot at tip ...... 54
Involucral bracts lacking blackish spot at tip ..........cceeseeseeees 54.

Leaves oblong, 1.5—3.0 cm wide, almost entire; outer involucral
bracts 5—7, linear, almost as long as inner; peduncles subglabrous
onde CATR 2 9h: 21. S. pseudoorientalis Schischk.
Leaves ovate-lanceolate, 2.5—5.0 cm wide, toothed; outer involucral
bracts 8-10, linear or lanceolate, a third to a half as long as inner
eis dedonnst taeda asb Parone SUT ets stab suoies 22. S. kolenatianus C.A.M.
Ligulate florets 12; lamina at base sometimes more or less cordate
(Carpathians)!..14. 4008. aa, 19. S. subalpinus Koch
Ligulate florets 8—10; lamina at base narrowed ............::ccesee 55.
Inflorescence racemose; leaves subglabrous above; pappus white
(Glavnyi)[Main]*Caucasus! Range) noone. De
NITRILE GORA: LOE NOIRE 27. S. racemosus (MB.) DC.
Inflorescence paniculate, leaves pilose above, particularly along
veins; pappus brownish (Transcaucasia) ..........:ccccsssccssssseeseeeeeees
stadt ao VERE ce AQAA, 8 28. S. thyrsophorus C. Koch
Leaves twice pinnate or ternate with divergent, linear or oblong
terminal lobes. 6h sciciteisd ge ARIMA RRR bis a7.
Leaves simple pinnate, sometimes lyrate .............:csscceesceeseeeees 59.
Achenes 3-4 mm long, all densely hispid (Caucasus) ............::cs+s+00+
4 kis an aseacuiveraed ceusiened tha iinenieatitewaiin 10. S. schischkinianus Sof.

706

61.

62.

63.

64.

65(9).

673

Achenes 1.5—2.5 mm long, all glabrous; or peripheral achenes
glabrous and those of disk florets pilose .............cc:cceeseeeeeeessees 58.
All achenes glabrous; leaves twice pinnately incised (Soviet Cen-
WBA SID ha asicde Teswyerciswessmvonnnremigs 10a. S. ferganensis Schischk.
Peripheral achenes glabrous; achenes of disk florets pilose, leaves
thrice pinnately incised ............. eee 9. S. borysthenicus Andrz.
Achenes pilose, peripheral ones sometimes glabrous ............... 60.
alia ones *plabrous "26.6.5, 2/2. Ue IRS HALE ones 65.
All achenes pilose and with pappus; leaf lobes long and acute;
PACS CHAIRS dase. nieeseier...cnpmads eee auenieeinetus 61.
Peripheral achenes glabrous, lacking pappus; pappus of ligulate
florets consisting of few hairs on ovary, detaching readily, leaf
lobes short and obtuse or subacute; creeping rhizome absent

Leaves densely hairy beneath, often grayish-tomentose, paasek
lobed or usually deeply pinnately divided ...............ecsceeeeseeeseeeeees
teviereeehtaapatangcostbeeenes dyeveaedeconseersarseetiss 2. S. grandidentatus Ldb.
Leaves glabrous or weakly tomentose beneath, deeply pinnately
GINA DORR ied bs PAN be TAGs Ler fants YL hka tlh tro Hive. senihed 62.
Stem 20-30 cm high; leaf lobes linear, cartilaginously thickened
along margin; achenes 3 mm long (Transcaucasia) ............
PUA, Mec desis hnestecpvasyrebenessscsvennsvases 100. S. polycephalus Ldb.
Stem 40-100 cm high; leaf lobes linear-lanceolate, lacking
cartilaginously thickened margin, long and acute; achenes 2 mm
DEVAS cotceoxaani hove RIMUIRD. GIR, £8... ak! 1. S. erucifolius L.
Ligulate florets 8—10(12), small, 4-5 mm long and about 2 mm
wide; plants divaricately branched from middle of stem ...............

MOA AA RAAT Shs SE Leader dh 8. S. erraticus Bertol.
Ligulate florets (10)12—25, longer (6-15 mm long) and broader
(2540. tittn Site 4. Wa leNON8..686i. UR LR Bla 64.

Capitula large, together with ligulate florets 2-3 cm in dia; ligu-
late florets 12-15 mm long, usually 15-30, twice as long as in-
GODS LR, ie Avr kee, RON SAL QU 5. S. ambraceus
Turcz.

Capitula 1.5—2.0 cm in dia; ligulate florets 6-10(12) mm long,
12—13(15), slightly or 1.5 times as long as involucre ..............

YOK). COnl sm.2...2voriels coins. OL ala, 4. S. jacobaea L.
Peg cs i a eee o'er 66.
Beialate: Moretet B54 ih: Bhilai elds WOE aetin 67.

Achenes 5-6 mm long; leaves pinnatifid, leaf lobes with upward-
directed teeth; plants 1-2 m high (Far East) .........cseeeeeees
Grerrsas PIA MGA eh eels 47. S. cannabifolius Less.
Achenes 3 mm long, teeth on leaves not upward directed; plants

707

674

67.

69.

70.

rey

20-60 cm high ............... 7. S. saposhnikovii Krasch. and Schipcz.
Ligulate florets 8, whole plant white-tomentose except upper sur-
face of leaves; achenes 1.5 mm long (Crimea, cultivated)
~ ae aR NE nian nspsnnnssstnnocneaesieeheiee 11. S. cineraria DC.
Ligulate florétsl0—15 sdi.s. csncsudkely. apmacion. levee cia 68.
Leaves lyrate or panduriform, with 2-5 pairs of deltoid or broadly
ovate, sharply toothed lobes, densely white-pubescent beneath
(rarely lacking tomentum of glandular hairs); terminal lobe usu-
ally cordate of owate at: base :.:::..o.08. cinoma ILA... eo
69.

Leaves not lyrate, pinnatifid and not pubescent beneath .......... 74.
Whole plant short glandular-hairy, disk florets 4-toothed ............
octyl, smal fanmemte. nates! i808 18. S. taraxacifolius (MB.) DC.
Plants arachnoid-hairy, lacking glandular hairs; leaves usually
white-tomentose beneath:....i:s:..0d.cued.“lnandd. aompsiom dee 70.
Terminal lobe of lyrately pinnate leaf ovate-deltoid, cordate at
Dyan ean onsen cianpneirncesi pense ermmc ehaeeeanaauanls tee nee e hd.
Terminal lobe of leaf truncate at base, not cordate ............... rex
Capitula 2-6; involucre 15 mm in dia (Ciscaucasia and Western
Transcaneasia)ic.:2psiyi!. 2acia!. teak. 17. S. pandurifolius C. Koch
Capitula numerous; involucre 10-12 mm in dia ..............eeeeeeee
mt cpp int 2 AI ds cn ckedassnksngtaneueeeresaies 16. S. kubensis Grossh.
Involucre 10 mm in dia; capitula numerous, 10-40 ........ 0. eee
aye opti. pe eet! weeeakes, dene 14. S. massagetovii Schischk.
Involucre 12-15 mm in dia; capitula 2-25 ooo... eeeseeseeeeeees 13.
Involucre 8-10 mm long; outer involucral bracts 1-3 ...............
Mg dt lite eet baton lean teed 12. S. lorentii Hochst.
Capitula up to 25. Involucre 5-6 mm long; outer involucral bracts
De] Lsicvcaky Spin fi beetiiewsiters teat tle ted As sates 13. S. lipskyi Lom.
Middle cauline leaves obovate or ovate, 8 cm long and 4-5 cm
wide; outer involucral bracts linear-subulate, about 5; ligulate flo-
Ptsyl Doris Maviabeastlla ties pepanenl cases ist 51. S. renardii Wink.
Middle cauline leaves oblong, 8-20 cm long and 0.54.0 mm
wide, pinnatifid; outer involucral bracts 2—3, narrowly linear, half
as long asidnnenal...sszliusk..2ah .aianie-Qewke! nleleeeahhu ate 75.
Inner involucral bracts with quite distinct blackish spot at tip;
ligulate florets 10, achenes glabrous, 4 mm long (Soviet Central
LACIE) ee aeee, Celene mee eee 3. S. korshinskyi Krasch.
Inner involucral bracts lacking blackish spot at tip; ligulate florets
12, yellow; achenes glabrous, 2 mm long. Involucre broadly cam-
panulate, 5 mm long, 6-7 mm in dia; outer involucral bracts
linear, 10, considerably shorter than inner bracts ..........:::csseeseeees
iit. brine, dot eons 2G atest anaes 6. S. argunensis Turcz.

708

76(21).

84.

85.

675

Stem branched in upper half, usually densely glandular through-
out with divergent, multicellular hairs; stem hollow; ligulate flo-
rets light yellow, numerous (20-25); pappus at anthesis scarcely
longer than disk florets, strongly elongated at fruiting ....................
eh Ab ah po pL ORR Le Ae oS eeaeicus. Rupr.
Stem usually solid (less often hollow); pubescence otherwise; ligu-
late florets yellow, usually less numerous; pappus not elongated at
MRUUNURENE a8 oi Nc cv ches cere cseusvedevecsscsusseteneectu nue cvsente okt ned neanuaaagte aa
Ligulate florets flame-red; high-mountain plants ...............6+++ 78.
Ligulate florets dark-purple, orange, yellow, or lemon-yellow
SMO teas eR ass scat as ctsc adh ak ece SEO Oren Dares 80.
Involucre 6-7 mm long and 8-10 mm in dia; achenes pubescent,
less Often glabrous 2.2... 79. S. caucasigenus Schischk.
Involucre 10 mm long and 10-15 mm in dia; achenes glabrous
EL ee Acs BA RE BE ATI A RO sacosenve 79.
Stem (25)30—40 cm high; basal and lower cauline leaves sessile
or with very short broad petiole (Carpathians) ............::e
yA a pee SRE heed ee 78. S. capitatus (Wahlb.) Sten.
Stem 15-30 cm high; basal and lower cauline leaves with petioles
as long as lamina or longer (Dzhungarian Alatau) ..................
RT yrnsseassiataanatarataat ai i88885 5548 80. S. pyroglossus Kar. and Kir.
Ligulate florets dark purple ..............0:ccssscssesssssenecsestecereenesesenees 81.
Ligulate florets orange, yellow, or lemon-yellow ...........::0 83.
Ligulate florets linear, 15-20 mm long and 1-2 mm wide ............
Brera eee OTs SCT lute tacs ote vucuranete 76. S. flammeus Turcz.

WY AEN ATES EN AT BLE ER Nia, oct 82.
Achenes pubescent; basal leaves 1-2(3) cm wide (Western
[od ig oe oy eo Re Ree EE 77. S. besserianus Mind.
Achenes glabrous; basal leaves broader, 2-5 cm wide (Eastern
SIGCHI) Heer reteccecees cadet ites recteseeN 75. S. porphyranthus Schischk.
Arctic or alpine plants, 10-30 cm high; capitula 2—5, rarely more,
ligulate florets Often OTaNge ............:.seccseeeeeteeeeteeeretseeteeeeeeees 84.
Taller plants, 30-100 cm high, usually with many capitula; ligulate
floret yelOW si.cccscc.cccceccscseccsescsscsssessssssssessosssssessenencsessosesensnensncsnene 86.
Ovaries and achenes pubescent; arctic plants ............scssessesessees
so ecco Ds AE Fi foe eee ee a 86. S. tundricola Tolm.
Ovaries and achenes glabrous; alpine plants .............::::sese00 85.
Rhizome rather thick and long, creeping; stem in upper part and
peduncles tomentose-villous (Altai) .........:.:sssesseeseeseeeeeteeerees
Lipa bite eae hi. aa nla ieee apa 29. S. sumneviczii Schischk. and Serg.
Capitula small; stem in upper part villous-glandular mixed with
white [and/or] dark-purple hairs (mountains of Central Siberia.

709

676

88.

89.

sive lub Bei ebay nlataeeedhcit asians. eee 85. S. turczaninovii DC.
Basal and lower cauline leaves with petioles longer than lamina;
lamina cordate at basescis%: ».26-<dk ied bho eae 87.
Lamina attenuate toward base (not cordate) ............cc:cceceeeeeee 88.
Involucre 10 mm long and almost as much in dia; achenes glabrous
(Western, Ukraine); stem, not; hollow. 3.2..04.<sskisicch os. bea
ci bah itawy cages td Rd ad ls cbc lve 72. S. rivularis Waldst. and Kit.
Involucre 6-7 mm long and 7-8 mm in dia; achenes hairy; stem
hollow: ayes Sheets AEH dereorens 62. S. cladobotrys Lab.
Basal and lower cauline leaves roundish-ovate, or broadly ovate,
with Lange! teeth. singh och ce Pei a ee 89.
Basal and lower cauline leaves oblong, ovate, elliptical, with small,
incanspievous teeth or almost entive %..isgisitoain tee 90.
Inflorescence umbellate; capitula on peduncles 2.5-4.0 cm long;
involucral bracts dark violet, wtih blackish tips (Far East)
stub eliediiel Heit naiteecelnel 2 70. S. sichotensis Kom.
Inflorescence corymbose, on rather unequal peduncles, 0.53.5
cm long; involucral bracts greenish-brown, lacking blackish spot

at tip (Altai)... 71. S. veresczaginii Schischk. and Serg.
Achenes\ pl abiiouis giisctesishs..crocegsendss ap teeth ee ae A D1,
Aghenes eben cen’ ifs 5.03.2i45 5h accectege SRI Re oh TIO ouaneinnseinannes 103.
Achenes,2:0-2.5 mam: Lomp: ..:,3issces-adsnnis. dedenasste Seeeceeeeae oe 92.
Achenes 35520 mani long jp ctieiie cangratie tte taal ot pe i eee 94.

Involucral bracts densely covered with grayish felt; involucre 10—
12 mm long and 12-15 mm in dia. Ligulate florets yellow, 3-4
mmiwide ws. 9h. Melee: Bees ee 68. S. aguilonaris Schischk.
Involucral bracts lacking grayish felt ...............:ccssccessesseseesreees 93.
Capitula small; involucre 4-5 mm in dia; stem hollow; ligulate
10) (loi i 9p aR PR 67. S. sukaczevii Schischk.
Capitula larger; involucre 8-10 mm in dia; ligulate florets 12-20
cpinsoa tterdoyes ean aR ew cae bakad 66. S. asiaticus Schischk. and Serg.
Ligulate florets often orange, 20—25; achenes 5 mm long ............
fis soa ee. dee ine cabal A ae de 81. S. karjaginii Sof.
Ligulate florets yellow, 5-15; achenes not longer than 4 mm
pin ip MUe Saie MARYS sales ens epinsln aineacis phcsuetaniSmeandaca ce ages Raeae Mecca ne 95.
Plant 20-30 cm high, densely tomentose; ligulate florets 10-15
mm long (Sakhalin), .............s0ssessesee. 65. S. kawakamii Makino.
Plants much taller and subglabrous ............:cssccesssseesreeeeseseenees 96.
Ligulate florets. epe-yellow: (Unals)) :.ss5issasteuz. denenpcndtaeinsds Anarene

Ligulate florets. orange.oryellow i. -chikc ane earns cae 97.
Involucral bracts blackish-violet; ligulate florets orange ............
Jeihaiiaicaincbane~ casan sei asieahed Berne mer gies 90. S. reverdattoi Sobolevsk.

710

98.

100.

677

Involucre 4 mm long and 6 mm in dia; involucral bracts 12, with
lang;aeute, ‘recurved, and. blackish ¢ipyg: 200552 20, wtsesledssayi onsets
SR ES --Reerrnsnt hn aie othe sees 73. S. hieraciiformis Kom.
Involucre 8-10 mm long and about 15 mm in dia; involucral
bracts 20-30, lacking blackish tips ...........ccccscesesseseeeeseeeenes 99.
Stem almost leafless, 15-30 cm high; ligulate florets about 10;
achenes glabrous, 4-5 mm long ........... 74. S. subscaposus Kom.
Stem leafy, ligulate florets 12-15; achenes 34 mm long .......
Bales ghee td tides «ondv ects Gan deactnde ees ta apsyantd geet tabegh Sos Leena 100.
Achenes hairy; involucre 8 mm long and as much wide; ligulate
florets 12—14 (Caucasus) ............... 63. S. subfloccosus Schischk.
Achenes glabrous; involucre 8-10 mm long and about 15 mm
wide; ligulate florets 15 (Carpathians) ................:cccsesscesseeseeeeess
SER Me atte ecnclia ls idguvetvaces tedesaensrinhadts 64. S. papposus (Rchb.) Less.
Stem hollow; involucral bracts with dark spot at tip ...............
SE a ee Be oe ae ee ee ae 61. S. lenensis Schischk.
Stem solid; involucral bracts lacking dark spot at tip .......... 102.
Basal and lower cauline leaves with petioles shorter than or as
long as lamina, rarely slightly longer ..............:ccccscesceseeseeseeeees 103.
Basal and lower cauline leaves with petioles 2-3 times as long as
LANIAS gk -<.i2 a eadlenesite- mninignd stheginh- tee ane 104.
Plants floccose-tomentose, later subglabrous, capitula 7-20 (Siberia
and northern European part of the USSR) ..........ccccccssecsecesseeeeeeees
brpbiie cient, rigs aghgchiedachans ssesssseeseeeeeee OO. S. Campester (Retz.)
Plants with dense, thin, white tomentum on stem and leaves;
SABIE a BCL iach, pct 0s J 505055020 ih ehinsras dae eed deagpaee Mees 105.
Leaves with remote, very small, blackish teeth; achenes 2.5 mm
Fei Se Pa oy ee, 59. S. amurensis Schischk.
Leaves lacking very small blackish teeth; achenes 3 mm long
ib Gide re ha dissdghl.-.cyete dd. daachdin diene 58. S. jalikola Juz.
Plants subglabrous (Kamchatka); capitula 3; involucral bracts 6—

7 mm long; achenes 4 mm long ........... 60. S. succisifolius Kom.
Plants pubescent; capitula 5-15; involucral bracts 4.5—5.5 mm
lonf;achenes 3 mm long’............0........... 57. S. czernjaevii Mind.

Section 1. Jacobaea DC. Prodr. VI (1837) 348; Boiss. FI. or III,
384; Gord. and Gr. FI. Fr. II, 112.—Plants perennial or biennial; leaves
pinnatisect or pinnatifid, or even lyrate, less often twice pinnately cut,
glabrous or tomentose; ligulate florets always present, 10—20, less often
fewer, yellow; involucre 2 rowed, outer involucral bracts usually few.

Series 1. Erucifolii (Rouy) Schischk. comb. n.—Subsect.
Erucifolius Rouy, Fl. Fr. VIII (1903) 335.—Leaves simple pinnate or
lyrate, glabrous or weakly pubescent.

711

678

1. S. erucifolius L. Sp. pl. (1753) 869; Ldb. Fl. Ross. II, 633;
Boiss. Fl. or. III, 351 ex parte; Schmalh. Fl. II, 87, var. excl.; Grossh.
Fl. Kavk. IV, 154; Kryl. Fl. Zap. Sib. XI, 2842.—S. jacobaea Ldb. FI.
Alt. IV, 110, non L.—S. praealtus Ldb. Fl. Ross. I, 634, p. min. p.
non Bertol.—S. erucifolius var. viridis Trautv. in Mélang. biol. II (1854)
130.—Jacobaea erucifolius Gaertn. May. and Scherb. Fl. Wett. III
(1801-1802) 208.—Ic.: Rchb. Ic. Fl. Germ. (1853) t. 966; Syreistsch.
Ill. Fl. Mosk. Gub. III, 284.—Exs.: GRF No. 178.

Perennial. Rhizome creeping, stem erect, glabrous or weakly
tomentose, weakly branched above, sometimes reddish, 40-100 cm
high. Basal and lower cauline leaves withering early, glabrous or weakly
pubescent, petiolate, lyrately pinnately cut; middle cauline leaves ovate,
5-13 cm long and 2.5-7.0(10.0) cm wide, pinnately cut, with lanceolate
or linear, obliquely upright, acuminate lobes; lobes entire or toothed,
or divided into few lobes. Capitula numerous, in corymbose
inflorescence. Involucre campanulate, 4-5 mm in dia, glabrous or
weakly tomentose; outer involucral bracts 5-6, subulate, a half as long
as lanceolate inner bracts; ligulate florets yellow, 10-15, oblong, 10
mm long and 2 mm wide; achenes about 2 mm long, ribbed, oblong-
terete, hairy in both disk and peripheral florets; all achenes with pappus.
Flowering June to August; fruiting September.

Forest edges in birch groves, pine woodlands, birch-aspen forests,
scrubs, wet floodplain meadows, gravel beds, alkaline steppes, dry
slopes, banks of irrigation canals, less often in standing grain crops.—
European part: Dvina-Pechora, Baltic Region, Upper Dnieper, Upper
Volga, Volga-Don, Volga-Kama, Trans-Volga, Lower Volga, Black
Sea Region, Bessarabia, Upper Dniester; Caucasus: Ciscaucasia,
Dagestan; Western Siberia: All regions; Eastern Siberia: Angara-
Sayans? Dauria? Lena-Kolyma; Soviet Central Asia: Aralo-Caspian
Region (north), Lake Balkhash Region, Dzhungaria-Tarbagatai, Tien
Shan, Syr-Darya. General distribution: Scandinavia (southern), Altantic
Europe, Central Europe, Balkans-Asia Minor (north), Mongolia.
Described from Western Europe. Type in London.

2. S. grandidentatus Ldb. FI. Ross. II (1844-1846) 636; Grossh.
Fl. Kavk. IV, 154.—S. arenarius MB. in Bess. Cat. Hort. Crem. (1816)
129, non Thunb. Prodr. pl. Capens (1800) 158.—S. erucifolius a.
canescens Trautv. in Mélang. biol. II (1858) 130.—?S. jacobaea L.
farinosus Lindem. FI. Elisabethgr. (1867) 183; Prodr. Fl. chersonens.
113.—S. erucifolius B. latifolius and y. grandidentatus Boiss. FI. or.
III (1875) 392.—S. erucifolius var. mollis Trautv. in Tr. Peterb. Bot.
Sada, X (1877) 117.—S. ponticus Grecescu. Suppl. Fl. Romanei (1904)
199.—S. mollis auct. Fl. cauc. non Willd.: Grossh. Fl. Kavk. IV, 154.—
Exs.: GRF No. 1779; Pl. orient. exs. No. 247.

71S

aS,
SZ 4 /
; 7 a\

Z—
<i S

Plate XXXIII.
1 — Senecio grandidentatus Ldb., habit; 2 — S. renardii Winkl., habit.

679

WAZ

680

Perennial. Rhizome short; stem simple, erect, finely ribbed, floc-
cose-arachnoid-hairy, 30-100 cm high. Basal and lower cauline leaves
withering early, middle cauline leaves ovate, deeply lobed or coarsely
toothed with acute lobes and teeth, weakly arachnoid-hairy-floccose
above, more densely hairy and grayish beneath, with small auricles, S—
10 cm long and 2.5-4.5 cm wide; upper leaves smaller, deeply di-
vided. Capitula numerous, in corymbose inflorescence. Involucre 5
mm long and almost as much in dia; inner involucral bracts ovate-
lanceolate, with wide, white border, acute or acuminate, weakly pubes-
cent on outer side; outer bracts 8-10, linear, up to a half as long as
inner bracts. Ligulate florets yellow, 10-15. Achenes pubescent, 2 mm
long. Flowering June to September; fruiting October (Plate XXXIII,
Fig. 1).

Scrubs, forest edges, river valleys, sandy steppes, wet, sometimes
alkaline meadows, mountains up to 1,500 m.—European part: Trans-
Volga, Volga-Don, Volga-Kama, Bessarabia, Lower Volga, Lower Don,
Black Sea Region, Crimea; Caucasus: Ciscaucasia, Dagestan, Eastern
and Southern Transcaucasia, Talysh; Western Siberia: Upper Tobol;
Soviet Central Asia: Aralo-Caspian Region (Mugodzhary, Chelkar, Irgiz,
and others). General distribution: Armenia and Kurdistan, Balkans
(Romania). Described from Sar Island near Baku. Type in Leningrad.

Note. D.I. Litwinow (op. cit.) considered the name S. arenarius
MB. to have priority for this species. But, apparently, he did not know
that, in the genus Senecio the epithet arenarius is a homonym of S.
arenarius Thunb. (1800) from southern Africa, described earlier by
Thunberg.

3. S. korshinskyi Krasch. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser.
1INe.. 3°¢l9ary a2:

Perennial. Whole plant more or less floccose-woolly or subglabrous.
Stem erect, finely ribbed, sometimes violet below, 30-60 cm high.
Basal and lower cauline leaves oblong, oblong-obovate, or elliptical,
8-20 cm long and 1-5 cm wide, undivided or almost lyrate, more or
less deeply sinuate-toothed, with acuminate teeth, narrowed into rather
long, thickish, basally expanded and serrately winged-petiole; middle
and upper cauline leaves identical to basal leaves, sessile, at base
expanded, with irregularly lobed-toothed auricles; uppermost leaves
linear. Capitula on more or less long peduncle. Involucre about 1.0
mm in dia; outer involucral bracts 2—3, linear, a half as long as inner,
latter lanceolate-linear, 4-5 mm long, blackish-brown in upper half,
glabrous or pubescent, with narrow yellowish-green border. Ligulate
florets 10, yellow, linear, 8 mm long. Achenes 4 mm long, very slightly
curved, indistinctly ribbed, glabrous, brownish; pappus 7-8 mm long.
Flowering July to August.

ais

681

Mountain slopes at 2,000—-3,600 m.—Soviet Central Asia: Pamiro-
Alai Region. General distribution: Afghanistan, Indian Himalayas
(Kashmir). Described from Pyandzh River. Type in Leningrad.

4. S. jacobaea L. Sp. pl. (1753) 870; Ldb. FI. Ross. II, 635;
Boiss. Fl. or. III, 392; Schmalh. Fl. II, 87, ex p.; Grossh. Fl. Kavk. IV,
154; Kryl. Fl. Zap. Sib. XI, 2845.—S. jacobaea var. typica Trautv. in
Mélang. biol. II (1854) 130.—S. erucaefolius Ldb. Fl. Alt. TV (1830)
110, non L.—Jacobaea vulgaris Gaertn. De fruct. et semin. II (1791)
445.—J. sicca Gilib. Fl. lithuan. III (1782) 149.—Ic.: Rchb. Ic. FI.
Germ. XVI, t. 964 (1853); Fedtsch. and Fler. Fl. Evrop. Ross. 993.—
Exs.: Fl. Polon. exs. No. 350; Pl. Finl. exs. No. 984; Eston. plants No.
136.

Perennial. Rhizome branched; whole plant arachnoid-hairy or
subglabrous; stem erect, sulcate, simple or branched from middle, 20-
100 cm high. Basal leaves clustered in rosette, withering before anthesis;
lower cauline leaves with more or less long petiole, oblong-obovate,
obtusely toothed or lyrately incised, with ovate terminal lobe, 2-10 cm
long and 2-8 cm wide; middle cauline leaves sessile, lyrately pinnate,
with linear or lanceolate, apically broadened and incised lobes, 2-8
cm long and 1-3 cm wide; upper leaves pinnately cut, their lobes 2—
10 mm wide, oblong, subobtuse, irregularly toothed or pinnately
divided. Capitula usually numerous, aggregated into corymbose panicle.
Involucre (5)8—10 mm in dia; outer involucral bracts 2—6, a third to a
half as long as broadly lanceolate inner bracts. Ligulate florets yellow,
12-15, as long as involucre, 2.5—4.0 mm wide. Achenes oblong-terete,
2-3 mm long, peripheral achenes glabrous, with pappus of few fragile
hairs; inner achenes pubescent, with well-developed persistent pappus.
Flowering June to August.

Thin forests, often in oak, birch, and linden forests and along their
edges, scrubs, meadow-steppes, sometimes on calcareous soils, often
as weed of roadsides and railroads.—European part: Dvina-Pechora
(introduced), Ladoga-IImen, Baltic Region, Upper Dnieper, Middle
Dnieper, Upper Volga, Volga-Kama, Trans-Volga, Lower Don, Lower
Volga, Black Sea Region, Bessarabia, Crimea, Upper Dniester
Caucasus: Ciscaucasia, Eastern, Western, and Southern Transcaucasia;
Western Siberia: All regions; Eastern Siberia: Angara-Sayans; Soviet
Central Asia: Aralo-Caspian Region, Lake Balkhash Region,
Dzhungaria-Tarbagatai, Syr-Darya, Tien Shan, Pamiro-Alai Region
(Alai Range). General distribution: Central Europe, Atlantic Europe,
Scandinavia, Mongolia, North America (introduced). Described from
meadows of western Europe. Type in London.

682

5. S. ambraceus Turcz. ex DC. Prodr. VI (1837) 348; Turcz. Fl.
baic.-dahur. II, 89.—?Cineraria ambracea Hornem. Hort. Hafn. Suppl.
(1819) 972.—S. jacobaea f. grandiflorus Turcz. ex. DC. Prodr. VI
(1837) 350; Ldb. Fl. Ross. II.—S. erucifolius var. (flor. majoribus)
Ldb. FI. Alt. IV (1833) 110.—S. krylovii Schischk. in Fl. Zap. Sib. XI
(1949) 2847.—S. jacobaea L. var. ambracea Trautv. and Mey. Florula
ochot (1856) 56.

Perennial. Rhizome short, with numerous, rather long roots. Stem
solitary, erect, branched above or sometimes from middle, with
obliquely upright branches, cylindrical, often violet, finely arachnoid-
hairy or subglabrous, 30-80 cm high. Basal leaves withering early,
with petioles as long as or longer than lamina, lamina ovate and oblong,
obtuse, irregularly toothed with obtuse teeth, at base sometimes
pinnately divided, 5—8 cm long, 1-3 cm wide, glabrous on both sides;
cauline leaves sessile, auriculate, pinnately cut, sometimes almost up
to midrib, lobes linear-oblong, subobtuse, in turn pinnately divided or
irregularly toothed; upper-most leaves small, linear, almost entire.
Capitula on long, weakly arachnoid-hairy peduncles, 1.5—6.0 cm long.
Involucre broadly campanulate, 5-6 mm long and about 10 mm in dia,
glabrous, only at base weakly arachnoid-hairy; outer involucral bracts
2-6, linear, acuminate, up to a half as long as inner; inner bracts
lanceolate or narrowly oblong, with scarious margin. Ligulate florets
yellow, 18-20. Achenes 2 mm long, terete, white-pubescent; pappus
white, as long as tubular florets. Flowering July to August.

Wet meadows, birch-aspen forests, sometimes on gravel beds of
rivers.—Western Siberia: Altai; Eastern Siberia: Angara-Sayans,
Dauria, Lena-Kolyma; Far East: Zeya-Bureya. General distribution:
Mongolia, northeastern China. Described from route between Okhotsk
and Yakutsk. Type in Geneva; cotype in Leningrad.

Note. Horneman (loc. cit.) described under the name Cineraria
ambraceus a plant grown from seeds obtained from the Goren Botanical
Garden, but the origin of the plant has remained unknown. It is not
possible to establish if it is identical to Senecio ambraceus Turcz. on
the basis of the short description. Ledebour (loc. cit.) supposed that
Cineraria ambracea Horn. is close and, perhaps, identical to Senecio
lyratifolius Rchb. Some authors have noted that the specific epithet
ambraceus cannot be used for the Siberian plant because the latter
does not have the resinous odor.

6. S. argunensis Turcz. in Bull. Soc. Nat. Mosc. XX (1847) 18.—
S. jacobaea grandiflorus Turcz. in DC. Prodr. Vi (1837) 350 p.—S.
praealtus davuricus DC. Prodr. VI (1837) 351.—S. argunensis f.
angustifolius and f. latifolius Kom. Fl. Manchzh. III (1907) 706.—S.

717

683

blinii Leveille in Fedde, Repert. VIII (1910) 138.—Ic.: Kom. and
Alis. Opred. Rast. Dalnevost. Kr. II, Plate 314; Sato, Willd. Fl. Manch.
and Mong. (1927) tab. 16.

Perennial. Rhizome creeping, 3-7 mm thick, short, scatteredly
covered with slender roots. Stem solitary, erect, 25—60(90) cm high,
together with leaves weakly arachnoid-hairy, sometimes subglabrous,
branched in inflorescence. Basal and lower cauline leaves withering
early; middle cauline leaves sessile, numerous, their laminas ovate, 6—
15 cm long and 2-5 cm wide, pinnately divided, sometimes almost up
to midrib, into linear or narrowly lanceolate, 1-2 cm long and 1-5 mm
wide lobes, terminal lobe often larger than rest. Capitula 5—20 (less
often 2-3), aggregated into corymbose inflorescence, peduncles white-
arachnoid-hairy below capitulum. Involucre broadly campanulate, 5
mm long and 6—7 mm in dia; outer involucral bracts linear, about 10,
considerably shorter than inner; inner bracts lanceolate, acuminate with
narrow scarious margin, weakly pubescent or subglabrous. Ligulate
florets yellow, 10-15, about 10-15 mm long and 2—3 mm wide. Achenes
glabrous, sulcate, oblong, 2.0-2.5 mm long. Flowering August to
September.

Scrubs, forest edges, in reedy and other meadows.—Eastern
Siberia: Dauria; Far East: ?Zeya-Bureya, Ussuri. General distribution:
Mongolia, northeastern China. Described from Urulungui Creek and
Argun River. Type in Leningrad.

7. S. saposhnikovii Krasch. and Schipcz. in Fedde. Repert. XX VI
(1924) 113.—A. doriaeformis auct. nonnul. fl. As med. non DC.

Perennial. Stem 40-50 cm high, erect, glabrous or sometimes
weakly arachnoid-hairy, sulcate, often branched. Leaves thick, lower
and middle ones lanceolate, 7-10 cm long and 1.5—2.5 cm wide, more
or less irregularly cristate-toothed, sometimes pinnatifid, basally
narrowed into petiole, subobtuse or acuminate; upper leaves entire or
with small teeth, linear-lanceolate. Inflorescence corymbose. Involucre
obconical-cylindrical, 5-6 mm long and about 3 mm in dia, on weakly
arachnoid-hairy peduncles and with few, narrowly linear, bracteal
leaves; outer involucral bracts 1-2, linear, a half as long as inner
bracts or almost as long; inner bracts broadly linear, obtuse or subacute,
with broadly scarious and ciliate margin. Ligulate florets 5, slightly
longer than involucre, yellow. Achenes terete, 3 mm long, glabrous;
pappus 6-7 mm long, white, at base stramineous-yellow. Flowering
August.

Turfy and stony slopes, talus, gravel beds and in subalpine meadows
at 2,000-2,500 m.—Soviet Central Asia: Tien Shan, Syr-Darya.
Described from Tien Shan. Type in Leningrad.

718

684

8. S. erraticus Bertol. Rar. Ital. pl. dec. III (1810) 62; Amoen.
Ital. 92 and FI. Ital. IX, 237; DC. Prodr. VI, 349; Ldb. Fl. Ross. II,
634; C. Koch in Linnaea, XVII, 47; Boiss. Fl. or. III, 392; Grossh. FI.
Kavk. IV, 155; M. Popov, Ocherki Rast. i Fl. Karpat. 246.—S.
barbareae folius Krock. Fl. Siles. II, 2 (1790) 421.—S. aquaticus
Huds. barbareifolius Wimm. and Grab. FI. Siles. II, 2 (1829) 151.—
S. erraticus (Bertol.) Hegi, Ill. Fl. IV, 2 (1928) 777.—S. erraticus
Bertol. subsp. barbareifolius (Wimm. and Gr.) Hegi, op. cit. p. 776.—
S. jacobaea var. erraticus Fiori and Paol. Icon. FI. Ital. (1903) 412.—
Ic.: Fior. and Paol. (1903) op. cit.—Exs.: Fl. Ital. exs. No. 1369 and
1369 bis; Fl. exs. Reipubl. Bogem.-Sloven. No. 957; Fl. exs. Austro-
Hung. No. 1808.

Root vertical, with several, more or less thick, lateral roots. Stem
solitary, erect, branched from middle, with long, glabrous or weakly
arachnoid-hairy branches, 20-70 cm high. Basal and lower cauline
leaves with petioles almost as long as lamina, latter glabrous, lyrately
lobed, 5-18 cm long and 2—5 cm wide, terminal lobe ovate, sometimes
with cordate base, with irregular acuminate teeth; middle cauline leaves
pinnately cut, sessile, with semiamplexicaul lobed auricles; uppermost
leaves oblong-linear, small, with few teeth or almost entire. Capitula
numerous at apices of stem and branches, in broadly corymbose panicle;
peduncles as long as involucre or several times as long, glabrous or
weakly hairy. Involucre glabrous, broadly campanulate, about 5 mm
long and 8 mm in dia; outer involucral bracts linear, 2-6, a third to a
half as long as inner; inner bracts oblong-rhombic, with narrow scarious
margin. Ligulate florets 8-12, yellow, 4-5 mm long and about 2 mm
wide. Pappus caducous; peripheral achenes subglabrous, inner ones
(of tubular florets) more or less pubescent. Flowering July to August.

Forest edges, in scrubs on marshy meadows, gravel beds, and
roadsides.—European part: Baltic Region (Kaliningrad), Upper
Dniester (mainly in Trans-Carpathian Mountains); Caucasus: Western
Transcaucasia. General distribution: Central Europe, western Mediter-
ranean Region, Atlantic Europe, Balkans-Asia Minor. Described from
Italy. Type in Bologna.

Note. In the Herbarium of the V.L. Komarov Botanical Institute
of the Academy of Sciences of the USSR, there is one specimen of this
species from Zheleznovosdsk (Northern Caucasus) from the herbarium
of I.Ya. Akinfiev the famous researcher of the Caucasus. Apparently
the label on this specimen is erroneous because other material of this
species from the Northern Caucasus could not be found. In Transcaucasia,
S. erraticus Bertol. is known from Western Transcaucasia, where it is a
common plant. There are certain grounds to suppose that in Western
Transcaucasia this plant was introduced from the Western Mediterranean
region, where suitable conditions for its survival are found.

719

685

Series 2. Borysthenici Schischk.—Leaves twice or thrice pinnately
incised, with narrowly linear terminal lobe; plants weakly pubescent.

9. S. borysthenicus Andrz. ex DC. Prodr. VI (1837) 351, nom. in
syn.—S. praealtus borysthenicus DC. ibid. (1837); Ldb. Fl. Ross. II,
634.—S. jacobaea var. borysthenica Trautv. in Mélang. biol. II (1854)
131 and in Bull. de la classe phys.-meth. AC. Sc. de Saint Pétersb. XII
(1854) 351; Schmalh. Fl. II, 87.—Exs.: GRF No. 3494.

Perennial. Rhizome vertical or ascending, short. Stems few, erect,
weakly flexuous, finely sulcate, branched from middle or only at apex;
branches upright, arachnoid-hairy, sometimes subglabrous, usually violet
below, 30-90 cm high. Basal and lower cauline leaves withering early,
long-petiolate, petioles as long as lamina, lamina oblong, 5-10 cm
long and 2—4 cm wide, twice or thrice pinnately incised, with linear,
5-10 mm long and 0.5—1.0(3.0) mm wide lobes; middle cauline leaves
like basal leaves, but smaller, short petiolate, upper leaves sessile; all
leaves arachnoid-hairy or sometimes subglabrous. Capitula numerous,
at apices of stem and branches, forming paniculate or more or less lax
corymbose inflorescence on 1-4 cm long peduncles, with small bracteal
leaves. Involucre campanulate, 5-7 mm long and almost as much in
dia in upper part; outer bracts 1-5, linear, acuminate, a half as long as
inner ones; inner bracts oblong, greenish, whitish along margin, with
darker stripe on dorsal side. Ligulate florets 10-15, oblong-linear,
almost twice as long as involucre. Achenes terete, ribbed, peripheral
achenes glabrous, lacking pappus, inner achenes finely pubescent, with
about 2.5 mm-long rudimentary pappus. Flowering June to August;
fruiting September.

River sands, sandy steppes and sandy pine woodlands.—European
part: Upper Dnieper, Middle Dnieper, Volga-Don, Black Sea Region,
Lower Don, Crimea (north), Bessarabia, Upper Dniester. Endemic.
Described from Podolia and Kherson. Type in Geneva.

Note. Christian von Steven also noted this plant. He collected it
from the shifting sands between Perekop and Alyoshka and near Kerch,
where it is found in abundance. Steven referred it to S. erucifolius L.,
but with the comment that it strongly differs from var. genuinus and
deserves to be treated as a separate species (which was already done
earlier by Andrzejowski).

10. S. schischkinianus Sof. in Izv. Akad. Nauk. Azerb. SSR, No.
1 (1957) 83.—S. erucifolius auct. Fl. cauc. non L.: Schmalh, FI. I, 87,
p. p.; Lipsky, Fl. Kavk., 351; Grossh. Fl. Kavk. IV, 554.—S. erucifolius
latilobus Boiss. FI. or. III (1875) 392.—S. borysthenicus Grossh. Opred.
Rast. Kavk. 470, non Andrz.—Ic.: Sof. op. cit. p. 85.

720

686

Perennial. Root ascending, slender. Stem erect, simple or branched
above, weakly sulcate, including leaves somewhat hairy or subglabrous,
40-70 cm high. Leaves twice pinnately incised, with linear or oblong,
sessile lobes; basal leaves withering early; lower cauline leaves short—
petiolate, obovate or oblong, 8—10 cm long and about 3 cm wide, their
lobes narrowly oblong, with occasional teeth; middle cauline leaves
smaller, sessile, amplexicaul, auriculate. Peduncles with small, 1-2
mm long, linear-subulate bracteal leaves. Capitula numerous, 20-30,
in corymbose-paniculate inflorescence. Involucre 6-8 mm long and
almost as much in dia; outer involucral bracts 3—5, linear-subulate, a
third as long as inner ones; inner bracts lanceolate or linear-lanceolate,
10-14, with scarious border, subglabrous. Ligulate florets yellow, 12-
15, 8-10 mm long and 1.5-2.0 mm wide. Achenes terete, 3-4 mm
long, densely hairy with stiff hairs; pappus white, 2-3 times as long as
achenes. Flowering June to August; fruiting August to September.

Sands, particularly river sands, rarely on talus.—Caucasus:
Ciscaucasia, Dagestan. Endemic. Described from Dagestan
(Kumtorkali). Type in Leningrad.

10a. S. ferganensis Schischk. sp. nov. in Addenda, XXV, 881.

Perennial. Stem erect, branched above, with obliquely upright
branches, including leaves arachnoid-floccose-pubescent, 60-75 cm
high. Leaves twice pinnately incised; basal and lower cauline leaves
withering early; cauline leaves numerous, narrowly ovate, sessile, at
base somewhat expanded and with inconspicuous auricles, 6-15 cm
long and 1.5-3.0 cm wide. Inflorescence corymbose, with upward
spreading branches; capitula numerous, up to 15-30, broadly
campanulate, 5 mm long and 10 mm in dia. Outer involucral bracts 3—
5, linear, adpressed to inner bracts or somewhat squarrose, a third to
two-thirds as long as inner; inner bracts narrowly ovate, subacute,
weakly hairy or subglabrous. Ligulate florets yellow, twice as long as
involucre. Achenes glabrous, about 2 mm long. Flowering June to
July.

Gravel beds of rivers, banks of irrigation canals, slopes.—Soviet
Central Asia: Syr-Darya, Pamiro-Alai Region (Alai Range). Described
from Alai Range. Type in Leningrad.

Section 2. Incani DC. Prodr. VI (1937) 355.—Velutini Boiss. FI.
or III (1875) 384.—Perennials; leaves hairy, particularly below,
variously pinnately lobed, with obtuse lobes, upper leaves often with
auricles or amplexicaul; involucre in 2 whorls with very few outer
involucral bracts; ligulate florets 5-12, yellow.

72

—

687

Series 1. Cinerariae Schischk.—Leaves twice pinnately incised,
dark green above, with sparse floccose-arachnoid-hairs, densely white-
tomentose beneath.

11. S. cineraria DC. Prodr. VI (1837) 355; Boiss. Fl. or. III, 395;
Fedtsch. and Fler. Fl. Evrop. Ross. 993.—S. maritimus Rchb. Fl. Germ.
exc. (1830-1832) 244, non L. fil. (1791).—Cineraria meritima L. Sp.
pl. (1753) 925.—Jacobaea tomentosa Moench. Meth. (1794) 587.—
Ic.: Sibth. and Sm. Fl. Graec. IX, tab. 871; Hegi, Ill. Fl. VI, 2, 725,
729; Jav. and Csapody. Iconogr. Fl. Hung. (1933) 531; Rchb. Ic. FI.
Germ. XVI, tab. 968 (1853); Gartenfl. XXXI (1882) 276.

Perennial. At base somewhat woody. Stem ascending or erect,
weakly flexuous, simple, usually branched only in inflorescence, less
often from base, white-tomentose, 20-40 cm high, and 3-5 mm thick.
Basal and lower cauline leaves long-persisting, ovate, twice pinnately
incised, with terminal lobe oblong, obtuse, S—10 mm long and 2-5 mm
wide, dark green above, with sparse floccose-arachnoid hairs, densely
white-tomentose beneath, petiolate, petiole shorter than lamina; cauline
leaves like basal, gradually reducing upward and less divided (simple
pinnate). Capitula numerous, in corymbose inflorescence, on white-
arachnoid-hairy peduncles. Involucre 5 mm long and 6 mm in dia;
inner involucral bracts linear-lanceolate, acuminate, on dorsal side
white-arachnoid-hairy; outer bracts 5, narrowly linear, much shorter
than inner ones. Ligulate florets yellow, about 8, twice as long as
involucre. Achenes glabrous, 1.5 mm long; pappus brownish. Flowering
June to July; fruiting August to September.

Coastal rocks. In USSR, as introduced plant that escaped
cultivation, found growing wild in places.—European part: Crimea
(southern coast). General distribution: Western Mediterranean Region,
in the east as far as Greece. Described from Mediterranean coast.
Type in Geneva.

12. S. lorentii Hochst. in Flora, XXVIII (1845) 27, non Griseb.
(1874).—S. gundelius C. Koch in Linnaea, XXIV (1851) 364.—S.
armenius Jaub. and Sp. Ill. pl. or. [V (1850-1853) 143; Boiss. Fl. or.
III, 397; Grossh. Fl. Kavk. IV, 162.—S. pedunculosus Trautv. in Tr.
Bot. Sada, III (1875) 277.—Ic.: Jaub. and Sp. op. cit. tab. 398.—Exs.:
Fl. cauc. exs. No. 221.

Perennial. Rhizome ascending, about 0.8 cm thick. Stem erect,
somewhat flexuous, striate, leafy up to middle or slightly above, apically
more or less branched, densely floccose-white-tomentose, 15-65 cm
high. Basal and lower cauline leaves with expanded petioles almost as
long as lamina, lamina oblong, 6-12 cm long and 2.5—7.0 cm wide,

722

688

pinnately incised, with 3-5 ovate, obtuse, entire or weakly divided
lobes, terminal lobe broadly deltoid, larger; middle cauline leaves (if
present) similar to basal, smaller, sessile; upper leaves small, linear-
lanceolate, toothed or entire, auriculate, all leaves, particularly beneath,
densely arachnoid-hairy. Inflorescence lax corymbose. Capitula 2-10,
on short or long, white-tomentose peduncles. Involucre white-tomentose,
broadly campanulate, 8-10 mm long and 12-15 mm in dia; outer
involcural bracts 1-3, linear or setaceous, a half as long as inner ones;
inner bracts linear, dorsally keeled. Ligulate florets yellow, 10-15, 15
mm long and 4.5 mm wide, glabrous. Flowering July to August.

Crevices of rocks, stony slopes.—Caucasus: Western Transcaucasia
(gorge of Chorokh River). General distribution: Armenia and Kurdistan.
Described from vicinity of Erzerum. Type was in Berlin.

13. S. lipskyi Lom. in Tr. Tifl. Bot. Sada, III (1899) 51; Grossh.
Fl. Kavk. IV, 151.—S. saxatilis Lom. in Tr. Tifl. Bot. Sada, II (1897)
28 non Wall. (1828) No. 3131, nom. nud., descr. apud DC. Prodr. VI
(1837) 367.—Exs.: Pl. or. exs. No. 246.

Perennial. Rhizome ascending or horizontal, 0.6—0.8 cm thick,
covered with long and slender roots. Stem usually solitary, erect, white-
tomentose at base, floccose-arachnoid-hairy and branched above, 40-
75 cm high. Basal leaves numerous, dark green and arachnoid-hairy
above, white-tomentose beneath, with petiole as long as, shorter or
longer than lamina, lamina ovate or oblong-ovate, 5-9 cm long, 3-6
cm wide, lyrate, with 2-4 pairs of lobes, terminal lobe deltoid, larger
than others, basally truncate or weakly cordate, lateral lobes gradually
reduced downward, usually remote, all lobes with coarse, unequal,
obtuse teeth or lobate, with broad base, decurrent on petiole; cauline
leaves fewer, lower and middle cauline leaves similar to basal but
sessile, with semiamplexicaul auricles; upper leaves lanceolate or linear-
lanceolate, divided or toothed, sometimes almost entire, small. Capitula
2-7, sometimes up to 25, forming corymbose inflorescence. Involucre
5-6 mm long, 12-15 mm in dia; outer involucral bracts 2-7, linear, a
third to a half as long as inner ones; inner bracts oblong, acute, with
scarious margin. Ligulate florets egg-yellow, about 10, 10-12 mm long
and 2-5 mm wide. Achenes terete, 4-5 mm long, ribbed, white-
pubescent along ribs, later subglabrous(?). Flowering June to July;
fruiting August.

Mountain-steppe and subalpine, often calcareous, slopes on talus,
at 1,600-2,400 m.—Caucasus: Southern Transcaucasia. General
distribution: Northern Iran. Described from Takali Mountains near the
village of Bechenakh. Type in Leningrad.

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689

14. S. massagetovii Schischk. sp. nov. in Addenda, XXV (1960)
882.—S. pedunculosus Trautv. in sched. ad Herb. Inst. bot. nom.
Komarovii, Ac. Sc. URSS non Trautv. in Tr. Bot. Sada, III, 2 (1875)
277.—Exs.: Herb. Fl. cauc. curante. Woronow, No. 98.

Perennial. Rhizome rather thick, 0.5 cm thick, with few long roots.
Stem solitary, floccose-arachnoid-hairy, sulcate, more or less branched
above, 50-75 cm high. Basal and lower cauline leaves with expanded
petioles as long as lamina or longer; lamina ovate-oblong, 6-20 cm
long and 3-10 cm wide, dark green and subglabrous above, densely
arachnoid-hairy beneath, lyrate, pinnate, with 2-3 somewhat remote
lobes reduced downward, unequally toothed, terminal lobe considerably
larger than others; middle cauline leaves similar to basal but sessile,
with semiamplexicaul auricles. Capitula numerous (10-40), forming
lax, corymbose paniculate inflorescence, peduncles as long or many
times exceeding capitula. Involucre floccose-arachnoid-hairy, 6 mm
long and 10 mm in dia; outer involucral bracts narrowly linear, 1-5,
a half as long as inner (sometimes absent); inner bracts oblong-linear,
dorsally keeled. Ligulate florets yellow, 13-15, about 10 mm long and
2-3 mm wide. Achenes terete, glabrous, weakly ribbed, 4.0-4.5 mm
long. Flowering June.

Stony slopes.—Caucasus: Eastern Transcaucasia. Endemic.
Described from vicinity of Borzhomi (Sakochavi). Type in Leningrad.

Note. Trautvetter identified the plant from Borzhomi as S.
pedunculosus Trautv. But this plant, described by him from the vicinity
of Erzerum, is identical to S. armenius Jaub. and Sp. The plant from
Eastern Transcaucasia differs from the Erzerum plant by smaller and
more numerous (10-40 and not 2-10) capitula, the uniform leafiness
of the stem, and later flowering. The range of this species is not yet
quite clear.

Series 2. Arnicoidei Schischk.—Stem hollow; leaves undivided,
ovate, unevenly toothed, green above, densely arachnoid-hairy beneath,
with suborbiculate auricles; outer involucral bracts 8, narrowly linear;
achenes about 7 mm long, glabrous.

15. S. pseudoarnica Less. in Linnaea, VI (1831) 240; Ldb. FI.
Ross. II, 642; Kom. Fl. Manchzh. III, 704.—Arnica maritima L. Sp.
pl. (1753) 884, non Senecio maritimus L. fil. (1781).—S. pseudoarnica
var. kurilensis Kudo, Fl. Paramush. (1922) 171.

Perennial. Rhizome short, covered with numerous long roots. Stem
erect, simple, hollow, glabrous below, about 1 cm thick, arachnoid-
hairy above, often violet, 15—70 cm high. Basal and lower cauline
leaves withering early, middle and upper leaves oblong, ovate, or

724

690

broadly ovate, undivided, 8-25 cm long and 2.5—9.0 cm wide, obtuse,
unevenly toothed, green and glabrous above, much paler beneath and
usually densely arachnoid-hairy, often whitish, cuneate toward base,
narrow, sessile and with suborbicular, indistinctly toothed auricles.
Capitula at apex of stem, 2-15, forming lax corymbose inflorescence,
on arachnoid-hairy or subglabrous peduncles, peduncles short at
beginning of anthesis, later elongated, with 1-4 cm long, narrowly
linear bracteal leaves. Involucre broadly campanulate, almost
hemispherical, 10-12 mm long and 20-25 mm in dia; outer involucral
bracts about 8, narrowly linear, as long as inner bracts, weakly
pubescent; inner bracts oblong, with broad scarious border, acuminate,
more or less pubescent in outer side. Ligulate florets yellow, about 15,
15 mm long and 3-4 mm wide. Achenes glabrous, lustrous, light brown,
sulcate, terete, 7 mm long and about 1 mm wide; pappus white.
Flowering July to first half of September; fruiting September.
Coastal sands and gravel beds.—Far East: Ussuri, Uda River Area,
Okhotsk, Kamchatka, Kuril Islands. General distribution: Beringia,
North America, Japan. Described from Kamchatka. Type in Berlin.

16. S. kubensis Grossh. in Tr. Azerb. Otd. Zakavk. Fil. Akad.
Nauk SSSR, Bot. 1 (1933) 59; Grossh. Fl. Kavk. IV, 159.

Perennial. Whole plant densely arachnoid-hairy. Stem 40-60 cm
high, branched. Basal leaves withering before anthesis; cauline leaves
with semiamplexicaul auricles, more or less long-petiolate, oblong,
lyrate, pinnatisect, with 2 pairs of lateral, ovate, obtuse lobes and
longer, shallow and obtusely sinuate-toothed terminal lobe; all lobes
subglabrous above, white arachnoid-hairy beneath; upper leaves much
smaller. Inflorescence corymbose, with many capitula; peduncles usually
short. Involucre 10 mm long and 10-12 mm wide; outer involucral
bracts linear, subobtuse, blackish, weakly pubescent. Ligulate florets
bright yellow, 12. Achenes glabrous, 4-5 mm long. Flowering July.

Subalpine zone, crevices of rocks.—Caucasus: Eastern
Transcaucasia. Endemic. Described from former Kuba District. Type
in Baku; cotype in Leningrad.

17. S. pandurifolius C. Koch in Linnaea, XVII (1843) 48; Ldb.
Fl. Ross. II, 644; Boiss. Fl. or. III, 399; Grossh. Fl. Kavk. IV, 153;
Kolak. Fl. Abkh. IV, 249.—S. conipes Somm. and Lev. Nuov. Giorn.
Bot. Ital. (1895) 88; Tr. Peterb. Bot. Sada, XVI, 239; Grossh. FI.
Kavk. IV, 153.—Exs.: Pl. cauc. exs. No. 50; Pl. or exs. No. 399.

Perennial. While plant white-arachnoid-hairy. Rhizome 5-10 mm
thick, with numerous roots. Stems solitary or few, erect, somewhat
branched above, floccose-pubescent, 20-50 cm high. Basal leaves with

2g

691

petioles shorter than lamina, lamina broadly ovate, undivided, basally
almost cordate, apically obtuse or panduriform, with large, terminal
lobe cordate at base, and 2-3 pairs of small lobes, whitish beneath
from dense arachnoid hairs, dark green above, with floccose pubescence,
2.5-12.0 cm long and 2-11 cm wide; cauline leaves 1-4, panduri-
form, sometimes almost undivided, usually sessile, with semiamplexicaul
auricles. Capitula 2-6, on long peduncles several times as long as
capitula. Involucre about 10 mm long and 12-15 mm in dia; outer
involucral bracts linear, 1-7, a fourth to a half as long as inner ones;
inner bracts linear-oblong, acuminate, densely arachnoid-hairy, later
subglabrous. Ligulate florets yellow, 12, about 10 mm long and 4 mm
wide. Achenes glabrous, terete, 4.0-4.5 mm long; pappus slightly shorter
than tubular florets. Flowering May; fruiting June.

Calcareous rocks.—Caucasus: Ciscaucasia (Teberda), Western
Transcaucasia. General distribution: Northern Turkey (Artvin). De-
scribed from Megrelin. Type in Berlin.

Section 3. Quadridentati Boiss. Fl. or. III (1875) 385.—Whole
plant pubescent; leaves lyrately pinnatifid, with 2-5 pairs of sharp-
toothed lobes; tubular florets with 4 teeth.

18. S. taraxacifolius (MB.) DC. Prodr. VI (1837) 348; Ldb. FI.
Ross. II, 633; Boiss. Fl. or. III, 413; Grossh. Fl. Kavk. IV, 153.—
Cineraria taraxacifolia MB. FI. taur.-cauc. II (1808) 313.—Ligularia
aucheri DC. Prodr. VII (1838) 300.

Perennial. Whole plant covered with short glands, pale green, very
rarely indistinctly pubescent. Rhizome ascending, 0.5—0.8 cm thick,
with numerous, long, slender roots. Stems solitary or 2—3, at base
covered with dark brown remnants of old petioles, branched in
inflorescence, erect, sulcate, 10-50 cm high. Basal (withering early)
and lower cauline leaves on more or less long, winged petioles,
expanded at base, semiamplexicaul, their laminas ovate, lyrately divided
with 2-5 pairs of deltoid or broadly ovate, sharp-toothed lobes, terminal
lobe larger, ovate or cordate; upper leaves sessile or subsessile, smaller,
usually auriculate, pinnately divided, uppermost leaves linear-lanceolate
or linear, long-acuminate, sharp-toothed. Capitula 3-12, more or less
long-pedunculate, forming lax corymbose panicle. Involucre about 10
mm long and 15 mm in dia; outer involucral bracts setaceous, shorter
than involucre, 1-3 or absent; inner bracts lanceolate, acuminate, flat
and densely glandular-pubescent on dorsal side. Ligulate florets 10—
12, yellow, 15 mm long and 1.5—3.0 mm wide; tubular florets with 4
teeth. Achenes glabrous, terete, 4-5 mm long; pappus brownish, as
long as tubular florets. Flowering July to August. (Plate XXXIV, Fig. 2).

726

692

Rocks in alpine zone, edges of glaciers, moraines, near melting
snow at 2,400-3,100 m.—Caucasus: Ciscaucasia, Dagestan, Eastern,
Western, and Southern Transcaucasia. General distribution: Armenia
and Kurdistan, Asia Minor (Pontic Range), Iran (north). Described
from Georgia. Type in Leningrad.

Note. Boissier segregated this species in a special “section,”
Quadridentati, mentioning that S. taraxacifolius (MB.) DC. differs
from all other groundsels. But Boissier reported this very character for
S. stenocephalus Boiss. (=S. pojarkovae Schischk.), in which the tubu-
lar florets also are four-toothed.

Section 4. Crociserides DC. Prodr. VI (1837) 384.—Plants peren-
nial; leaves undivided, entire or toothed, less often lobate or runcinate,
basal leaves narrowed into long petiole; involucre 2-rowed, less often
many bracts in outer whorl.

Series 1. Subalpini Schischk.—Involucre 8 mm long and 10-15
mm in dia, usually with 8-10 bracts; ligulate florets 10—12(15), yellow;
achenes about 4.5 mm long. High mountain plants growing mainly in
subalpine zone.

19. S. subalpinus C. Koch in Flora, XVII (1834) 614, in textu.—
S. cordifolius auct. Carp. non Clairv.

Perennial. Rhizome horizontal, apically ascending, about 5 mm
thick. Stem solitary, sulcate, erect, branched in inflorescence, arachnoid-
hairy when young, later subglabrous, 40-80 cm high. Basal and lower
cauline leaves with petioles longer than lamina, lamina broadly ovate,
less often almost round, acute, or obtuse, sometimes roundish at apex,
usually cordate at base, unevenly toothed with coarse teeth, glabrous,
2.5—6.0 cm long and 2.5—-5.5 cm wide; middle cauline leaves on short,
often expanded petiole, auriculate; upper leaves linear-oblong or linear,
entire. Capitula 2-11, forming lax, corymbose inflorescence. Involucre
broadly campanulate, 8 mm long and 12-15 mm in dia; involucral
bracts linear-oblong, with narrow scarious margin, green or dark violet,
dorsally arachnoid-hairy or subglabrous. Ligulate florets about 12,
yellow or slightly orange, 10 mm long and 2.0—-2.5 mm wide. Achenes
ovate, glabrous; about 2 mm long; pappus brownish. Flowering July to
August.

Mountain zone, at 1,600-1,700 m, mainly in alder stands.—
European part: Upper Dniester (Goverla Mountain, Pope Ivan, Petros,
Chorna Gora [Black Mountain] and other mountains). Endemic. De-
scribed from Carpathians. Type was in Berlin.

727

693

20. S. buschianus Sosn. in Zhurn. Russk. Bot. Obshch. XIV (1929)
84; Grossh. Fl. Kavk. IV, 150.

Perennial. Rhizome short, with numerous roots. Base of stem
covered with thin, brownish, fibrous remnants of dead leaves. Stem
erect, sulcate, solid, branched in inflorescence, usually densely
arachnoid-hairy, 40—60 cm high. Basal leaves with petioles as long, as,
shorter or longer than lamina, their laminas ovate or lanceolate, obtuse
or somewhat acute, unevenly toothed with short, acute, deltoid teeth,
abruptly or gradually narrowed at base into petiole, 6-18 cm long and
3-6 cm wide, on both sides covered with semiappressed entangled
hairs, particularly on veins; middle cauline leaves like basal but sessile,
semiamplexicaul; upper leaves smaller, lanceolate or almost linear,
weakly toothed, sometimes almost entire. Inflorescence indistinctly
corymbose; capitula 3-12, on long, arachnoid-hairy peduncles with
few small bracteal leaves. Involucre about 12 mm long and 15 mm in
dia; outer involucral bracts 8-10, linear or lanceolate, half as long as
or very slightly shorter than inner ones; inner bracts lanceolate,
acuminate, along margin whitish, more or less densely arachnoid-hairy.
Ligulate florets about 15, yellow, oblong, 2 cm long and 3-5 mm
wide. Young achenes ribbed, glabrous; pappus white, shorter than
tubular florets. Flowering July to August.

Subalpine meadows at 2,100-2,400 m.—Caucasus: Ciscaucasia
(Balkaria). Endemic. Described from Balkaria. Type in Leningrad.

21. S. pseudoorientalis Schischk. nom. nov.—S. orientalis Willd.
Sp. pl. III (1800) 2006, non Mill. Gard. Dict. ed. VIII (1768) No. 10;
Ldb. Fl. Ross. II, 642; Boiss. Fl. or. III, 406, ex p.; Grossh. Fl. Kavk.
IV, 150.

Perennial. Rhizome horizontal, with numerous, thick and long
fibrous roots; tip of rhizome floccose-woolly. Stem solitary, glabrous,
sulcate, erect, uniformly hairy, simple, only in inflorescence more or
less branched, 20-60 cm high. Basal leaves withering early, glabrous
or weakly pubescent beneath, oblong or oblong-lanceolate, gradually
narrowed into long petiole, including petiole 10-25 cm long and 1.5-—
3.0 cm wide, acute or acuminate, finely toothed or almost entire; middle
leaves short-petiolate, short-pubescent beneath; upper leaves sessile,
lanceolate or linear, indistinctly toothed or entire, usually adpressed to
stem. Capitula at stem apices, 1-5 or numerous and then forming lax
corymbose or paniculate-racemose inflorescence. Involucre semiglobose,
about 10 mm long and 15 mm in dia, outer involucral bracts about 10,
linear, acuminate, usually weakly floccose, a third to a half as long as
inner bracts, less often almost as long; inner bracts lanceolate-linear,
subglabrous, obtusely keeled on dorsal side, with scarious margin, acute

728

694

and with a black spot at apex. Ligulate florets 10-12, yellow, 15-25 mm
long and 3-4 mm wide. Achenes terete, ribbed, glabrous or sometimes
weakly pubescent, 3.0-4.5 mm long; pappus white, considerably longer
than involucre. Flowering June to July; fruiting August.

Mountain-steppe slopes, in thin mountain forests, subalpine
meadows, at 1,600-2,500 m.—Caucasus: Southern Transcaucasia,
Talysh. General distribution: Armenia and Kurdistan, Balkans-Asia
Minor. Described from Armenia. Type in Berlin.

22. S. kolenatianus C.A.M. in Beitr. Pfl. Russ. Reich. VI (1849)
34; Schischkin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk. SSSR, XV,
409.—S. amphibolus C. Koch in Linnaea, XXIV (1851) 361; Boiss.
Fl. or. III, 405; Grossh. Fl. Kavk. IV, 150.—S. longiradiatus Trautv.
in Ind. sem. Horti Petrop. (1865) 37.

Perennial. Whole plant glabrous or weakly arachnoid-hairy; rhizome
short, densely covered with thin roots. Stem solitary, usually erect,
simple or somewhat branched, sulcate, uniformly leafy, 20-80 cm high.
Basal and lower cauline leaves withering early, with petioles as long
as lamina, their laminas elliptical or elliptical-lanceolate, short-ciliate
along margin, with small, subacute, somewhat uneven teeth, obtuse or
subacute, with prominent veins, 3-10 cm long and 2.5-5.0 cm wide;
middle cauline leaves ovate-lanceolate or lanceolate, gradually narrowed
into short petiole or almost linear, semiamplexicaul, acuminate with
aristate tip. Inflorescence lax corymbose. Capitula 2—7(10) or solitary.
Involucre 8 mm long and 10 mm in dia, outer involucral bracts 5-7,
linear, almost as long as inner bracts, glabrous or arachnoid-hairy;
inner bracts linear-oblong, acuminate; all involucral bracts with a
black spot near apex. Ligulate florets yellow, about 10, twice as long
as involucre, 2—-3(4) mm wide. Achenes ribbed, weakly hairy or
subglabrous, 4.0-4.5 mm long. Flowering July to August.

Subalpine zone in meadows, at 2,000-2,700 m.—Caucasus:
Ciscaucasia, Dagestan, Eastern and Western Transcaucasia. Endemic.
Described from Kazbek Mountain. Type in Leningrad.

Series 2. Seravschanici Schischk.—Leaves on more or less long
petioles, often 1.5 times as long as lamina, lamina undivided or basally
pinnately divided; involucre about 15 mm long and 10 mm in dia;
outer involucral bracts usually numerous; ligulate florets 6-10; achenes
terete, 6-8 mm long and about 1 mm wide, with short and appressed
hairs or glabrous.

23. S. olgae Rgl. and Schmalh. in E. Regel. Putesh. v Turkestan,
A.P. Fedtschenko in 18 (1881) 45.—S. acrabatensis Franch. in Ann.

695

729 Sc. Nat. 6 sér. Bot. XVI (1883) 312.—S. serwschanicus Winkl. in Tr.
Bot. Sada, XIV (1895) 153.—Exs.: GRF No. 4198.

Perennial. Rhizome ascending, 0.7-1.0 cm thick. Stem solitary,
finely floccose-hairy or subglabrous, simple, branched only in
inflorescence, 30-80 cm high. Basal and lower cauline leaves with
usually long petioles, 1.5 times as long as lamina, lamina undivided or
pinnately divided at base, with remote, few, often solitary lobes, broadly
or oblong-ovate, 10-15 cm long and 5—7 cm wide, on both sides covered
with soft, curly, semiappressed hairs, particularly on veins, sometimes
almost subglabrous above, acute or subobtuse, with uneven, broadly
deltoid, subobtuse or acute teeth, middle cauline leaves like lower, but
on short petiole, often sessile and amplexicaul; upper leaves sessile.
Inflorescence at stem apex, corymbose, with 3-7 capitula on more or
less long peduncles. Involucre about 15 mm long and 10 mm in dia;
outer involucral bracts 4—5, sparsely pubescent, narrowly linear, more
or less appressed to inner bracts; inner bracts narrowly oblong,
pubescent with scarious-coriaceous margin. Ligulate florets 8-10,
yellow, oblong, 1.5—2 times as long as involucre, 3-4 mm wide. Achenes
terete, covered with short and compactly semiappressed hairs, 6-8 mm
long, 1 mm wide. Flowering May to June; fruiting June to July.

Stony and grassy slopes near rocks in juniper zone. At 1,000-
1,200 m.—Soviet Central Asia: Pamiro-Alai Region. Endemic.
Described from Bastand Gorge. Type in Leningrad.

24. S. franchetii Wink]. in Tr. Peterb. Bot. Sada, XI (1889) 165.—
S. bucharicus Winkl. op. cit. (1889) 335.

Perennial. Rhizome ascending. Stem solitary, simple, arachnoid-
hairy, 50-80 cm high. Leaves lanceolate or ovate, arachnoid-hairy
beneath, subglabrous above; basal and lower cauline leaves on long,
broadened petioles as long as lamina, at base often auriculate; upper
leaves strongly reduced; all leaves subacute, usually coarse teeth.
Inflorescence corymbose; capitula 3-13. Involucre 15 mm long and
almost as much in dia; outer involucral bracts 1—5, linear; inner bracts
18-20, lanceolate, keeled, dorsally pubescent, with scarious border.
Ligulate florets 8, yellow. Achenes pubescent, terete, 6-8 mm long.
Flowering April to May; fruiting June to July.

Pistachio forests and stony slopes, at 1,000—-2,000 m.—Soviet
Central Asia: Pamiro-Alai Region. Endemic. Described from Hissar.
Type in Leningrad. &

Note. A significant error crept into Winkler’s original description
of this species: the achenes are characterized as glabrous. On the basis
of the available type specimen, it is not possible to decide if the achenes

730 are glabrous or pubescent, because mature achenes are absent. However,

696

new collections from the same localities, and moreover in the fruiting
stage, show undoubtedly that an error had crept into the author’s
description. In this series of species, only S. paulsenni O. Hoffm. has
glabrous achenes.

25. S. paulsenni O. Hoffm. ex Paulsen in Kjoeb. Vidensk. Meddel.
(1903) 152.

Perennial. Plant glabrous or weakly floccose-hairy. Rhizome
creeping. Stems few or numerous, at base densely covered with whitish
or brownish remnants of petioles of dead leaves, simple, erect, whitish,
more or less lustrous and sulcate, 20-70 cm high. Basal leaves with
petioles as long or almost twice as long as lamina, their laminas elliptical
or oblong, 5-10 cm long and 1.5—5.0 cm wide, basally long-narrowed,
with large and unequal teeth or toward base sometimes sinuate-lobate
or almost pinnatifid; cauline leaves like basal, but on much shorter
petioles; upper leaves smaller and sessile. Capitula at stem apices,
solitary or 2-4, on 3-6 cm-long peduncle, peduncles thickened below
capitulum. Involucre 15 mm long and 15—20 mm in dia, at base weakly
pubescent; outer involucral bracts nemerous, linear, 6-8 mm long and
about | mm wide, leafy, short-acuminate, apically scarious; inner bracts
20-25, linear-oblong, green, purple above and covered with thin hairs.
Ligulate florets 12, lemon-yellow 2 cm long and about 6 mm wide;
tubular florets slightly longer than involucre. Achenes terete, glabrous,
lustrous, 6-7 mm long and about 0.7 mm wide, longitudinally ribbed.
Flowering June; fruiting August to September.

Stony and rubbly slopes, on talus and rocks, at 3,000—4,100 m.—
Soviet Central Asia: Pamiro-Alai Region, Tien Shan (west). Endemic.
Described from Pamiro-Alai Region. Type in Stockholm.

Note. Initially, I and other taxonomists determined this species to
be S. acrabatensis Franch. Thanks to the courtesy of the Herbarium of
the Paris National Museum, I received an authentic specimen of S.
acrabatensis for study. Although the achenes were still immature in
the specimen sent to me, they appeared pubescent. According to
Czerneva’s oral communication, attempts to collect a topotype of S.
acrabatensis Franch. from the Ak-Rabat Pass proved futile, as only S.
olgae was found there.

Series 3. Macrophylli Schischk.—Inflorescence large, corymbose
paniculate or racemose; lower leaves 15-40 cm long and 6—10(15) cm
wide; ligulate florets 5, less often 8.

26. S. macrophyllus MB. FI. taur.-cauc. II (1808) 308, III (1819)
571; Ldb. Fl. Ross. II, 639, ex p.; Boiss. Fl. or II, 406; Gross. FI.
Kavk. IV, 151.—S. umbrosus Bess. Enum. pl. Volh. (1822) 33, 108.—

697

731 §. microphyllus (errore typogr.) Spreng. Syst. veg. III (1826) 556.—
S. doria auct. fl. cauc. C. Koch in Linnaea, XVII (1843) 48, non L.
Perennial. Rhizome short, horizontal, thick. Stem 0.7 cm thick,
solid, sulcate, more or less arachnoid-hairy, sometimes subglabrous,
100-150 cm high. Basal and lower cauline leaves undivided, obovate-
oblong, gradually narrowed into long winged petiole, along margin
unevenly cristate-toothed, teeth broadly triangular, apically
cartilaginous, including petiole 20-40 cm long and 6—10(15) cm wide,
subglabrous above, pubescent beneath; upper leaves smaller, sessile,
with semiamplexicaul base, sometimes almost entire; uppermost leaves
lanceolate, acuminate. Capitula numerous, forming corymbose
paniculate inflorescence. Involucre broadly-campanulate, 5 mm long
and almost as much ‘in dia, glabrous; outer involucral bracts linear
subulate, 1-3, a third to a half as long as inner; inner bracts oblong-
linear with scarious border, obtuse. Ligulate florets yellow, about 5,
8-10 mm long and about 2 mm wide. Achenes glabrous, narrowly
ovate, 3 mm long and about 1 mm wide, finely ribbed, with narrow
black striations between ribs; pappus white-brownish. Flowering August;
fruiting September.
Slopes in middle mountain zone, in oak forests, near ditches and
irrigation canals.—Caucasus: Ciscaucasia? Endemic. Described from
Terek and Kuma rivers. Type in Leningrad.

27. S. racemosus (MB.) DC. Prodr. VI (1837) 358, ex p.; Ldb. FI.
Ross. II, 643; Boiss. Fl. or. III, 402.—S. racemosus L. puberus DC.
op. cit. (1837).—S. caspicus Less. in Linnaea, IX (1834) 170.—S.
pseudoracemosus Grossh. in Tr. Azerb. Otd. Zakavk. Fil. Akad. Nauk
SSSR, Bot. 1 (1933) 60; Grossh. Fl. Kavk. IV, 149.—Cineraria
racemosa MB. Tableau prov. sit. sur la céte occid. de la mer casp.
(1798) 119; Fl. taur.-cauc. I, 314, II, 573.—Cineraria caspica Pers.
syn. II (1807) 43.

Perennial. Rhizome short, thick, with long thin fibrous roots. Stem
solitary, at base covered with brown fibrous remnants of dead leaves,
finely sulcate, simple or somewhat branched in inflorescence, 15-70
cm high. Basal and lower cauline leaves with petioles shorter or longer
than lamina, lamina oblong-lanceolate or narrowly ovate, usually acute,
with sharp, irregular teeth along margin, stiff-hairy above, particularly
on veins, more densely beneath, 4-18 cm long and 1.5—3.0(6.0) cm
wide; upper leaves like basal but smaller, narrowly linear-lanceolate,
sessile. Capitula 3-25, in racemose inflorescence. Involucre about 10
mm long and 15 mm in dia, more or less pubescent; outer involucral
bracts oblong-linear, 6-12, a third to a half as long as inner ones;
inner bracts oblong lanceolate, scarious along margin, flat on spine.

732 Ligulate florets yellow, 10-20 mm long and 3-5 mm wide, 8-10.

698

Achenes terete, oblong, ribbed, glabrous, weakly lustrous, 4 mm long;
pappus brownish, almost as long as tubular florets. Flowering July to
August. (Plate XXXIV, Fig. 1).

Rubble slopes in subalpine and alpine zones, at 2,600—3,600 m.—
Caucasus: Ciscaucasia, Dagestan, Eastern Transcaucasia. Endemic.
Described from alpine zone of Glavnyi [Main] Caucasian Range. Type
in Leningrad.

Note. Marschall Bieberstein described two species under the name
Cineraria racemosa: one from the mountains, of the Caucasus “in
alpestribus caucasi,” and the other from the region south of Saratov.
Since the Caucasus was mentioned as the first locality, I retain the
name Senecio racemosus (MB) DC. for the Caucasian Mountain plant.

28. A. thyrsophorus C. Koch in Linnaea, XXIV (1851) 362;
Grossh. Fl. Kavk. IV, 149.—S. cyri C. Koch, ibid. 1851.—S.
schelkovnikovii Grossh. in Tr. Azerb. Otd. Zakavk. Fil. Akad. Nauk
SSSR, Bot. 1 (1933) 60; Gorssh. Fl. Kavk. IV, 149.—S. racemosus
var. schelkovnikovii Grossh. in Grossh. and Schischk. Plantae orient.
I (1924) 44; Grossh. Fl. Kavk. IV, 149.—S. racemosus var. araxina
Trautv. in Tr. Peterb. Bot. Sada, II (1873) 550.—S. racemosus var.
glabra DC. Prodr. VI (1837) 358.—Exs.: Pl. or No. 173.

Perennial. Rhizome short, thick, with long fibrous roots. Stem
solitary, basally covered with dark brown fibrous remnants of dead
leaves and also of leaf petioles, finely sulcate, solid glabrous or weakly
arachnoid-hairy, branched in inflorescence, 25-100 cm high. Basal
and lower cauline leaves with petioles more or less longer than lamina
and gradually expanded toward base; lamina lanceolate or narrowly
ovate, 8-20 cm long and 2.0-5.5 cm wide, with sharp teeth, larger
teeth at base, sometimes basally pinnatifid, subglabrous above, with
stiff hairs beneath; upper leaves smaller, linear-lanceolate, sessile, with
semiamplexicaul auricles. Capitula numerous, aggregated in corymbose
or racemose panicle. Involucre about 10 mm long and 15 mm in dia;
involucral bracts glabrous, white-ciliate, only apically and sometimes
along margin; outer involucral bracts linear, 6-12, a third to a half as
long as broadly lanceolate inner bracts. Ligulate florets 8-10, yellow,
8-20 mm long and 2—3(5) mm wide. Achenes angular-ribbed, glabrous,
about 4.5 mm long; pappus brownish, almost as long as tubular florets.
Flowering July to August.

Rubble and herb slopes.—Caucasus: Southern and Eastern
Transcaucasia. General distribution: Armenia and Kurdistan. Described
from Darichichag. Type was in Berlin; cotype in Leningrad.

Note. This species is very close to S. racemosus (MB) DC. but
differs from it by having a taller, subglabrous stem, almost glabrous

733 involucral bracts, shorter ligulate florets, and a paniculate inflorescence.

699

29. S. paucifolius Gmel. R. d. Russ]. I (1770) 171.—S. kirghisicus
DC. Prodr. VI (1837) 362; Fl. Yugo-Vost. Evrop. Ch. SSSR, VI, 371;
Kryl. Fl. Zap. Sib. XI, 2852.—Cineraria glabrata Ldb. Icon. pl. FI.
Ross. I (1829) tab. 94, non Swartz (1806) nec Senecio glabratus Hook.
and Arn. Bot. Beech Voy. (1841) 32.—Cineraria auriculata Ldb. FI.
Alt. IV (1833) 105, non S. auriculatus Desf. (1800).—S. racemosus
Ldb. FI. Ross. II, 643, ex p.; Schmalh. Fl. II, 88, p.—Branicia insignis
Andrz. ex Trautv. in A.H.P. VIII (1883) 476; Rogov. Obozr. Semenno.
i Vysshikh Spor. Rast. Kievsk. Uch. Okr. 302, nomen.—Ic.: Gmel.
S.G. loc. cit. tab. 38, f. 2.

Perennial. Rhizome with numerous, rather thick root strands. Stem
erect, solitary, branched in inflorescence, sulcate, whitish or slightly
violet, glabrous or with occasional hairs, 35—90 cm high. Basal leaves
numerous, ovate or lanceolate-ovate, narrowed into more or less long
petiole, including petiole 9-25 cm long and 1.0-—7.5 cm wide, unevenly
toothed, obtuse, pubescent beneath; lower cauline leaves identical with
basal leaves, upper leaves smaller, narrowly lanceolate or oblong,
sessile, with semiamplexicaul base. Inflorescence racemose-paniculate
or narrowly paniculate. Involucre 9 mm long and almost as much in
dia in upper part; outer involucral bracts linear, 12-15, a third to a
half as long as broadly linear inner bracts; all bracts weakly pubescent
or subglabrous. Ligulate florets 8-12, yellow, 12-14 mm long and
3.0-3.5 mm wide. Achenes terete, about 5 mm long and 1 mm wide,
ribbed, pubescent along ribs; pappus slightly longer than tubular florets.
Flowering July to August; fruiting September.

Alkaline meadows, sheep’s fescue steppes, occasionally on
solonchaks [salt marshes].—European part: Volga-Don, Lower Don,
Middle Dnieper, Black Sea Region, Upper Dniester (Podolia), Trans-
Volga, Lower Volga, Bessarabia; Western Siberia: Upper Tobol, Irtysh,
Altai; Soviet Central Asia: Aralo-Caspian Region (north), Lake
Balkhash Region (north). Endemic. Described from southern European
part of USSR. Type unknown.

30. S. schvetzovii Korsh. Tent. Fl. Ross. or. (1898) 519; Fl. Yugo-
Vost. Evrop. Ch. SSSR, VI, 370.—S. doria Ldb. Fl. Ross. II, 639, p.
non L.—S. auriculatus Desf. (1800).—S. racemosus Ldb. FI. Ross. II,
643, ex p.; var. microcephala Trautv. in Bull. phys.-math. Acad. Sc.
Pétersb. XII (1854) 351 and var. macrocephala Trautv. op. cit. (1854)
352.—S. doria B. biebersteinii Lindem. in Bull. Soc. Nat. Mosc. XL
(1867) 133; Elisabethgrad. 85; Prodr. Fl. Cherson. (1872) 113.—S.
biebersteinii Grec. Suppl. Consp. Fl. Romanei (1909) 91.—S. doria B.
macrophylla Schmalh. Fl. II (1897) 88 ex p.—Ic.: Fl. Yugo-Vost.
Evrop. Ch. SSSR, 691.

734

700

Perennial. Rhizome horizontally creeping or obliquely ascending,
about 1 cm thick. Stem erect, simple, branched only in inflorescence,
weakly arachnoid-hairy, sulcate, 60-100 cm high. Leaves glabrous,
glaucescent-green; lower leaves ovate, acute, gradually narrowed into
winged petiole, in basal leaves petiole almost as long as lamina or a
half as long, with irregularly toothed, often involute margin, lamina
15-30 cm long and 4.5-8.5 cm wide; middle cauline leaves sessile,
ovate-lanceolate or lanceolate, with round or somewhat cordate base,
unevenly toothed or almost entire. Inflorescence corymbose-paniculate;
capitula numerous, on 2-20 mm long peduncles, with 2-3 small linear
bracteal leaves. Involucre broadly campanulate, 5—6 mm long and 4—
6 mm in dia above; outer involucral bracts linear, a fourth to a third
as long as inner ones; inner bracts oblong, apically attenuate into
fimbriate-ciliate, sometimes deflexed appendage. Ligulate florets usually
5, 10 mm long and 4 mm wide. Achenes glabrous, ribbed, 3-4 mm
long. Flowering July to August.

Meadows, chalk slopes, forest edges, scrubs occasionally in saline
meadows and ravines.—European part: Middle Dnieper, Volga-Don,
Trans-Volga, Volga-Kama, Lower Volga, Lower Don, Black Sea
Region, Bessarabia, Upper Dniester; Western Siberia: Upper Tobol;
Soviet Central Asia: Aralo-Caspian Region (north). General
distribution: Balkans (Romania). Described from Bashkiria
(Davlekanovo). Type in Leningrad.

31. S. umbrosus Waldst. and Kit. Descr. et Icon. pl. rar. Hung. III
(1806) 232.—S. doria umbrosus Schmalh. FI. II (1897) 89.—Ice.:
Waldst. and Kit. op. cit. tab. 210; Hegi, Ill. Fl. VI, 2, 757; Jav. and
Csapody, Iconogr. Fl. Hung. (1933) 532.—Exs.: Fl. Hung. exs. No.
987, I and II.

Perennial. Rhizome about 1 cm thick. Stem 5—7 mm thick, white-
pubescent below, subglabrous above, erect, simple, branched only above
(in inflorescence), 50 cm high. Basal and lower cauline leaves ovate,
25-30 cm long and about 10 cm wide, subacute, long-attenuate toward
base, sessile, uniformly toothed, green above, paler beneath, on both
sides glabrous (only sometimes with occasional hairs on midrib); middle
leaves smaller, narrowly-ovate, 7-10 cm long and 2.0-3.5 cm wide,
sessile, amplexicaul; upper leaves lanceolate, acute, 2-5 cm long and
0.5—1.0 cm wide. Inflorescence paniculate-corymbose, with numerous
capitula. Involucre 12-13 mm long and 10 cm in dia; inner involucral
bracts narrowly ovate, with very wide, scarious border, acute, glabrous;
outer bracts linear, 4—S. Ligulate florets eight, yellow-orange. Achenes
glabrous. Flowering August.

IBS

701

Forest edges in river valleys.—European part: Upper Dniester.
General distribution: Balkans. Described from Arva River Valley
(Hungary). Type in Budapest.

Series 4. Racemuliferae Schischk.—Leaves almost entire, lower
leaves short-petiolate; ligulate florets 8-10, yellow.

32. S. racemulifer Pavl. in Vestn. Akad. Nauk. Kaz. SSR, No. 8
(1950) 21.—S. doriaeformis var. thyrsoidea Franch. in Ann. Sc. Nat.
VI, Bot. XV (1883) 311.—S. aksuensis Pavl. in Vestn. Akad. Nauk.
Kaz. SSR, No. 4 (73) (1951) 97.—Ic.: Pavl. op. cit. (1950) 21.

Perennial. Rhizome woody, vertical, many-headed above. Stems
few, weakly glaucous, glabrous, sometimes pubescent only in
inflorescence, simple or branched above, 45-70 cm high. Leaves ovate
or oblong—ovate, 5.0—8.5 cm long and 1.0—-3.5 cm wide, glabrous, obtuse
or short-acuminate, entire or less often with few teeth, narrowed toward
base; basal and lower cauline leaves on short (1 cm long) petiole,
withering before flowering, upper leaves sessile. Inflorescence
corymbose; capitula on 1.5—4.5 cm long peduncle, with narrowly linear,
0.7—1.0 cm long bracteal leaves. Involucre 6-8 mm long and 6-10 mm
in dia; outer involucral bracts 2-4, linear, 4-5 mm long, inner bracts
oblong-obovate, 6-8 mm long, acute, glabrous. Ligulate florets 8-10,
yellow, twice as long as involucre. Achenes not fully matured, 2-3 mm
long, obpyramidal, dark brown. Flowering June to August; fruiting
September.

River gravel beds in alpine and subalpine zones (2,300—2,400 m).—
Soviet Central Asia: Tien Shan (west). Endemic. Described from Talass
Alatau Range. Type in Alma-Ata; isotype in Leningrad.

Series 5. Paludosi Schischk.— Stem hollow, densely leafy; leaves
unevenly toothed or serrate with acute and upright teeth; ligulate florets
15-18.

33. S. paludosus L. Sp. pl. (1753) 870 s. str. var. excl.—S.
paludosus . vulgaris Ldb. Fl. Ross. II (1845) 640; Schmalh. FI. II.
89.—S. paludosus glabriuscula DC. Prodr. VI (1837) 353.—S.
paludosus &. nudiusculus Ldb. (1845) op. cit.—S. paludosus grandidens
Rupr. in Bull. XII (1854) 94; Rupr. Fl. ingr. 600.

Perennial. Rhizome long, horizontal or ascending, with long fibrous
roots at base of stem. Stem solitary, erect, simple, branched only in
inflorescence, glabrous or weakly pubescent, finely sulcate, hollow,
80-200 cm high. Leaves numerous, lower ones withering early, middle
and cauline leaves narrowly lanceolate, acute, gradually narrowed toward

736

702

base, with sharp upright teeth, dark green and glabrous above, lighter
beneath and sparsely pubescent, 10-15 cm long and 1-2 cm wide,
upper leaves smaller and narrower, sometimes almost entire. Capitula
numerous, in corymbose inflorescence at apices of stem; peduncles
weakly arachnoid-hairy. Involucre broadly campanulate, 8 mm long
and 10 mm in dia; outer involucral bracts 8-10, linear, usually recurved,
glabrous, a half as long as inner ones; inner bracts linear, with narrow
scarious margin, subacute, glabrous or weakly arachnoid-hairy. Ligulate
florets 15—16, 2-3 times as long as involucre, yellow or orange-yellow.
Achenes terete, glabrous, brownish, angular, 3.5-4.0 mm long.
Flowering July to August.

Along banks of lakes and marshes, rivers and seas, in willow
thickets.—European part: Ladoga-IImen, Baltic Region, Upper Dnieper.
General distribution: Scandinavia, Central Europe, Balkans (north).
Described from Europe from coastal marshes. Type in London.

34. S. tataricus Less. in Linnaea, IX (1834) 192.—S. lanatus S.G.
Gmelin R. d. Russl. I (1774) 155 non L. (1768) nec Scop. (1771).—
Cineraria aureus L. Sp. pl. 2 (1763) 1244 non Senecio aureus L.
(1753).—S. tomentosus Hos Fl. Austr. II (1831) 476, non Michx.
(1803).—S. auratus DC. Prodr. VI (1837) 348.—S. paludosus
hypoleucus Ldb. FI. Ross. II (1846-1848) 640.—S. paludosus auratus
Schmalh. Fl. II (1897) 89.—S. hypoleucus Ldb. and Wolosz. Fl. Polon.
exs. No. 445.—Ic.: S$.G. Gmel. loc. cit. tab. 35, fig. 1.—Fl. Yugo-
Vost. Evrop. Ch. SSSR, Fig. 690.—Exs.: Fl. Polon. exs. No. 445.

Perennial. Rhizome creeping, with long, thick, fibrous roots at
nodes. Stem solitary, erect, hollow, simple or branched above, densely
leafy, finely sulcate, arachnoid-woolly, 60-200 cm high. Lower cauline
leaves withering early; middle and upper leaves narrowly oblong,
lanceolate, or linear-lanceolate, acute, narrowed toward base, sessile,
unevenly toothed or serrate with acute, upright teeth, green above and
weakly arachnoid-woolly, floccose-white-tomentose beneath, 8-15 cm
long and 1-3 cm wide. Capitula 4—7 or numerous, at apices of stem on
arachnoid-hairy peduncles, in corymbose panicle. Involucre broadly
campanulate, about 8 mm long and 12-15 mm in dia; outer involucral
bracts linear, 8-10, a half as long as inner; inner bracts lanceolate,
acuminate, scarious along margin, with black spot at apex, 18-20; all
involucral bracts arachnoid-hairy. Ligulate florets yellow or orange-
yellow, about 18, 1.5—2 times as long as involucre. Achenes glabrous,
terete, ribbed, 3.5 mm long. Flowering June to July.

Floodplain meadows, on mounds among alders, banks of lakes.—
European part: Dvina-Pechora, Ladoga-IImen (Novgorod), Baltic
Region, Volga-Kama, Trans-Volga, Upper Dnieper, Upper Volga,

703

Plate XXXIV.
Senecio racemosus (MB.) DC., habit; 2—S. taraxicifolius (MB.) DC.,

TEM

1

habit.

739

704

Volga-Don, Middle Dnieper, Black Sea Region, Lower Don, Lower
Volga; Western Siberia: Ob River Area, Upper Tobol. General distri-
bution: Central Europe. Described from marshes along Ilek River. Type
was in Berlin.

Section 5. Oliganthi Boiss. FI. or. III (1875) 385 pp. emend. Sof.
in Izv. Akad. Nauk Azerb. SSR (1957) 88.—Plants perennial; leaves
pinnately cut or undivided; inflorescence corymbose-paniculate;
involucre cylindrical, 3-6 mm wide, usually with 4-6 inner and a few
outer involucral bracts; capitula few-flowered with 5—15 flowers, ligulate
florets absent, less often 1—3(5).

35. S. pojarkovae Schischk. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XV (1953) 409.—S. stenocephalus Boiss. FI. or. III (1875)
408, non Maxim. (1871); Grossh. Fl. Kavk. IV, 151.—Exs.: GRF No.
3247.

Perennial. Whole plant glabrous; rhizome short, obliquely
ascending, covered with long slender roots. Stem usually solitary,
somewhat sulcate, densely leafy, branched in inflorescence, 100-150
cm high. Basal leaves withering early, lower leaves short-petiolate,
lanceolate or narrowly lanceolate, acute or acuminate, narrowed toward
base, almost uniformly toothed with short, acute teeth; cauline leaves
identical with lower leaves, 6-24 cm long and 1-5 cm wide; uppermost
leaves shorter, linear-lanceolate, sessile or gradually narrowed into short
petiole. Capitula numerous, with 4—5 flowers, forming large corymbose
panicle; common peduncles obliquely upright, with very short,
indistinctly arachnoid hairs, some peduncles as long as, longer or shorter
than involucre, with few, linear, scale-like bracteal leaves. Involucre
campanulate, 4.5—6.0 mm long and 2.0—2.5 mm in dia; outer involucral
bracts 1-2, very small, sometimes absent; inner bracts 5, linear-oblong,
obtuse. Tubular florets whitish, with 4 teeth; ligulate florets lacking.
Achenes glabrous, terete, 3.0-3.5 mm long; pappus white, twice as
long as involucre. Flowering end of June to July.

Subalpine meadows, edges of mountain forests on river gravel
beds, at 1,100-2,100 m.—Caucasus: Ciscaucasia, Dagestan, Western
Transcaucasia. Endemic. Described from Dagestan from Dindidag
Mountain. Type in Geneva.

36. S. rhombifolius (Willd.) Sch. Bip. in Flora, XXVIII (1845)
499.—S. platyphyllus DC. Prodr. VI (1837) 352; Ldb. Fl. Ross II,
637; Boiss. Fl. or III, 409; Grossh. Fl. Kavk. IV, 151.—Cacalia
macrophylla MB. Fl. taur.-cauc. II (1808) 286, non Senecio
macrophyllus DC. (1837).—C. rhombifolia Willd. Enum. Horti Berol.
(1809) 850; DC. Prodr. VI, 331.

740

705

Perennial. Whole plant glabrous; rhizome horizontal, densely cov-
ered with fibrous root. Stem solitary, solid, sulcate, branched in inflo-
rescence, 100-120 cm high. Basal and lower cauline leaves long-peti-
olate, their laminas undivided, with unequal acute teeth, acuminate, at
base deeply cordate, 14-15 cm long and 20-25 cm wide; middle cauline
leaves like basal but smaller, with short, sometimes expanded petiole,
upper leaves lanceolate, basally attenuate, reduced. Capitula cylindri-
cal, with 5-8 flowers, numerous, in corymbose panicle.
Involucre 5 mm long and 2 mm in dia; outer involucral bracts linear-
subulate, 1-3, a third to a half as long as inner ones; inner bracts 4—
5, linear-oblong, obtuse. Ligulate florets absent. Achenes terete,
glabrous, weakly ribbed, about 3 mm long; pappus brownish, almost
as long as tubular florets. Flowering July to August; fruiting August to
September.

In scrubs, banks of mountain streams, mixed pine birch-beech and
mountain pine forests, talus, at 1,800—2,500 m in apline meadows.—
Caucasus: Ciscaucasus, Dagestan, Eastern Transcaucasia, Western
Transcaucasia. Endemic. Described from the Caucasus. Type in
Leningrad.

Economic Importance. The rhizomes and leaves contain an alka-
loid—platyphylline—that dilates the pupil.

37. S. platyphylloides Somm. and Lev. in Tr. Bot. Sada, XII
(1892) 153; Grossh. Fl. Kavk. IV, 251.—S. platyphylli var.
platyphylloides (Somm. and Lev.) Grossh. Opred. Rast. Kavk. (1949)
468.—S. similiflorus Kolak. Fl. Abkh. IV (1949) 252.—S. platyphylli
Boiss. Fl. or. II] (1872) 4098, quo ad pl. lazicam.—Ic.: Kolak. FI.
Abkh. IV, Plate XXV.

Perennial. Whole plant glabrous; rhizome thick, horizontal. Stem
solitary, erect, sulcate, branched above, 50-75 cm high. Basal and
lower cauline leaves long-petiolate, their laminas deltoid-reniform, with
sharp teeth, usually acute, at base deeply cordate, 10-17 cm long and
15-30 cm wide; middle cauline leaves identical to basal in shape,
short-petiolate, usually with large auricles; uppermost leaves lanceolate:
or ovate, sometimes almost entire, narrowed toward base. Capitula
numerous, with 10-15 flowers, 5-7 mm long and about 3 mm in dia.
Outer involucral bracts 1—3, linear-subulate, a third to a half as long
as inner; inner bracts linear-oblong, 5-8. Ligulate florets absent. Achenes
terete, glabrous, weakly ribbed, 4-5 mm long; pappus brownish, almost
as long as tubular florets. Flowering July to August.

Mountain birch forests, subalpine tall-herb meadows, Caucasian
rhododendron scrubs, at 1,500—2,500 m.—Caucasus: Ciscaucasia (west),
Western Transcaucasia. General distribution: Asia Minor (? Artvin

741

706

District, Pontic Range). Described from Danais-Paraul. Type in Flo-
rence.

Note. This species is very close to the preceding species, but has
morphological differences and a rather separate range (mainly Western
Transcaucasia).

38. S. paucilobus DC. Prodr. VI (1837) 352; Boiss. Fl. or III, 41;
Grossh. Opred. Rast. Kavk. 470.

Perennial. Whole plant covered with thin, appressed arachnoid-
hairs. Stems few, arising from thick rhizome, somewhat branched in
inflorescence, 50-60 cm high. Lower cauline leaves long-petiolate,
petiole twice as long as lamina; lamina broadly ovate, 7-8 cm long and
5-6 cm wide, green, arachnoid-hairy, densely whitish-arachnoid-hairy
beneath, usually pinnately cut, with 2 pairs of lobes, terminal lobe
larger, basally truncate or almost cordate, with uneven coarse teeth,
lateral lobes oblong, toothed, lower lobes shorter, remote; upper cauline
leaves smaller, short-petiolate. Inflorescence dense or lax corymb;
capitula numerous, conical-cylindrical, 7 mm long and 2 mm in dia,
glabrous. Outer involucral bracts few, obtuse, many times into inner
oblong bracts. Ligulate florets absent. Achenes terete, ribbed, glabrous,
about 3.5 mm long. Flowering July to August.

Stony slopes at about 2,300 m.—Caucasus: Southern Transcaucasia.
General distribution: Armenia and Kurdistan, Iran. Described from
route to India from Iran. Type in Geneva.

39. S. othonnae MB. FI. taur.-cauc. II (1808) 308, III, 570; Ldb.
Fl. Ross. II, 637; Boiss; Fl. or III, 410; Grossh. Fl. Kavk. IV, 153.—
S. rariflorus C. Koch, in Linnaea, XVII (1843) 47.—S. pinnatus Sch.
Bip. in Flora, XXVIII (1845) 499, non Poir. (1817).—S. othonnae var.
genuinus C. Koch and var. rariflorus C. Koch, in Linnaea, XXIV
(1851) 366.—Cacalia pinnata Willd. Enum. Horti Berol. (1809) 580.—
Jacobaea othonnae C.A.M. Verzeichn. Pfl. Cauc. (1831) 81.—Ie.:
Parsa, Fl. de Iran. III (1943) fig. 187.

Perennial. Rhizome thick, ascending, covered with long roots. Stem
glabrous, sulcate, branched in inflorescence, 80-175 (200) cm high.
Basal and lower cauline leaves short-petiolate, including petiole 40-50
cm long and 12—15(25) cm wide, their laminas oblong, simple pinnate
or almost twice pinnate, with 6-10 pairs of lobes, sessile and somewhat
decurrent on rachis, with deeply pinnatifid or pinnately cut into ovate-
oblong or oblong-lanceolate, acute lobes with irregularly toothed margin,
pubescent beneath, glabrous above; upper leaves sessile, smaller, with
narrower lobes. Capitula with 5—8 flowers, numerous, forming
corymbose panicle. Involucre cylindrical, 4 mm long and 2 mm in dia;

742

707

outer involucral bracts 1-3, linear, a third to a half as long as inner
ones; inner bracts oblong, with scarious margin, apically obtuse, with
black spot, dorsally pubescent. Ligulate florets yellow, 1-2, 8-10 mm
long and 2.0—2.5 mm wide. Achenes terete, angular-ribbed, pubescent,
2.0-2.5 mm long; pappus brownish, as long as tubular florets. Flowering
July to August; fruiting August to September.

Mountain forests, their edges banks of mountain streams, at 1,200-
2,300 m.—Caucasus: Ciscaucasia, Dagestan, Eastern, Western, and
Southern Transcaucasia, Talysh. General distribution: Armenia and
Kurdistan, Iran. Described from Georgia. Type in Leningrad.

Note. This species has been reported by Boissier and other au-
thors also for the Balkans, but it is replaced there by another similar
species, S. anomalus DC.

Section 6. Pseudo-Oliganthi Sof. in Izv. Akad. Nauk Azerb. SSR
(1957) 88.—Plants perennial; usually glabrous or subglabrous; leaves
undivided, with toothed or serrate margin. Inflorescence corymbose;
inner involucral bracts 6-12; capitula many-flowered (not less than
15); ligulate florets 5-8; achenes glabrous.

40. A. nemorensis L. Sp. pl. (1753) 870; Ldb. Fl. Alt. IV, 109;
Ldb. Fl. Ross. I, 641, p.—S. octoglossus DC. Prodr. VI (1837) 354;
Fisch. and Mey. Ind. sem. Horti Petropol. VIII (1841) 71; Kryl. FI.
Zap. Sib. XI, 2850.—M. Popov, FI. Sr. Sib. Il, 744.—S. nemorensis
var. turczaninovii Hara in Bot. Mag. Tokyo, LII (1938) 126.—S.
fuchsii* turczaninovii DC. op. cit. (1837) 353.—S. nemorensis var.
polyglossus Rupr. Fl. Samojed. cisural. (1845) 43.—S. nemorensis
octoglossus C. Koch, Taschenb. (1844) 284; Ldb. Fl. Ross. II, 641;
Kryl. Fl. Alt. III, 676.—S. nemorensus subsp. macer Printz. The veg-
etation of the Siberian Mongolian frontiers. (1921) 428.—S. obscurus
Fisch. ex Ldb. Fl. Ross. II (1845-1846) 638; ?Turcz. in Bull. Soc.
Nat. Mosc. XX (1847) 220.—S. octoglossus macer DC. op. cit. (1837)
354.—Ic.: Rchb. Ic. Fl. Germ. XVI (1853) t. 971; Printz, op. cit. tab.
XIV, fig. 1 (photo).—Exs.: GRF No. 1224; Fellm. Pl. arct. No. 136.

Perennial. Rhizome short, creeping, covered with fibrous root. Stem
erect, branched above, weakly hairy or subglabrous, 50-150 cm high
and up to 1 cm thick in lower part, often reddish. Leaves ovate-
lanceolate or ovate, 5-22 cm long and 1-7 cm wide, with short and
straight teeth, less often teeth long, 3-8 mm long and curved (f.
longidentatus Serg.), pubescent beneath or subglabrous; lower leaves
on petioles a half as long as lamina; middle leaves on short and

*fuhsii in Russian original—General Editor.

743

708

broadened petioles; upper leaves smaller, sessile. Inflorescence corym-
bose. Involucre 5-6 mm long and 4-5 mm in dia; involucral bracts
subglabrous or slightly hairy; outer bracts 4—5, linear, as long as inner
ones; inner bracts lanceolate, apically brownish and ciliate. Ligulate
florets 8—10(13), ligules 12-18 mm long and 4-5 mm wide. Achenes
glabrous, ribbed, 4.0-4.5 mm long. Flowering June to August.

In coniferous and birch aspen forests, their edges, in pine
woodlands, scrubs, meadows along edges of marshes, river banks,
mountains ascending almost to the upper treeline.—Arctic: Arctic
Europe (Kanin Peninsula, Bolshezemelskaya tundra, Arctic Urals—
Usa and Sablya rivers); European part: Dvina-Pechora, Karelia-
Lapland, Baltic Region? Ladoga-IImen, Volga-Kama, Upper Dnieper,
Trans-Volga, Upper Dniester; Western Siberia: All regions; Eastern
Siberia: Angara-Sayans, Yenisei, Dauria, Lena-Kolyma; Far East: Zeya-
Bureya, Ussuri, Uda River Area, Okhotsk, Sakhalin (and Kuril Is-
lands); Soviet Central Asia: Aralo-Caspian Region (north), Dzhungaria-
Tarbagatai, Tien Shan. General distribution: Central Europe,
Scandinavia, Mongolia, Northeastern China. Described from Germany
and Siberian forests. Type in London.

Note. Given its eight ligulate florets, De Candolle proposed to
name this species S. octoglossus instead of S. nemorensis. But in
Linnaeus’s description of S. nemorensis it is also indicated that this
species has eight ligulate florets. It seems to me that the name pro-
posed by Linnaeus should be retained.

41. S. iljinii Schischk. sp. nov. in Addenda, XXV, 882.

Perennial. Stem erect, simple, branched only in inflorescence, solid,
pubescent with white crisped hairs, 40-50 cm high, densely leafy. Leaves
ovate, sessile or short-petiolate, acute, irregularly toothed with large,
usually obtuse teeth, dark green above, lighter beneath, subglabrous or
scarcely arachnoid-hairy, 8-12 cm long and 3-5 cm wide. Inflorescence
paniculate, with numerous capitula; peduncles densely white-pubescent.
Involucre about 5 mm long and almost as much in dia; outer involucral
bracts 3-4, linear, slightly longer than inner bracts and appressed to
them; inner bracts ovate, subacute, 8-10, like outer bracts, dorsally
covered with short and curly hairs, ciliate on margin. Ligulate florets
8, yellow, 3 times as long as involucre. Young achenes glabrous, 3-4
mm long. Flowering August.

Mountain slopes.—Soviet Central Asia: Dzhungaria-Tarbagatai.
Endemic. Described from Dzharkent District from Tyshkan River. Type
in Leningrad.

744

709

42. S. propinquus Schischk. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XV (1953) 406.—S. fuchsii Grossh. Fl. Kavk. IV (1934)
152, non Gmel.—S. jacquinianus Grossh. Fl. Kavk. IV, 151, non Rchb.

Perennial. Rhizome short, densely covered with fibrous roots. Stem
solitary, straight or indistinctly flexuous, slightly violet, glabrous or
sparsely hairy, sulcate, densely leafy, branched in inflorescence, 80-—
150 cm high. Basal leaves withering early; cauline leaves ovate, acute,
or subacuminate, narrowing toward base, sessile; middle cauline leaves
10-20 cm long and 3-8 cm wide, cartilaginously toothed, sometimes
with 2 teeth, short-ciliate between teeth, dark green above and usually
glabrous, paler beneath and pubescent along veins, less often
subglabrous. Inflorescence corymbose; branches of inflorescence and
peduncles arachnoid-hairy; capitula mostly numerous. Involucre 5 mm
long and 3.5-4.0 mm in dia; outer involucral bracts about 5, narrowly
linear, longer than inner bracts, crisped-ciliate along margins; inner
bracts oblong, subobtuse, with an indistinct dark spot at apex, scarious
along margin and sparsely hairy. Ligulate florets 5, yellow, 12-14 mm
long and 2-3 mm wide. Achenes terete, glabrous, ribbed, about 4 mm
long; pappus white or brownish, almost as long as tubular florets.
Flowering June to July; fruiting August to September.

Mountain pine, birch, pine-birch-beech and fir-spruce forests, their
edges, mountain meadows, thickets of rhododendron and box trees, at
800—2,400 m.—Caucasus: Ciscaucasia, Dagestan, Eastern and Western
Transcaucasia. Endemic. Described from Kazbek Mountain. Type in
Leningrad.

43. S. lapsanoides DC. Prodr. VI (1837) 348 (“lampsanoides’’);
Ldb. Fl. Ross. II, 633; Boiss. Fl. or. III, 408; Grossh. Fl. Kavk. IV,
150.—Cineraria parviflora MB. FI. taur.-cauc. II (1808) 316; Willd.
Enum. Horti Berol. (1808) 893, non S. parviflorus All. (1785).—Exs.:
GRF No. 1328.

Perennial. Whole plant glabrous; rhizome obliquely ascending, 0.8
mm thick. Stem erect, finely sulcate, simple, branched only in
inflorescence, 35-120 cm high. Basal and lower cauline leaves with
petioles longer than lamina, gradually expanded toward base into
semiamplexicaul sheath, withering early, their laminas undivided, ovate-
cordate, green and lustrous above, paler beneath, acute or subobtuse,
with broad, somewhat obtuse teeth, 4-8 cm long and 1.5—12 cm wide;
middle and upper leaves sessile or short-petiolate; with broad
amplexicaul auricles, oblong-lanceolate or lanceolate, acute, with
deltoid, acute teeth. Capitula 4-50, with 15-20 flowers. Involucre
obconical, 8 mm long and 4 mm in dia; outer involucral bracts lacking,
inner oblong linear, obtuse or subacute, with very short hairs and usually

74

nn

710

a black spot at apex, membranous on margin. Ligulate florets yellow,
5-6, 8-10 mm long, and 2-3 mm wide. Achenes ribbed, glabrous,
oblong-terete, 4.5 mm long; pappus brownish, slightly shorter than
tubular florets. Flowering July; fruiting August.

Base of rocks, on cliffs in mountain birch and pine forests in
subalpine and alpine zone, at 1,500-2,400 m.—Caucasus: Ciscaucasia,
Central Caucasus, Eastern Transcaucasia, Dagestan. Endemic. Described
from Mount Kazbek. Type in Geneva.

44. S. quinqueligulatus Wink. in Del. sem. quae hortus bot. imp.
Petropol. pro mutua commutatione offert (1881) 15; O. and B. Fedtsch.
Perech. Rast. Turk. IV, 218.

Perennial. Root adventitious. Whole plant subglabrous. Stem
branched from base, about 40 cm high. Leaves oblong or ovate, basally
narrowed into petiole, unevenly toothed and indistinctly ciliate, 3-8
cm long and 1—5 cm wide, somewhat acuminate. Capitula at apices of
stem and branches, forming lax panicle. Involucre 2-rowed; outer
involucral bracts 2-3, linear, short-ciliate along margin, longer than
inner; inner bracts ovate, acuminate, dark brown. Ligulate florets 5,
yellow, 2.0—-2.5 times as long as involucre. Achenes glabrous. Flowering
June to July.

Mountain slopes.—Soviet Central Asia: Dzhungaria-Tarbagatai
(Dzhungarian Alatau-Kashgaria), Tien Shan (Muzart and Issyk-Kul).
Endemic. Described from cultivated specimens raised from seeds
obtained from Dzhungarian Alatau (Kashgaria) and Tien Shan (Muzart).
Type in Leningrad.

Note. Until now this plant is known only from specimens raised in
the St. Petersburg Botanical Garden from seeds obtained by Regel
from the Dzhungarian Alatau and Tien Shan. In 1959, one more
specimen of this plant was obtained from the Main Botanical Garden
of the Academy of Sciences in Moscow, grown from seeds collected
near Lake Issyk-Kul (Tien Shan).

45. S. fuchsii K. Gmel. Fl. Badens. III (1808) 444.—S. nemorensis y.
fuchsii Ldb. Fl. Ross. II (1844-1846) 641; Boiss. Fl. or. III, 408.—?S.
ovatus Willd. Sp. pl. III (1800) 2004.—Jacobaea fuchsii C.A.M. Verzeichn.
Pfl. Cauc. (1831) 81, quo ad nom.—Ic.: Rchb. PI. crit. III, t. 293.

Perennial. Rhizome slender, horizontal or ascending, covered with
slender and long roots. Stem solitary, angular-sulcate, more or less
purple, weakly flexuous, branched in inflorescence, 70-100 cm high.
Basal leaves withering early; cauline leaves numerous, short-petiolate,
ovate-lanceolate, 15-20 cm long and 4—6 cm wide, dark green above,
lighter beneath, glabrous on both sides, acuminate, toothed, sometimes

746

711

with 2 teeth, with acute, deltoid teeth, short-ciliate between teeth;
uppermost leaves linear-lanceolate, smaller. Capitula more or less
numerous, forming corymbose inflorescence. Involucre 6 mm long and
about 4 mm in dia, glabrous; outer involucral bracts 5-6, narrowly
linear, somewhat shorter or slightly longer than inner ones; inner bracts
linear, 9-10; all bracts with blackish apices. Ligulate florets yellow, 5,
linear, 12 mm long and about 2 mm wide. Achenes glabrous, terete,
about 5 mm long. Flowering July to August, fruiting September.

Open forests and edges.—Upper Dniester (Carpathians). General
distribution: Central Europe, Balkans. Described from Alsace. Type in
Karlsruhe.

46. S. fluviatilis Waller. in Linnaea, XIV (1840) 64; Bordz. in
Vestn. Tifl. Bot. Sada, 4-5 (1930) 64.—Grossh. Fl. Kavk. IV, 151.—
S. sarracenicus L. Sp. pl. (1753) 871, ex. p.; Ldb. Ross. II, 640;
Schmalh. FI. II, 89; Kryl. Fl. Zap. Sib. XI, 2850.—S. sarracenicus B.
sclerophyllus C. Koch, in Linnaea, XXIV (1850) 359.—S. hajastanicus
Sof. in Izv. Akad. Nauk Azerb. SSR, I (1957) 86.—Jacobaea
sarracenica Moench. Meth. (1794) 588.—Ic.: Sof. loc. cit. Fig. 2;
Syreistsch. Ill. Fl. Mosk. Gub. III, 28 (1910).—Exs.: GRF No. 1478.

Perennial. Rhizome creeping or ascending. Stem erect, densely
leafy, sparsely branched and sparsely hairy above, glabrous, elsewhere,
sulcate, 100-200 cm high. Leaves glabrous, sessile, elliptical-lanceolate
or almost lanceolate, 12-20 cm long and 1.5-4.0 cm wide, acute,
cuneately narrowed toward base, serrate-toothed and short-ciliate, teeth
acute, antrorse. Inflorescence corymbose; capitula numerous. Involucre
pubescent, 6 mm long and 3.5 mm in dia, 2-rowed, campanulate-
cylindrical; outer involucral bracts 3-5, linear, as long as inner ones;
inner bracts lanceolate, apically ciliate and with black spot. Ligulate
florets 6-8, bright yellow, ligule linear-oblong, 1.0—1.2 cm long and
2.5-3.0 cm wide. Achenes ribbed, 4 mm long, usually glabrous.
Flowering June to July; fruiting August to September.

Shrubby thickets along riverbanks, riverine meadows and willow
stands, edges of marshes.—European part: Dvina-Pechora (south),
Ladoga-IImen, Baltic Region, Upper Volga, Volga-Kama, Trans-Volga,
Upper Dnieper, Volga-Don, Middle Dnieper, Lower Don, Black Sea
Region; Caucasus: Southern Transcaucasia; Western Siberia: All
regions; Eastern Siberia: Angara-Sayans; Soviet Central Asia: Aralo-
Caspian Region, Dzhungaria-Tarbagatai (rarely). General distribution:
Atlantic and Central Europe, northern Balkans. Described from southern
border of Hartz [Mountains]. Type in Berlin.

Note. Recently, Sofieva described the new species S. hajastanicus
Sof. from the Armenian SSR, of the S. fluviatilis Wallr. complex. The

747

712

range of this new species is far removed from the range of the latter
species, and it would appear that establishing the new species S.
hajastanicus Sof. was completely justified. However, a careful study
of both these species revealed that it is impossible to find any morpho-
logical differences between them. In the comparison, the author men-
tions narrower leaves and shorter and comparatively fewer ligulate
florets. But the indicated characters are not confirmed from the mate-
rial. The leaves are sometimes broader, sometimes narrower; the dif-
ferences in the number of ligulate florets and their sizes are not sup-
ported by the author with any numerical data. I do not think it possible
to distinguish a separate species without morphological differences
only on the basis of its range alone. It would be more appropriate to
consider S. fluviatilis Wallr. as a relict in Armenia, persisting since the
Ice Age. In fact in Southern Transcaucasia, many species are known,
that in no way differ from the typical northern species and represent
a group of disjunct northern relicts.

Section 7. Extremiorientales Schischk. sect. n. in Addenda XXV,
883.—Plants 1-2 m high, with palmate or undivided leaves, often with
large auricles. Capitula numerous, small; involucre about 5 mm long
and as much in dia; achenes glabrous, 3-5 mm long.

47. S. cannabifolius Less. in Linnaea, VI (1831) 242; DC. Prodr.
VI, 349; S. Turcz. Fl. baic.-dahur. I, 92.—S. palmatus Pall. Reise, III
(1776) 321, nom. nud.; Ldb. Fl. Ross. II, 636, descr. non La Peyrousse
(1818) nec Less. (1823); Kom. Fl. Kamch. III, 331; Hult. Fl. of
Kamchatka, IV, 199.—Solidago palmata Pall. Reise. III (1776) 437,
nom. nud.—Senecio dahuricus Fisch. ex DC. Prodr. VI (1837) 349,
pro syn.; Turcz. in Bull. Soc. Nat. Mosc. XX, 2, 19, non Schultz Bip.

Perennial. Rhizome short, covered with numerous slender roots.
Stem erect, simple, finely sulcate, glabrous, 1-2 m high. Leaves almost
sessile, cuneately narrowed toward base, and with or without round-
falcate auricles, oblong-lanceolate, acuminate, with 2-3 pinnately
divided lobes on each side, serrate, with upright teeth, decurrent, dark
green and glabrous above, lighter beneath, 8-10 cm long and 1-2 (5)
cm wide. Inflorescence at stem apex, paniculate corymbose; capitula
on more or less long, slender; peduncles, with small subulate bracteal
leaves. Involucre 5 mm long and 5 mm in dia above; inner involucral
bracts ovate-lanceolate, sparsely pubescent, subacute. Ligulate florets
5, yellow, 10 mm long and 2 mm wide. Achenes glabrous 3.5-4.0 mm
long; pappus ginger-colored. Flowering July.

Scrubs, coastal forests, floodplain meadows, reed thickets, edges
of deciduous forests.—Eastern Siberia: Dauria, Lena-Kolyma; Far East:

748

#15

Zeya-Bureya, Ussuri, Kamchatka, Sakhalin (Kuril Islands). General
distribution: China, Japan, Beringia. Described from Kamchatka and
Bering islands. Type in Berlin.

Economic Importance. In spring, the rhizome of the plant is used
in raw form as food by the people of Kamchatka, and its leaves, dried
and ground to a powder, are sometimes added to food.

48. S. litvinovii Schischk. nom. nov.—S. otophorus Maxim.
Mélang. biol. VIII (1871) 11, non Wedd. (1855); Kom. Fl. Manchzh.
III. 708; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1057.

Perennial. Rhizome short, covered with numerous slender roots.
Stem erect, simple, finely sulcate, glabrous, 80-120 cm high. Basal
and lower cauline leaves withering early, more or less long-petiolate,
20 cm long and 5-7 cm wide; middle cauline leaves ovate, dark green
and glabrous above, lighter beneath and crisped-hairy, particularly on
veins, acuminate, unevenly toothed with sharp, upright teeth, narrowed
toward base into short petiole or sessile, with 2 oblong or suborbicular
auricles recurved and adpressed to stem; upper leaves smaller, lanceolate,
uppermost leaves linear, almost entire. Capitula numerous, in corymbose
panicle, inflorescence branches and peduncles pubescent. Involucre
campanulate, 5 mm long and about 5 mm in dia; outer involucral
bracts about 6, linear, pubescent, a third to a half as long as inner;
inner bracts oblong, acute, with narrow scarious margin, dorsally
pubescent. Ligulate florets 5-8, yellow, 10-12 mm long and 3 mm
wide. Achenes terete, ribbed, glabrous, 3 mm long and about 1 mm
wide; pappus brownish, 8-9 mm long. Flowering July to August; fruiting
September.

Mixed montane forests, stream banks, edges of alder forests.—Far
East: Ussuri. General distribution: Northeastern China, Japan.
Described from vicinity of Vladivostok and from Japan. Type in
Leningrad.

Section 8. Carniolici Schischk. sect. nov. in Addenda XXV, 883.—
Basal leaves long persistent, pinnately divided, with broadly linear
lobes; inner involucral bracts with black spot at apex; ligulate florets
2-4, dark yellow; achenes glabrous; pappus sordid brown.

49. S. carniolicus Willd. Sp. pl. III, 3 (1800) 1993; DC. Prodr.
VI, 356.—S. incanus Scop. Fl. Carn. ed. 2 (1772) non L. (1753).—S.
incanus subsp. carniolicus (Willd.) Br. Bl. efr. Dostal, Kvetena CSR
(1950) 1633.—S. incanus var. carniolicus Heat Fl. Engadine (1841)
tab. 121.—Jacobaea carniolica Schrank in Denkschr. Acad. Muench.
(1814) 316.—Ic.: Rchb. Ic. Fl. Germ. XVI, tab. 698 (1853); Hegi and

749

714

Dunc. Alp. Fl. (1913) tab. 29; Dostal, op. cit. Fig. 537, 2; Javorka and
Csapody, Iconogr. Fl. Hung. (1933) 530.—Exs.: Fl. Hung. exs. Cent.
1, No. 351; Fl. exs. Reipubl. Bohem.-Sloven. No. 959; Fl. Polon. exs.
No. 351.

Perennial. Rhizome ascending, 0.4—0.5 cm thick. Stem ascending
at base or erect, simple, arachnoid-floccose-hairy, slightly violet, finely
sulcate, 6-20 cm high. Basal leaves more or less numerous, long-
persistent, pinnately divided (not up to midrib), with petiole usually
longer than lamina, subglabrous above, finely arachnoid-hairy beneath
and bluish-dark bluish, including petiole, 5—10 cm long and 1-2 cm wide;
leaf lobes broadly linear, obtuse, with occasional teeth or entire; cauline
leaves 2—4(8), like basal, but smaller, lower among these usually long-
petiolate, upper sessile. Inflorescence corymbose or lax, head of 5—15
capitula on short peduncles, arachnoid-hairy and with linear, acuminate,
bracteal leaves. Involucre 5 mm long and 6-7 mm in dia, 2 rowed;
inner involucral bracts lanceolate, subacute, dorsally finely arachnoid-
hairy, with dark violet or almost black spot at apex. Ligulate florets 2—
4, dark yellow, somewhat longer than involucre; pappus dirty brown.
Achenes terete, glabrous, about 3 mm long. Flowering July to August.

Stony slopes, granite rocks, and meadows in alpine zone at 1,700—
2,300 m.—European part: Upper Dniester (Carpathians). General dis-
tribution: Balkans, mountains of Central Europe. Described from
Carnolia. Type in Berlin.

Section 9. Montani Schischk. sect. n. in Addenda XXV, 883.—
Basal leaves with petioles as long as lamina or 2-3 times as long,
ovate, irregularly toothed with coarse teeth or pinnately divided; outer
involucral bracts 2—8, linear; ligulate florets yellow, about 10-12;
achenes glabrous.

50. S. tianschanicus Rgl. and Schmalh. in Tr. Peterb. Bot. Sada,
VI'CL87 9) Ste.

Perennial. Rhizome short, with numerous long roots. Stem erect,
sometimes at base ascending, simple, 5—20 cm high, glabrous or
arachnoid-hairy, sulcate. Lower leaves ovate or ovate-lanceolate, with
long petioles often many times as long as lamina, their laminas 1.5—
4.0 cm long and 0.6—1.2 cm wide, obtuse or subacute, glabrous above,
usually arachnoid-hairy beneath, irregularly toothed with coarse teeth
or deeply pinnatifid; upper leaves sessile. Capitula in corymbose
inflorescence or even solitary. Involucre 7-8 mm long and 5—6 mm in
dia; outer involucral bracts 2—8, linear, a fourth to two-thirds as long
as inner; inner bracts broadly linear; all bracts weakly pubescent or
even subglabrous. Ligulate florets yellow, about 10, twice as long as

750

715

involucre. Achenes glabrous, ribbed, about 3.5 mm long. Flowering
July to August; fruiting September.

Old moraines, gravel beds, scrubs, forest edges at treeline, on
stony slopes.—Soviet Central Asia: Tien Shan. General distribution:
Dzhungaria-Kashgaria (Sin-Tsian). Described from Tien Shan. (Tersk
Alatau near Sary-Kungei). Type in Leningrad.

51. S. renardii Winkl. in Trautv. Rg]. Maxim. and Winkl. Decas
pl. nov. (1882) 6.—Exs.: GRF No. 4199.

Perennial. Rhizome obliquely ascending, 0.5—1.0 cm thick, whole
plant densely floccose-hairy. Stems many, often violet below, erect,
60-90 cm high, somewhat branched from middle. Basal and lower
cauline leaves obovate or ovate-lanceolate, with petiole as long as
lamina or longer, expanded at base, semiamplexicaul, their laminas 8
cm long and 4—6 cm wide, subobtuse, irregularly toothed, sometimes
pinnately divided at base; upper leaves smaller, sessile, semiamplexicaul,
deeply toothed or almost lyrate. Inflorescence corymbose panicle;
capitula usually numerous. Involucre 7 mm long and about 10 mm in
dia; outer involucral bracts about 5, linear-subulate, half as long as
oblong-lanceolate, acuminate, inner bracts. Ligulate florets about 12,
yellow, 1.5-2 times as long as involucre. Achenes glabrous, ribbed,
about 3.5 mm long. Flowering July to August; fruiting September.
(Plate XXXIII, Fig. 2).

Juniper stands at treeline, stony beds of streams, on loess slopes
and rocks at 2,000—3,500 m.—Soviet Central Asia: Pamiro-Alai Region.
General distribution: Indo-Himalayas (Kashmir). Described from
Baldzhuan, Darvaz and other places. Type in Leningrad.

Section 10. Scaposi (Hoffm.) Schischk. ex Kryl. Fl. Zap. Sib. XI
(1949) 2840; Popov, FI. Sr. Sib. I], 748.—Scaposi Hoffm. in
Pflanzenfam. IV, 5 (1889) 297 p. p.; Addenda XXV, 883.—Perennial
low herbs; stem always with single capitulum; leaves pinnate or lyrate;
high-mountain wholly glabrous plants.

52. S. resedifolius Less. in Linnaea, VI (1831) 243; Ldb. Fl. Ross.
II, 631; Turcz. Fl. baic.-dahur. II, 88; Kryl. Fl. Zap. Sib. XI, 2840.—
S. resedifolius B. lyratus DC. Prodr. VI (1837) 347.—S. resedifolius
Q. genuina Vv. multicaulis 6. multiflkora Rgl. in Bull. Soc. Nat.-Mosc.
XL (1867) 429.—S. lyratus Turcz. in Bull. Soc. Nat. Mosc. V (1838)
non L. fil. (1791).—Cineraria heterophylla Fisch. in Mém. Soc. Nat.
Mosc. III (1812) 79, non Senecio heterophyllus Thunb. (1794).—C.
lyrata Ldb. in Mém. Acad. Sc. Pétersb. V (1815) 576.—Ic.: Rchb. Pl.
crit. II, t..101, f. 211; Popov. Fl. Sr. Sib. II, Fig. 88.

75

=

716

Perennial. Whole plant glabrous; rhizome thick. Stems solitary or
few, simple, 5-20 cm high, always terminating into single capitulum.
Basal leaves almost round, with 10-20 cm long and as wide lamina,
with coarse and round teeth, less often almost entire, abruptly narrowed
at base into long petiole; lower and middle cauline leaves lyrate,
pinnately divided or even pinnately lobed; uppermost leaves lanceolate
or linear, entire. Involucre 8 mm long and 10-12 mm in dia, usually
violet with upper part of stem, in single whorl, with broadly linear
involucral bracts. Ligulate florets yellow, often with violet veins, 2.5
times as long as involucre. Achenes glabrous, brownish, 3.5—-4.0 mm
long, finely ribbed.

Moss-lichen and rubbly tundras near glaciers and snow in apline
zone (2,500—3,000 m), and in Arctic.—Arctic: Arctic Europe, Novaya
Zemlya, northern Urals, Arctic Siberia, Chukotka, Anadyr; Western
Siberia: Altai; Eastern Siberia: Angara-Sayans, Dauria, Lena-Kolyma;
Far East: Okhotsk, Kamchatka. General distribution: Mongolia, Arctic
America. Described from shores of Gulf of St. Lawrence. Type in
Berlin(?).

53. S. carpathicus Herb. Addit. fl. Galic. (1831) 43; Boiss. FI. or.
Ill, 394; Szafer, Kulcz. Pawl. Rosliny Polskie (1953) 705.—S.
abrotanifolius Wahlb. Fl. Carp. (1814) 266 non L.—S. abrotanifolius
var. carpaticus Hayek. Prodr. Fl. Balc. II (1931) 679.—S. monocephalus
Schur. Enum. plant. Transsilv. (1866) 349.—Herbichia carpathica
Steud. Nomencl. ed. 2, I (1840) 750.— Senecillis carpathica Schott in
Nym. and Kotschy, Garten-Zeit. Berlin, III (1884) 572.—Ic.: Prodan,
Fl. determ. et desct. pl. Roman. II, tab. 111 (1923); Javorka and
Csapody, Iconogr. Fl. Hung. 531 (1933), tab. 38 (1934).—Exs.: PI.
Polon exs. No. 276.

Perennial. Rhizome long, ascending, 2—3 mm thick. Stems many or
solitary, basally ascending, usually purple, simple, glabrous, 10-15 cm
high. Leaves crowded in lower half of stem, broadly ovate, 2.5—-3.0 cm
long and about 1.5 cm wide, with petioles almost as long as lamina,
glabrous, twice or almost thrice pinnately cut, terminal lobes lanceolate
or linear, 2-5 mm long and 0.5—1.5(2.0) mm wide; middle and upper
cauline leaves smaller, simple pinnate, with linear lobes, sessile, often
brownish, uppermost leaves linear, acuminate, undivided. Capitulum
solitary, at stem apex. Involucre 5-8 mm long and 1.0—1.5 cm in dia;
outer involucral bracts 2—5, linear, shorter than inner ones; inner bracts
linear-oblong, white-scarious along margin, dorsally somewhat dark.
Ligulate florets orange-yellow, oblong, with 3 teeth in upper part, 10
mm long and 1.5-2.0 mm wide. Achenes terete, glabrous, 2.5 mm
long. Pappus as long as florets. Flowering August; fruiting September.

752

717

Alpine grass-mixed herb meadows, at about 2,000 m.—European
Part: Upper Dniester (Carpathians, Chernaya Gora [Black Mountain],
Petros, and other mountains). General distribution: Western Carpathians,
Tatry. Described from the Carpathians.

Section 11. Karelinoidei O. and B. Fedtsch. Perech. Rast. IV
(1911) 220 nom.; Addenda XXV, 883.—Perennial plants with almost
entire, ovate or lanceolate-ovate leaves; inner involucral bracts 8, outer
bracts 1-2, or even absent; ligulate florets 3-5, yellow.

54. S. karelinioides Winkl. in Tr. Peterb. Bot. Sada, XI (1890)
280.

Perennial. Rhizome woody. Stems numerous, glabrous, only in
upper part floccose-hairy, whitish, sulcate, erect, flexuous, 25-30 cm
high, leaf axis floccose-hairy. Basal leaves withering early, cauline
leaves ovate or lanceolate-ovate, sessile, paired toward apex, almost
entire, 5-6 cm long and 3 cm wide; upper leaves smaller; bracteal
leaves linear or subulate. Capitula numerous, forming broadly
corymbose inflorescence. Involucre campanulate-cylindrical, 6 mm long
and 6 mm in dia in upper part; outer involucral bracts 1-2, or even
absent, linear; inner bracts 8, oblong, herbaceous, with wide scarious
border and finely ciliate in upper part, sharply arachnoid-hairy on dorsal
side, connate at base. Ligulate florets 3-5, about 6 mm long and 3 mm
wide. Achenes obconical, glabrous, ribbed, 2-3 mm long, blackish.
Flowering July.

Mountain slopes.—Soviet Central Asia: Pamiro-Alai. Endemic.
Described from vicinity of the village of Dzhegen. Type in Leningrad.

Note. So far only one locality is known. Unfortunately, the collec-
tor (Skornyakov) did not mention the habitat conditions.

Section 12. Eriopappus Dumort. Fl. belg. (1827) 65.—Cineraria
sect. Holoseris Rchb. Fl. Germ. exc. (1832) 341.—Senecio sect.
Holoserides DC. Prodr. VI (1837) 362.—Stems hollow; inner involucral
bracts numerous, outer bracts absent; ligulate florets 20-25, yellow;
achenes glabrous; pappus at flowering slightly longer than tubular
florets, during fruiting strongly elongated. Annual or biennial plants.

55. S. arcticus Rupr. Fl. Samojed. cisural. (1845) 44.—L. congestus
B. intermedius DC. Prodr. VI (1837) 363.—S. palustris (L.) Hook. FI.
bor. am. I (1833) 334, non Velloso (1827); DC. Prodr. VI (1837) 363;
Turcz. Fl. baic.-dahur. II, 99; Ldb. Fl. Ross. II, 648; Schmalh. FI. II,
84 (paluster); Kryl. Fl. Zap. Sib. XI, 2858; Samojed, cisural. 44; Kryl.
Fl. Zap. Sib. XI, 2859.—S. paluster var. genuina Trautv. Fl. Bogan.

753

718

(1856) 162; Tr. Peterb. Bot. Sada, VI (1879) 25..—S. palustris. ~
congestus Hook. Fl. bor. am. I (1833) 334; Ldb. op. cit. 648; Hult. FI.
of Kamtch. IV, 202.—S. tubicaulis Mansf. in Fedde, Repert. XLVIII
(1940) 264; Ej. Verzeichn. der. Farn und Bliitenpflanzen des Deutschen
Reiches (1940) 253; J. Dostal, Kvétena CSR, 1626.—S. congestus var.
palustris (L.) Nyl. in Uppsala universitets Arsskrift. 7 (1945) 319.—
Othonna palustris L. Sp. pl. (1753) 924.—Cineraria palustris L. Sp.
pl. (1763) 1243; Ldb. Fl. Alt. IV, 102.—C. unctuosa Gilib. Fl. Lith.
III (1781) 211; Exerc. phytol. I (1792) 172, nom. illeg.—Ic.: Syreistsch.
Ill. Fl. Mosk. Gub. III (1910) 281; Fedtsch. and Fler. Fl. Evrop. Ross.
995.—Exs.: GRF No. 3495.

Biennial, sometimes annual. Roots adventitious. Whole plant
glandular-floccose-arachnoid-hairy, less often glabrous in lower part
or even all over stem, excluding inflorescence. Stem hollow, simple or
branched above, 20-100 cm high, 0.5—2.0 cm thick, densely leafy.
Leaves oblong, lanceolate or lanceolate-linear, obtuse, usually
pubescent, less often subglabrous, 5-15 cm long and 0.7—5.0 cm wide,
sessile, with semiamplexicaul base, sinuate-coarsely toothed or almost
pinnately lobed; upper leaves smaller, almost entire; some leaves in
rosette narrowed into short petiole, withering before flowering; lower
cauline leaves like basal, but smaller, 2-6 cm long; middle and upper
leaves sessile, expanded at base and 5-15 mm wide, ovate-lanceolate,
long-acuminate, with long, usually entangled hairs along margin,
sparsely appressed hairy above, or subglabrous. Capitula in dense or
lax corymbose inflorescence; peduncles arachnoid-hairy, with long hairs.
Involucre in single whorl, with linear, acuminate, involucral bracts
densely hairy only at base, sparsely appressed-hairy in remaining part,
sometimes almost glabrous above. Ligulate florets light yellow, 1.0—
2.5 mm wide, with 2-3 obtuse teeth or entire. Achenes finely ribbed,
glabrous, 1.5—2.0 mm long. Flowering July to August.

Sandy-clayey banks of lakes, marshes and rivers, marshy tundras.—
Arctic: Arctic Europe, Novaya Zemlya, Arctic Siberia, Chukotka,
Anadyr; European part: Ladoga-IImen, Baltic Region, Upper Volga,
Upper Dnieper, Middle Dnieper, Volga-Kama, Volga-Don, Lower Don,
Trans- Volga, Black Sea Region (rarely); Western Siberia: All regions;
Eastern Siberia: All regions; Far East: Zeya-Bureya, Okhotsk,
Kamchatka, Ussuri; Soviet Central Asia: Aralo-Caspian Region. General
distribution: Central Europe, Atlantic Europe, Scandinavia. Described
from European Arctic. Type in Leningrad.

Note. I could not find any morphological differences between S.
tubicaulis Mansf. (=S. paluster (L.) Hook.), S. congestus (R. Br.) DC.,
and S. arcticus Rupr.

754

719

Section 13. Tephroseris (Rchb.) DC. Prodr. VI (1837) 359
(‘“‘Tephroserides”).—Cinerariae sect. Tephroseris Rchb. Fl. germ.
excurs. (1831) 241.—Leaves undivided, alternate, entire or toothed;
inflorescence corymbose or umbellate, very rarely capitulum solitary
on stem; involucre campanulate in single whorl; involucral bracts
acuminate, flat, green or purple, glabrous to completely pubescent,
usually recurved at achene maturity; disk florets tubular, bisexual;
peripheral florets ligulate, with 3 teeth in upper part, pistillate. Plant
perennial, rarely biennial.

Series 1. Campestres Schischk.—Perennial, usually densely
pubescent plants with ovate or lanceolate leaves on short or more or
less long petioles; leaves entire or with short, inconspicuous teeth,
ligulate florets usually about 12, often orange; achenes pubescent.

56. S. campester (Retz.) DC. Prodr. VI (1837) 361. p. min. p.;
Ldb. FI. Ross. II, 646, ex p.; Turcz. Fl. baic.-dahur. II, 23, p.; Trautv.
in Tr. Peterb. Bot. Sada, VI, 2 (1880) 552; Schmalh. FI. II, 85, ex p.;
Fedtsch. and Fler. Fl. Evrop. Ross. 995.—S. kochii Ldb. Fl. Ross. II
(1845-1846) 643.—S. integrifolius auct. plur. fl. USSR; Kryl. Fl. Zap.
Sib. XI, 2853, non Clairv.—Cineraria alpina Retz. Observ. bot. I
(1779) 30, non L. C. campestris Retz. Prodr. Fl. Scand. ed. 1 (1779)
159.—C. crispa y. schkurii Ldb. Fl. Alt. [V (1833)104.—Tephroseris
campestris Rchb. Fl. saxon. (1842) 147, quo ad nomen.—Ic.: Gmel.
Fl. sib. II, tab. 71.

Perennial. Rhizome ascending, short, with numerous slender roots.
Stem erect, simple, including leaves floccose-arachnoid-hairy or
glabrescent, 15—60 cm high. Basal (often withering early) and lowermost
cauline leaves with somewhat expanded petiole as long as lamina,
longer, or considerably shorter, lamina broadly ovate, ovate, or narrowly
ovate, obtuse, with small and large teeth, 2-5(10) cm long and 1-2 cm
wide; cauline leaves usually sessile, often semiamplexicaul, oblong or
oblong-lanceolate, almost entire or indistinctly toothed, 2-8 cm long
and 0.5-2.0 cm wide; upper leaves lanceolate-linear or linear.
Inflorescence terminal, usually corymbose or umbellate, of 7-20
capitula, on unequal (0.5—4.5 cm long) peduncles with linear bracts,
less often capitula solitary. Involucre S-6 mm long and 8-10 mm in
dia, with linear-lanceolate involucral bracts sometimes persisting up to
fruiting. Ligulate florets yellow or light yellow, less often orange, 10—
12, 6-10 mm long and about 3 mm wide. Achenes narrowly ovoid,
about 2-5 mm long and 0.75 mm wide, white-pubescent. Flowering
July to August.

720

Plate XXXV.
Senecio karjaginii Sof., habit; 2 — S. atropurpureus (Ldb.) B. Fedtsch.,

WEE

1

habit.

757

721

Patchy and clayey tundra, birch groves, juniper thickets, willow
stands, rocky slopes, talus, sandy and gravelly banks, wet meadows.—
Arctic: Arctic Europe, Arctic Siberia (Polar Urals, Yamal Peninsula);
European part: Karelia-Lapland. Dvina-Pechora; Western Siberia: All
regions; Eastern Siberia: Angara-Sayans. General distribution:
Scandinavia, Central Europe. Described from Siberia. Type in
Stockholm.

Note. The plant is highly variable over its vast range in the degree
of pubescence of the stem, leaves and inflorescence, the number of
capitula, the color of the ligulate florets, the size of the whole plant
and also the leaves.

57. S. ezernjaevii Mind. in Ukr. Bot. Zhurn. XIII, 3 (1956) 55.—
S. campestris auct. plur. Fl. Ross. non DC.—Ic.: Mind. op. cit. Fig. 1,
B, p. 56.—Exs.: Wol. Fl. Polon. exs. No. 640.

Perennial. Plant with short rhizome. Stem erect, simple, finely
sulcate, 25-70 cm high, weakly floccose-arachnoid-hairy or
subglabrous. Basal leaves with petioles as long as lamina or twice as
long, lamina ovate, obtuse, sinuate-toothed, crenate, or almost entire,
2-8 cm long and 1.0—3.5 cm wide; middle cauline leaves sessile, oblong-
lanceolate, acuminate; upper leaves linear-lanceolate. Capitula 5-15,
aggregated in umbellate panicle. Involucre in single whorl, slightly
arachnoid-woolly; involucral bracts almost linear, 4.5-5.5 mm long
and 0.7-1.5 mm wide, acuminate. Ligulate florets, 10-15, yellow, 7-
11 mm long and 2-3 mm wide; disk florets numerous, 5.0—-7.5 mm
long. Achenes almost terete, about 3 mm long, ribbed, densely
pubescent; pappus white, 2-3 times as long as achene. Flowering May
to July; fruiting July to August.

Groves, scrubs, pine woods, forest edges, steppes and meadows.—
European part: Ladoga-I|men (? introduced), Dvina-Pechora (extreme
south), Upper Dnieper, Middle Dnieper, Upper Volga, Volga-Kama,
Volga-Don, Trans-Volga, Lower Don, Lower Volga, Bessarabia;
Western Siberia: Upper Tobol. General distribution: Central Europe.
Described from Kharkov Region. Type in Kiev.

58. S. jailicola Juz. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XV (1953) 399.—S. aurantiacus B. tomentosus Ldb. FI. Ross.
II (1844-1846) 647, quo ad pl. taur.—S. campestris Y. capitatus DC.
Prodr. VI (1837) 361, quo ad pl. taur.—S. campestris Boiss. Fl. or.
III, 412 quo ad pl. taur.—Exs:.: Callier, Iter. taur. tert. (1900) No. 639.

Perennial. Rhizome obliquely ascending or almost horizontal, short,
densely covered with slender and long roots. Stem solitary, erect, simple

758

722

or somewhat branched in upper part, ribbed, densely white-tomentose,
when young, later glabrescent, 15-40 cm high. Basal leaves ovate or
oblong, gradually narrowed into petiole as long as lamina or shorter,
somewhat broadened; lamina 3-7 cm long and 1-3 cm wide, obtuse,
with small teeth or almost entire; cauline leaves oblong or lanceolate
or oblong-linear, sessile, subacute. Inflorescence corymbose at apices
of stem and branches (if present). Capitula 3-6, on short, but later
unequal, elongated peduncles. Involucre broadly campanulate, S—7 mm
long and 10 mm in dia; outer involucral bracts lacking, inner bracts
linear-oblong, subacute, with narrow scarious margin, arachnoid-hairy.
Ligulate florets about 10, yellow, 8 mm long and 2—3 mm wide. Achenes
terete, ribbed, pubescent, about 3 mm long. Flowering end of May to
July; fruiting July to August.

Stony slopes, talus, in montane pine forests.—European Part:
Crimea. Endemic. Described from Babugan-Yaila. Type in Leningrad.

59. S. amurensis Schischk. in Addenda, XXV, 885.—S. pratensis
Maxim. Prim. Fl. Amur. (1859) 166.—S. pratensis var. polycephalus
Rgl. Tent. Fl. Ussur. (1861) 93, non S. polycephalus Ldb. (1844—
1846).—S. campestris auct. Fl. Orient. Extrem.: Maxim. Prim. Fl. Amur.
165: Kom. Fl. Manchzh. III, 628; Kom. and Alis. Opred. Rast.
Dalnevost. Kr. II, 1057, non DC.—Ic.: Kom. and Allis. op. cit. p. 315
(sub S. campestris*).

Perennial. Root adventitious; stems solitary or 2—3, erect, densely
white arachnoid-hairy, (10)20—60 cm high. Basal leaves ovate, narrowly
ovate, or sometimes lanceolate, obtuse, or subacute, basally narrowed
into flattened petiole, on both sides arachnoid-hairy, with very small,
blackish, remote teeth, their laminas 1.5-8.0 cm long and 1-5 cm
wide; cauline leaves numerous, gradually reduced upward. Lowermost
cauline leaves identical with basal ones, short-petiolate, rest sessile,
amplexicaul, lanceolate, acute, 4-8 cm long and 1.0-1.5 cm wide;
upper leaves lanceolate linear, smaller. Capitula 3-10, in umbellate
inflorescence on arachnoid-pubescent peduncles usually many times as
long as capitulum, with linear, acute, small, bracteal leaves at base of
umbel. Involucre in single whorl, broadly-campanulate, 6-7 mm long
and about 10 mm in dia. Ligulate florets about 12, light yellow, oblong,
10 mm long and 2.0-2.5 mm wide, ovary and anthers pubescent.
Achenes terete, 2.0-2.5 mm long. Flowering May to July.

Sedge-mixed herb meadows, marshy meadows, mixed and oak
forests and edges, sandy meadows, slopes and occasionally in cultivated
fields\—Far East: Zeya-Bureya, Ussuri. General distribution:
Northeastern China. Described from vicinity of Blagoveshchensk. Type
in Leningrad.

*sampastri here but correctly given below and in index—General Editor.

759

723

60. S. succisifolius Kom. Fl. Kamch. III (1930) 167.

Perennial. Stem erect, simple, sulcate, violet in lower half, sparsely
floccose-hairy, 50-60 cm high. Basal and lower cauline leaves on
floccose-hairy petioles longer than lamina (4—10 cm long), withering
early, their laminas ovate or narrowly ovate, 3.5—9.0 cm long and 2.5—
3.0 cm wide, acute, with fine teeth or entire, cuneately narrowed toward
base, glabrous; middle cauline leaves like basal but narrower; upper
leaves narrowly lanceolate, smaller. Capitula 3, on short, 4 cm-long,
pubescent peduncles in umbellate inflorescence. Involucre in single
whorl, about 1 cm in dia; involucral bracts about 20, linear-lanceolate,
acuminate, 6-7 mm long, covered with multi-cellular brownish hairs.
Ligulate florets pale yellow, about 12, 12 mm long and 3 mm wide.
Achenes pubescent, 4 mm long; pappus white. Flowering July to August.

Riverine meadows.—Far East: Kamchatka. Endemic. Described
from Koryatskaya Creek, a tributary of Avacha River. Type in
Leningrad.

Note. Along with S. succisifolius, V.L. Komarov recognized the
presence of typical S. campestris (Retz.) DC. also in Kamchatka but,
in my opinion, the latter species is not found at all in Kamchatka and
is replaced here by S. succisifolius Kom.

61. S. lenensis Schischk. in Addenda, XXV, 885.

Perennial. Rhizome ascending, 8-10 mm thick, densely covered
with numerous slender roots. Stem glabrous or floccose-hairy, uniformly
leafy in lower half, almost lacking leaves in upper half, hollow, finely
sulcate, 25-50 cm high. Basal leaves with petioles shorter than lamina
or slightly longer, lamina ovate or lanceolate-ovate, 4-6 cm long and
1.5—3.0 cm wide, obtuse or acute, with remote, fine, blackish teeth,
glabrous or floccose-hairy; lower cauline leaves narrower than basal
leaves, usually lanceolate, acuminate or sometimes acute, sessile, very
rarely long-petiolate, 4-12 cm long and 0.5—2.0 cm wide, almost linear
and small-in upper part of stem. Inflorescence umbellate, with 3-7
capitula on peduncles usually of unequal length. Involucre in single
whorl, 5-8 mm long and 10-15 mm in dia, sparsely pubescent;
involucral bracts lanceolate-linear, with thin, acuminate, blackish apex.
Ligulate florets about 12, yellow, sometimes orange yellow, 12 mm
long and 3 mm wide. Achenes 2.0-3.5 mm long, pubescent. Flowering
July to August.

Meadows, floodplains of rivers, tundra.—Eastern Siberia: Lena-
Kolyma. Endemic. Described from Yakutian ASSR, the village of
Amginskoe. Type in Leningrad.

62. S. cladobotrys Ldb. FI. Ross. II (1845-1846) 644; Boiss. FI.
or. III, 411; Grossh. Fl. Kavk. IV, 148.—S. cladobotrys var. typicus

760

724

Cuf. in Fedde, Repert. LXX, 1, III (1933) 93.—Cineraria longifolia
MB. FI. taur.-cauc. I (1808) 315, non Jacq. (1774).—C. campestris €.
dentatus DC. Prodr. VI (1837) 361, ex p.; Hohenack. in Bull. Soc.
Nat. Mosc. VI, 62.—C. campestris var. longifolia Trautv. in Tr. Peterb.
Bot. Sada, V, 2 (1878) 471.—Tephroseris cladobotrys Griseb. and
Schenk in Wiegmann’s Arch. XVIII, 1 (1852) 342.—S. brachychaetus
DC. Prodr. VI (1837) 362. p. min. p.

Perennial. Rhizome short, oblique, with numerous roots. Stem
solitary, simple, erect, sparsely arachnoid-hairy, with readily detaching
hairs or glabrous, sulcate, hollow, 0.5—1.5 cm thick and 60-160 cm
high. Basal leaves withering early lower cauline leaves on expanded
petioles almost as long as lamina, their laminas broadly ovate or ovate,
cordate, subacute, with coarse, irregular, subacute teeth, 6-12 cm long
and 3.5—10.0 cm wide; middle cauline leaves like lower but on shorter
petioles; upper cauline leaves lanceolate or linear-lanceolate, long-
acuminate, basally narrowed into short petiole. Inflorescence corymbose,
dense when young, later lax; capitula numerous (20-60), on usually
long, glabrous or weakly arachnoid-hairy peduncles, sometimes with
narrowly linear, entire, long-acuminate bracteal leaves. Involucre
broadly campanulate, 6-7 mm long and 7-8 mm in dia, arachnoid-
hairy or subglabrous; outer involucral bracts absent, inner linear-oblong,
subacute or obtuse. Ligulate florets yellow, 12—15(20), oblong, 7-12
mm long and 1.5—2 mm wide. Achenes terete, ribbed, about 4 mm
long, pubescent or sometimes subglabrous. Flowering June to July;
fruiting August.

Mountain forests, tall-herb meadows.—Caucasus: Ciscaucasia,
Western Transcaucasia. Endemic. Described from Western Georgia.
Type in Leningrad.

63. S. subfloccosus Schischk. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XV (1953) 402.—S. brachychaetus DC. Prodr. VI (1837)
362, ex p. quo ad pl. cauc.; Boiss. Fl. or. III, 411.—Grossh. Opred.
Rast. Kavk. IV, 148, non DC.—S. cladobotrys var. pseudorivularis
Cuf. in Fedde, Repert. LXX, I-III (1933) 94.—S. campester var.
brachychaetus Schmalh. Fl. II (1897) 85.—S. brachychaetus var.
xerophilus Grossh. Fl. Kavk. IV (1934) 148; var. mesophilus Grossh.
op. cit. 148; var. longifolius Grossh. op. cit. 149.—S. integrifolius
Grossh. op. cit. 149.

Perennial. Rhizome short, with slender fibrous roots. Stem simple,
solitary, including leaves floccose-arachnoid-hairy, sulcate, 40-120 cm
high. Basal leaves numerous or few, with expanded petioles as long as
lamina, shorter, or longer, lamina ovate, obtuse, or sometimes sub-
acute, irregularly finely toothed, with obtuse or acute, broadly deltoid

725

teeth or almost entire, 3.5-10.0 mm long and 2-5 cm wide; lower
cauline leaves identical with basal ones; middle leaves sessile, lanceolate
or narrowly lanceolate, obtuse or acute, 3-10 cm long and 1.5-2.0 cm
wide; upper leaves also sessile, but smaller. Capitula 12—50 (less often
3-10), aggregated into umbellate inflorescence at apex of stem, on
arachnoid-hairy peduncles of unequal length; bracteal leaves 2-10,
linear, 1-2 cm long and about 1 mm wide, floccose-hairy or subglabrous
at base of inflorescence. Involucre about 8 mm long and almost as
much in dia; outer involucral bracts lacking, inner linear, with narrow
scarious margin, dorsally weakly arachnoid-hairy. Ligulate florets 12—
14, yellow or orange, 12-14 mm long and 1-2 mm wide. Achenes
pubescent, terete, finely ribbed, brownish, 3-4 mm long; pappus white,
shorter than tubular florets, about 6 mm long at fruiting.

Forest edges, cut-over mountain forests, subalpine and alpine
meadows up to 2,500 m, rarely ascending higher.—Caucasus:
Ciscaucasia, Dagestan, Eastern, Western (rarely), and Southern
Transcaucasia. Endemic. Described from vicinity of Bakuriani. Type
in Leningrad.

Series 2. Papposi Schischk.—Perennial, subglabrous or weakly
pubescent plants; leaves oblong or lanceolate, more often finely toothed
or almost entire; ligulate florets about 15.

64. S. papposus (Rchb.) Less in Linnaea, VI (1831) 244 p. p.;
DC. Prodr. VI, 360; Cuf. in Fedde, Repert. LXX, I-III, 187.—Cineraria
761 papposa Rchb. Icon. II, Fl. critic. (1824) 13.—Tephroseris papposa
Schur. Enum. plant. Transsilv. (1866) 344.—Ic.: Rchb. op. cit. (1824)
t. 124.—Exs. Degen, Pl. Banatus exs. a. 1897, sine numero.
Perennial. Rhizome short. Stem sulcate, floccose-arachnoid-hairy,
basally subglabrous, scabrous above, 40-130 cm high. Basal leaves
long-persistent, with petioles longer or shorter than lamina, lamina
ovate or narrowly ovate, less often spatulate, 6-12 cm long and 2.0—
3.5 cm wide, with 5 remote teeth, less often coarsely toothed, or both
sides arachnoid-hairy (more densely beneath), subacute, gradually
narrowed toward base. Inflorescence umbellate, at stem apex; capitula
on arachnoid-hairy peduncles of unequal length. Involucre in single
whorl, 8-10 mm long and about 15 mm in dia. Involucral bracts dark
green, less often purple, loosely arachnoid-hairy. Ligulate florets 15,
about 3 cm long and 3 mm wide, yellow. Achenes glabrous, 3.0—3.5
mm long. Flowering June to July.
Mountain meadows.—Upper Dniester (Eastern Carpathians).
General distribution: mountains of Central Europe. Described from
Western Europe. Type in Berlin.

762

726

65. S. kawakamii Makino in Bot. Mag. Tokyo, XX VI (1912) 291;
Cuf. in Fedde, Repert. LXX, I-III (1933) 259.—Ic.: Miyabe and
Miyake, Fl. Saghal., t. 8 (1915).

Perennial. Rhizome short, erect or ascending, with numerous roots,
root collar covered with dark brown petioles of dead leaves. Stem
sulcate, simple, floccose-arachnoid-hairy, later sometimes subglabrous,
20-30 cm high. Basal leaves finely arachnoid-hairy, more or less long-
petiolate, ovate, ovate-oblong or elliptical, sometimes almost round,
mostly obtuse, gradually or abruptly narrowed into petiole, with rather
distinct, remote, acute teeth, 2.5—8.0(11.0) cm long and 2.0—3.0(5.5)
cm wide, petiole usually shorter than lamina, less often much longer;
cauline leaves usually sessile, semiamplexicaul, narrowly ovate, oblong-
lanceolate, or linear-oblong, finely toothed or entire, acute, turning
distinctly black when dry. Capitula 3-8, at apices of stem, in corymbose-
umbellate inflorescence, with linear, bracteal leaves. Involucre in single
whorl, broadly campanulate, 7-8 mm long and 12-15 mm in dia;
involucral bracts 10-15, linear-lanceolate, indistinctly narrowly scarious
along margin, glabrous, dorsally crisped-hairy. Ligulate florets yellow,
12-15, 10-15 mm long and 1.5—3.5 mm wide. Achenes terete, glabrous,
ribbed, about 4 mm long. Flowering June to July.

Stony mountain slopes.—Far East: Sakhalin (southern). General
distribution: Japan (Hokkaido, Hondo). Described from southern
Sakhalin. Type in Tokyo.

66. S. asiaticus Schischk. and Serg. nom. nov.—S. praticola
Schischk. and Serg. in Sistem. Zam. Gerb. Tomsk. Univ. 1-2 (1949);
Kryl. Fl. Zap. Sib. XI, 2855, non S. praticola Rouy, Fl. Franc. VIII
(1903) 316, nom. nud.—S. spathulifolius v. praticola Rouy (1903) 1.
c. descr.—S. pratensis Ldb. Fl. Ross. II, 644, p. min. p.; Turez. Fl.
baic.-dahur. II, 94, non DC.—S. glabellus (Turez.) DC. Prodr. VI (1837)
360, non Poir. (1806).—S. campestris 5. glabratus DC. L. (1837)
381.—S. subdentatus (Bge.) Turcez. var. glabellus (Turez. ex DC.)
Cuf. in Fedde, Repert. LXX, I-III (1933) 83.—Cineraria glabella
Turcz. ex DC. (1837) |. c. in synon.

Perennial. Rhizome short, densely covered with slender roots. Stem
solitary, simple, glabrous or weakly arachnoid-hairy, ribbed, solid, 20—
40 cm high. Basal leaves long-persistent with expanded petioles as
long as lamina or shorter, lamina ovate, 2.0-4.5 cm long and 1—2 cm
wide, obtuse, usually narrowed toward base, with short, wide, acute
teeth, indistinctly arachnoid-hairy or subglabrous; lower cauline leaves
like basal; middle leaves usually lanceolate, acute, sessile,
semiamplexicaul, toothed or almost entire; upper leaves linear,
acuminate or acute. Capitula 2-12, on weakly arachnoid-hairy peduncles

763

T27

of variable length or almost equal. Involucre in single whorl, 5-8 mm
long and 8-10 mm in dia; involucral bracts linear-lanceolate, acuminate,
weakly arachnoid-hairy or subglabrous. Ligulate florets yellow, 12-
20, linear, about 8-12 mm long and 2.5-3.5 mm wide. Achenes
glabrous, 2.0—-2.5 mm long. Flowering June to July.

In steppe vaileys of mountain rivers, cut-over larch forests, alpine
and subalpine meadows.—Western Siberia: Altai; Eastern Siberia:
Angara-Sayans, Dauria. General distribution: Mongolia? Described
from alpine Shebel. Type in Leningrad.

67. S. sukaczevii Schischk. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVI (1954) 437.

Perennial. Rhizome short. Stem erect, simple, glabrous or weakly
arachnoid-floccose, sulcate, hollow, 30-70 cm high. Basal leaves
withering early, lanceolate or narrowly ovate, gradually narrowed into
petiole as long as lamina, longer, or shorter, lamina 3-8 cm long, 0.5-
2.0 cm wide, green above, lighter beneath, subobtuse, entire or with
indistinct teeth; lower cauline leaves like basal leaves; middle and
upper leaves sessile, semiamplexicaul, shorter and narrower, usually
obliquely upright, almost appressed toward stem; uppermost leaves
linear. Inflorescence with linear, sometimes almost filiform bracts at
base; capitula 5—20. Involucre in single whorl, broadly campanulate, 5
mm long, 8-10 mm in dia. Involucral bracts lanceolate-linear,
acuminate, glabrous. Ligulate florets 12-15, light yellow, 5-6 mm long,
1-2 mm wide. Achenes glabrous, ribbed, about 2 mm long; pappus
white. Flowering June to July.

Marshy meadows and gravel beds of rivers —Eastern Siberia: Dauria,
Lena-Kolyma. Endemic. Described from Shilka River. Type in Leningrad.

68. S. aquilonaris Schischk. in Addenda, XXV, 684.

Perennial. Rhizome quite long, 3-8 mm thick, covered with few,
lone, slender roots; stems straight or somewhat flexuous, simple, few
or solitary, usually violet, sulcate, weakly arachnoid-hairy or
subglabrous, 20-30 cm high. Basal leaves with expanded petioles,
shorter or longer than lamina, lamina ovate or sometimes oblong, obtuse
or subacute, along with few, short, broadly deltoid sharp teeth, less
often entire, on both sides glabrous, 2-7 cm long and 1.0—-2.5 cm
wide; cauline leaves 2—4, sessile, amplexicaul, lanceolate, entire,
glabrous, 3-6 cm long and 0.8—-1.2 cm wide, smaller above. Capitula
2-5 at apices of stem, on short, densely arachnoid-hairy peduncles,
sometimes capitula solitary. Involucre 10-12 mm long and 12-15 mm
in dia, in single whorl. Involucral bracts lanceolate-linear, dark purple,
densely grayish-arachnoid-tomentose. Ligulate florets 10-12, yellow,

764

728

1.5 times as long as involucre, 3-4 mm wide, often lacking. Achenes
terete, glabrous, 3.5 mm long; pappus 2-3 times as long as achene.
Flowering August.

Riverbanks, scrubs.—Arctic: Arctic Siberia; Far East: Lena-Kolyma.
Endemic. Described from Medvezhei River valley. Type in Leningrad.

69. S. igoschinae Schischk. in Addenda, XXV, 885.

Perennial. Rhizome very long, horizontal. Stem glabrous, 30—40
cm high. Basal leaves 2-3; lower cauline leaves long-petiolate, petiole
2-3 times as long as lamina, lamina narrowly ovate, 2-5 cm long and
0.7-1.5 cm wide, apically round, cuneately narrowed toward base,
glabrous, with small teeth or entire; middle cauline leaves lanceolate,
subacute, gradually narrowed into petiole as long as lamina; upper
cauline leaves linear, entire. Capitula 5, at apices of stem in umbellate
inflorescence. Involucre 5 mm long and about 10 mm in dia, in single
whorl; involucral bracts linear, almost glabrous or indistinctly
pubescent. Ligulate florets about 15, egg-yellow, twice as long as
involucre. Achenes terete, about 4 mm long, glabrous or pubescent.
Flowering July.

Mountain tundra, near melting snows, in grassy pine forests at
900-1,500 m.—European part: Volga-Kama (Urals). Endemic. De-
scribed from Southern Urals, from Belaya River. Type in Leningrad.

Series 3. Alpestres Schischk.—Perennial, subglabrous plants;
leaves ovate, sometimes ovate-cordate, with more or less coarse teeth,
long-petiolate. Ligulate florets 12-20. Achenes glabrous.

70. S. sichotensis Kom. Bot. Mat. Gerb. Glav. Bot. Sada, VI
(1926) 15; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1654.

Perennial. Rhizome obliquely ascending, 3 mm thick. Stem solitary,
sulcate, glabrous or weakly arachnoid-hairy, simple, 20-50 cm high.
Basal leaves numerous, with petioles shorter or almost as long as lamina,
lamina ovate, subobtuse, irregularly coarsely toothed, dark green above,
lighter beneath, weakly floccose-arachnoid-hairy, 3.0—6.5 cm long and
1.5-3.0 cm wide; lower cauline leaves also petiolate; middle leaves
sessile, irregularly toothed, ovate-lanceolate or lanceolate, acute, 3—10
cm long and 0.8-3.0 cm wide; upper leaves lanceolate-linear, with a
few teeth. Inflorescence umbellate; peduncles weakly pubescent, 3-6
cm at base, with linear bracteal leaves. Involucre in single whorl, 4—
5 mm long, about 5 mm in dia in upper part. Involucral bracts 12-15,
dark violet, pubescent attenuate into almost blackish tip. Ligulate florets
10-15, yellow, 10 mm long and 1.5 mm wide. Achenes glabrous, ribbed,
about 3 mm long; pappus white. Flowering August.

765

729

Herb meadows at 1,800 m.—Far East: Ussuri. Endemic. Described
from Botcha River valley. Type in Leningrad.

71. S. veresczaginii Schischk. and Serg. in Sistem. Zam. Gerb.
Tomsk. Univ. I (1944).—Kryl. Fl. Zap. Sib. XI, 2856.—S. crispa
dilatata Ldb. Fl. Alt. IV (1833) 103.—S. alpestris Ldb. Fl. Ross. II,
645 ex parte non DC.

Perennial. Rhizome ascending; stem simple, erect, finely sulcate, basally
sometimes reddish-violet, glabrous or with leaves weakly arachnoid-floccose,
20-40 cm high. Basal and lower cauline leaves with petiole almost as long
as lamina, lamina ovate or roundish, coarsely toothed, 3-5 cm long and
2.54.0 cm wide; middle cauline leaves oblong or oblong-ovate, basally
narrowed into short, expanded, semiamplexicaul petiole, 3-6(9) cm long
and 1.5-—2.5 cm wide, toothed; upper leaves sessile, lanceolate or linear,
toothed or entire. Inflorescence corymbose; capitula 2-7 (very rarely
solitary), on peduncles with lanceolate-linear, acuminate, small, bracteal
leaves at base. Involucre 6 mm long and 7-15 mm in dia, glabrous or
somewhat arachnoid-hairy, in a single whorl. Ligulate florets yellow, 12—
20. Achenes glabrous; pappus a half or slightly more as long as tubular
florets. Flowering July to August.

Rocks and herb slopes in subalpine zone.—Western Siberia: Altai.
Endemic. Described from Altai. Type in Tomsk.

72. S. rivularis (Waldst. and Kit.) DC. Prodr. VI (1837) 359;
Szafer, Kulcz. Pawl. Rosliny Polskie (1953) 700.—S. rivularis (Waldst.
and Kit.) DC. var. typicus Domin and Podpera, Klic k uplné kvetene
rep. Ceskoslovenske, 2 (1928) 651.—S. crispus Kittel. Taschenbuch
Fl. Deutsch. Ed. 2, II (1844) 588, non Thunb. (1794).—S. crispatus
DC. Prodr. VI (1837) 359; Fedtsch. and Fler. Fl. Evrop. Ross. 995.—
S. kochii Ldb. Fl. Ross. II (1844-1846) 643, p. min. p.—Cineraria
crispa Jacq. Fl. Austr. Icones, II (1774) 48.—C. rivularis Waldst. and
Kit. Ic. pl. rar. Hung. III (1812) 265.—Thephroseris crispa Rchb. FI.
Saxon. (1842) 142.—Ic.: Waldst. and Kit. op. cit. tab. 239; Hegi, III.
Fl. VI, 445.—Exs.: Fl. exs. Reipubl. Bohem.-Sloven. No. 379, 380,
955; Fl. Stir. exs. No. 586; Fl. Siles. exs. No. 837, 838, 839; FI. exs.
Bavar. No. 538; Hayek, Fl. Styr. exs. No. 586.

Perennial. Stem erect, branched above, 40-80 cm high, arachnoid-
hairy when young, glabrous when old. Basal leaves ovate-cordate,
obtuse, with sharp teeth, on long thickened petiole sometimes 2-3
times as long as lamina, lamina subglabrous and dark green above,
paler and pubescent beneath, 3-7 cm long and 2-6 cm wide; lower
cauline leaves like basal but with shorter petiole; upper leaves sessile,
lanceolate or narrowly ovate. Inflorescence umbellate, with 4-15,

766

730

somewhat irregular, slightly pubescent rays, with many linear-lanceolate
bracteal leaves at base. Involucre about 10 mm long and almost much
in dia, in single whorl, weakly pubescent. Ligulate florets linear-oblong,
about 20. Achenes glabrous, about 2 mm long. Flowering July to August.

Wet subalpine meadows, turfy meadows, scrubs and stony slopes
at 600-1,600 m.—European part: Upper Dniester. General distribu-
tion: Balkans. Described from Hungary. Type in Budapest.

Note. S. subalpinus Koch is very close to S. rivularis (Waldst.
and Kit.) DC. in habit, but is easily distinguished by the absence of the
outer involucral bracts.

Series 4. Hieraciiformes Schischk.—Plants perennial with ovate
or lanceolate leaves on petioles almost as long as lamina and with
small remote teeth; achenes glabrous; ligulate florets 10-13, yellow or
egg-yellow. .

73. S. hieraciiformis Kom. in Bot. Mat. Gerb. Gl. Bot. Sada, VI
(1926) 16 (S. hieraciformis); Kom. and Alis. Opred. Rast. Dalnevost.
Kr. II, 1054.

Perennial. Rhizome 3 mm thick, obliquely ascending. Stem simple,
glabrous or weakly arachnoid-hairy, 10-25 cm high. Cauline leaves
crowded in lower half of stem with petiole usually shorter than lamina,
very rarely as long; lamina ovate or lanceolate, less often broadly
ovate, cuneately narrowed at base, with small remote teeth, 2-7 cm
long and 0.8—2.5 cm wide and with short curly hairs (mainly along
margin). Capitula 2 or solitary. Involucre 4 mm long and 6 mm in dia;
involucral bracts 12, oblong, long-acuminate with recurved blackish
apex, glabrous. Ligulate florets yellow, 10-13, 10-18 mm long and 2—
4 mm wide. Achenes glabrous, terete, 4 mm long and 1 mm wide,
longitudinally ribbed; pappus snow-white. Flowering July.

Riparian rocks and stony slopes.—Far East: Ussuri. Endemic.
Described from Bocha River valley (Sikhote-Alin). Type in Leningrad.

74. S. subscaposus Kom. in Bot. Mat. Gerb. Gl. Bot. Sada, VI
(1926) 16.—Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1057.

Perennial. Stem almost leafless, 15-30 cm high. Basal and lower
cauline leaves elliptical, ovate, or oblong-ovate, broadly cuneate at
base, with petiole as long as lamina or longer; their laminas 2.5—7.0
cm long and 1.5—4.0 cm wide. Capitula 1-8. Involucre about | cm in
dia; involucral bracts 20-30, linear, acuminate, not densely glandular-
hairy, only sometimes almost villous. Ligulate florets about 10, 1.5—
1.8 cm long and 2-4 mm wide. Achenes glabrous, 4.5 mm long; pappus
white. Flowering July. |

767

731

Crevices of rocks and in stony herb meadows.—Far East: Ussuri.
Described from Suchan River from Chernyi Kust Mountain, and from
Yalu-Dzyan River. General distribution:*

Series 5. Porphyranthi Schischk.—Ligulate florets violet-purple
or flame-red, 8-15, capitula many, very rarely solitary; achenes most
often glabrous; perennials.

75. S. porphyranthus Schischk. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVI (1954) 432.—S. integrifolius (L.) Clairv. subsp.
capitatus (Nym.) Cuf. var. aurantiacus Cuf. f. gmelini Cuf. in Fedde,
Repert. LXX, I-III (1933) 26, quo ad pl. sibir. orient.—S. aurantiacus
a. glabratus Turcz. Fl. baic.-dahur. II (1856) 97.—S. aurantiacus 6.
flosculosus Turcz. ibid. (1856).

Perennial. Rhizome short, covered with long, slender roots. Stem
erect, simple, sulcate, with leaves sparsely arachnoid-hairy, sometimes
subglabrous, 25-80 cm wide. Basal leaves long-persistent, abruptly or
gradually narrowed into short petiole or petiole as long as lamina;
lamina broadly or narrowly ovate, sometimes almost obovate, 2.5-14.0
cm long and 1-6 cm wide, obtuse or round, entire or with few, small,
remote teeth; lower cauline leaves like basal, middle leaves sessile,
semiamplexicaul, narrower; upper leaves lanceolate-linear, acuminate.
Capitula 3-9, on sparsely pubescent peduncles of variable length
forming corymbose inflorescence. Involucre 4-5 mm long and 7-8 mm
in dia; outer involucral bracts absent; inner bracts lanceolate-linear,
usually violet, sparsely pubescent. Ligulate florets purple, broadly linear,
8-10 mm long and 3 mm in dia, 8-15. Achenés terete, finely ribbed,
glabrous or with occasional hairs, 2.0-2.5 mm long; pappus silvery-
white, shorter than tubular florets. Flowering June to July.

Birch, pine-birch, pine-larch, and spruce forests, also their edges,
scrubs, often in peat-marsh meadows.—Eastern Siberia: Angara-Sayans,
Yenisei, Lena-Kolyma, Dauria. Endemic. Described from Krasnoyarsk
Territory (Ermakovsk District). Type in Leningrad.

76. S. flammeus Turcz. ex DC. Prodr. VI (1837) 362; Ldb. FI.
Ross. II, 647; Turcz. Fl. baic.-dahur. II, 25.—S. flammeus var. typicus
Cuf. in Fedde, Repert. LXX, I-III (1933) 89.—S. flammeus var.
glabrifolius Cuf. Ibid. (1933) 90.—S. flammeus f. glabrescens Hara in
Journ. Jap. bot. K (1934) 437.—S. flammeus subsp. glabrifolius

*In the original text on p. 766, the end of the description of Series 5 (“very rarely
solitary; achenes most often glabrous; perennials.”) is repeated here in place of the
distribution, which is missing.—General Editor.

768

732

Kitamura in Acta Phyt. et Geobot. VI (1937) 169.—Cineraria flammea
Turcz. ex DC. loc. cit. (1837); Herd. in Bull. Soc. Nat. Mosc. XLI,
415.—C. ignea Fisch. ex Herd. ibid. (1867).—Senecio longeligulatus
Lev. and Van. ex Lev. in Fedde, Repert. VII (1910) 139.—Ic.: Gartenfl.
XII, tab. 394 (1863); Trans. Russ. Hort. Soc. tab. 120 (1863).—Exs:.:
GRF No. 3281.

Perennial. Rhizome short, densely covered with slender roots; stem
simple or somewhat branched in upper part, erect, sulcate, together
with leaves and peduncles floccose-arachnoid-hairy, with readily
detaching hairs, 26-70 cm high. Basal leaves early-withering; lower
cauline leaves with petiole almost as long as lamina and somewhat
expanded lamina ovate-lanceolate or lanceolate, acute or obtuse, on
both sides arachnoid-floccose, more densely beneath, dark green above,
paler beneath, with widely spaced fine or coarse teeth; middle leaves
lanceolate or narrowly lanceolate, sessile, acute, 5-12 cm long and
1.5-3.0 cm wide; uppermost leaves lanceolate-linear or linear,
acuminate, shorter. Capitula 3-20, at apices of stem and upright
branches, aggregated in corymbose inflorescence. Involucre 5-6 mm
long and 8-10 mm in dia; outer involucral bracts absent, inner linear,
acuminate, glabrous or weakly arachnoid-hairy, dark violet. Ligulate
florets 8-15, linear, 15-20 mm long and 1-2 mm wide, flame-red or
dark violet. Pappus brownish, distinctly shorter than tubular florets;
achenes pubescent, somewhat ribbed, about 2 mm long. Flowering June
to July.

Wet and mixed herb meadows, scrubs.—Eastern Siberia: Dauria,
Lena-Kolyma; Far East: Zeya-Bureya, Ussuri. General distribution:
Northeastern China, Korea. ?Japan, Mongolia. Described from vicinity
of Nerchinsk. Type in Geneva; cotype in Leningrad.

77.8. besserianus Mind. in Ukr. Bot. Zhurn. XIII, 3 (1956) 58.—
S. aurantiacus f. flosculosus Bess. ex DC. Prodr. VI (1837) 362.—S.
campestris B. aurantiacus and f. capitatus Schmalh. Fl. Sr. i Yuzhn.
Ross. II (1897) 85.—S. besserianus f. radiatus Mind. op. cit. (1956)
60.—Ic.: Mind. loc. cit. Fig. 2.

Perennial. Rhizome short; stem simple, erect, sulcate, often with
violet striations, weakly arachnoid-hairy or subglabrous, 40-70 cm
high. Basal and lower cauline leaves on 1.5—6.0 cm long petioles, their
laminas elliptical or oblong-elliptical, 3-5 cm long and 1-3 cm wide,
subobtuse, irregularly toothed or sometimes entire, almost glabrous;
middle cauline leaves short-petiolate or sessile, lanceolate or oblong,
smaller than basal; upper leaves sessile, almost linear. Capitula 3-10,
aggregated in umbellate inflorescence. Involucre 6-7 mm long and
10-12 mm in dia, in single whorl, basally arachnoid-hairy; involucral

769

733

bracts oblong-linear, 6-8 mm long and 1-2 mm wide. Ligulate florets
deep violet. Achenes densely pubescent, 2.5-3.0 mm long; pappus
white, 3 times as long as achene. Flowering June to July; fruiting
August.

Oak forests and edges.—European part: Upper Dniester. Endemic.
Described from Pochaev, Ternopol District. Type in Kiev.

78. S. capitatus (Wahlb.) Steud. Nomencl. bot. II (1841) 559.—
S. aurantiacus (Hoppe) Less. in Linnaea, IX (1834) 191, non Rchb.—
S. aurantiacus Y. tomentosus and . capitatus DC. Prodr. VI (1837)
362.—S. aurantiacus b. lanatus Rchb. Ic. Fl. Germ. et Helv. XVI
(1854) 44.—S. integrifolius (L.) Clairv. subsp. capitatus Cuf. in Fedde,
Repert. LXX, I-III (1933) 10.—Cineraria capitata Wahlb. Flora
Carpath. (1814) 271.—C. alpina All. Flora pedemont. I (1785) T.
XXXVIII, f. 2, non L.—C. aurantiaca Hoppe ex Willd. Sp. pl. III
(1804) 2081.—Tephroseris aurantiaca Griseb. and Schenk. Iter Hung.
(1852).—Exs.: Fl. exs. Austro-Hung. No. 3773; Hayek, Fl. Stir. exs.
No. 149.

Perennial. Rhizome short. Stem 25-40 cm high, erect, simple,
densely white-tomentose. Basal leaves in rosette, ovate or lanceolate,
subobtuse, narrowed toward base into very short, flattened petiole or
sessile, 2.5—-7.0 cm long and 2-3 cm wide, entire or with fine, sometimes
very fine teeth, arachnoid-tomentose beneath, subglabrous above;
cauline leaves like basal leaves but narrower, gradually becoming
narrower in upper part, sessile, 2.0-8.5 cm long and 0.5—2.0 cm wide.
Capitula 3-6, on short peduncles in lax heads. Involucre 10 mm long
and 15 mm in dia; involucral bracts in single whorl, lanceolate, acute,
dark violet. Ligulate florets 10-15, flame-red to dark violet, less often
absent. Flowering July to August.

Alpine meadows at 2,100-2,250 m.—European part: Upper
Dniester (Carpathians). General distribution: Tatry. Described from
Carpathians. Type in Sweden.

Note. The ligulate florets are often absent, which is observed es-
pecially often in plants from the Carpathians. However, according to
Woloszczak’s data, occasionally a form with ligulate florets is found
here. It is impossible to treat the form without ligulate florets as a
separate species because the occurrence of ligule-less florets is a
common feature of a majority of the species in section Tephroseri.

79. S. caucasigenus Schischk. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVI (1954) 427.—S. aurantiacus B. leiocarpus Boiss. FI.
or. III (1875) 412.—S. campester var. pyroglossa Trautv. in Tr. Bot.
Sada, V (1877) 445.—S. aurantiacus Grossh. Opred. Rast. Kavk. (1949)
467, non DC.—S. pyroglossus Grossh. ibid. non Kar. and Kir.

770

734

Perennial. Rhizome ascending, densely covered with numerous
slender roots. Stem erect, simple, basally covered with fibrous dark
brown remnants of dead leaves, densely arachnoid-hairy, finely sulcate,
10-45 cm high. Basal leaves ovate, gradually or abruptly narrowed
into expanded petiole shorter or longer than lamina, obtuse or subacute,
along with irregular, broadly deltoid, subacute or almost obtuse teeth,
usually arachnoid-hairy, 2-7 cm long and 1.5-3.5 cm wide; lower
cauline leaves like basal leaves but narrower; middle cauline leaves
lanceolate, acute, sessile; upper leaves lanceolate-linear or linear,
smaller, acuminate or acute; all cauline leaves more or less appressed
toward stem. Capitula at stem apex, 3-7 (9), on peduncles of unequal
length, forming corymbose inflorescence, less often capitula solitary.
Involucre 6-7 mm long and 8-10 mm in dia; outer involucral bracts
absent, inner bracts linear, acuminate, dark violet. Ligulate florets 10—
15, 10-20 mm long, violet or flame-red. Achenes usually pubescent,
less often glabrous, narrowly oblong, 3.5 mm long. Flowering June to
August.

Alpine and subalpine meadows, mountain forests, rocks,
occasionally on talus, at 1,300-3,500 m.—Caucasus: Ciscaucasia,
Dagestan, Eastern, Western, and Southern Transcaucasia. Endemic.
Described from Tskhra-Tskhro Range. Type in Leningrad.

Note. S. caucasigenus Schischk. is widely distributed in the Glavnyi
[Main] Caucasian Range in the alpine and subalpine zones
and in the mountains of Transcaucasia. In habit, it is similar to S.
pyroglossus Kar. and Kir., which so far is known only from Dzhungarian
Alatau.

Possibly, the nonligulate form, described by Steven from the
Glavnyi [Main] Caucasus Range under the name Cineraria fulva Stev.
loc. cit., belongs to this species, and then, maybe, the plant should be
called S. fulvus (Stev.) Schischk. and not S. caucasigenus Schischk.
However, I am not fully convinced of the identity of these [two] spe-
cies and therefore am retaining the latter name as the name assigned
to the specimens with ligulate florets.

80. S. pyroglossus Kar. and Kir. in Bull. Soc. Nat. Mosc. XV
(1842) 386; Ldb. Fl. Ross. II, 647.—S. campestris var. pyroglossa
Trautv. in Tr. Peterb. Bot. Sada, V, 2 (1878) 415.—S. integrifolius
subsp. capitatus var. pyroglossus Cug. in Fedde, Repert. LXX, I-III
(1933) 22.—Exs.: Kar. and Kir. No. 1624.

Perennial. Root vertical or ascending, 3-4 mm thick. Stem erect,
simple, 15-30 cm high, floccose-arachnoid-hairy. Lower leaves
narrowly ovate, narrowed into petiole almost as long as lamina or
longer; middle and upper leaves sessile, semiamplexicaul, 4.5-6:0 cm

—

735

long and 0.8-1.2 cm wide; all leaves acute, with fine, sharp irregular
teeth, floccose-arachnoid-hairy. Capitula at stem apex, 2-5 or (less
often) solitary. Involucre 10 mm long and 10-15 mm in dia, floccose.
Ligulate florets 12-20, 2.0-2.5 cm long and 2-3 cm wide, flame-red,
twice as long as involucre. Achenes glabrous, terete, ribbed, about 3.5
mm long. Flowering July to August.

Alpine zone, alpine meadows, moraines and mountain tundra, at
2,200-3,000 m.—Soviet Central Asia: Dzhungaria-Tarbagatai
(Dzhungarian Alatau). Endemic? Described from the upper reaches of
Lepsa and Sarkan rivers. Type in Leningrad.

81. S. karjaginii Sof. in Izv. Akad. Nauk Azerb. SSR, (1957)
89.—S. primulifolius Somm. and Lev. in Nuov. Giron. Bot. Ital. ser.
2, II (1895/89); Tr. Peterb. Bot. Sada, XVI, 245; Grossh. Fl. Kavk.
IV, 148 non Vell. (1827) nec. Miill. in Hook. (1857).—S. pyroglossus
var. macrocephala Lipsky in Tr. Peterb. Bot. Sada, XIII (1894) 305,
cfr. Ibid. XIV (1898) 294.—S. integrifolius subsp. capitatus var.
pyroglossus f. primulifolius Cuf. in Fedde, Repert. LXX, I-III (1933)
23.—Ic.: Somm. and Lev. in Tr. Peterb. Bot. Sada, XVI (1900) Plate 23.

Whole plant more or less densely floccose-arachnoid-hairy.
Rhizome horizontal, somewhat thick, densely covered with numerous
roots. Stem somewhat thick, weakly sulcate, leafy, 5-20 cm high. Basal
leaves broadly ovate, narrowed into short or long-winged petiole, their
laminas at base cuneate or almost round, sometimes cordate, in young
leaves floccose-arachnoid-hairy along margin, later subglabrous, more
or less coarsely toothed with obtuse teeth, 3.5—4.0 cm long and about
2.5 cm wide; lower cauline leaves like basal leaves but narrower, oblong-
linear, with petiole expanded at base; upper leaves sessile, with
semiamplexicaul auricles; uppermost leaves linear. Capitula 1-8, at
stem apex, in corymbose inflorescence on thick and arachnoid-hairy
peduncles, longer in outer capitula, usually with 1 linear bract. Involucre
about 10 mm long and 15 mm in dia, dark purple; outer involucral
bracts absent, inner bracts linear-lanceolate, dorsally pubescent, apically
glabrous. Ligulate florets 20-25, 9-12 mm long and about 2 mm wide,
often orange; tubular florets 13 mm long. Achenes glabrous, 5 mm
long and 0.75 mm wide; pappus white, slightly shorter or two-thirds as
long as tubular florets. Flowering July to August. (Plate XXV, Fig. 1).

Alpine zone, on talus, moraines, in alpine meadows at 2,300-—
3,500 m.—Caucasus: Ciscaucasia (Glavnyi[Main] Range), Dagestan,
Eastern Transcaucasia (Southern Osetia). Endemic. Described from
Nakhar, Teberdinsk, Kukurtia and Elbrus ranges. Type in Florence;
cotype in Leningrad.

vie

736

Series 6. Frigidi Schischk.—Plants perennial, arachnoid-hairy,
sometimes densely tomentose particularly on peduncles below
capitulum; ligulate florets yellow or sometimes orange, about 15;
capitula often solitary (less often) 3; plants usually 10-20 cm high,
rarely up to 30 cm.

82. S. atropurpureus (Ldb.) B. Fedtsch. in Fedtsch. and Fler.
Evrop. Ross. (1911) 992.—S. frigidus Less. in Linnaea, VI (1831)
289; Ldb. Fl. Ross. II, 632, ex p.; Rupr. in Beitr. Pfl. Russ. VII, 28.—
S. frigidus Less. var. radiata and var. discoidea Trautv. in Tr. Peterb.
Bot. Sada V (1878) 538.—S. frigidus Less. var. radiata Trautv. FI.
ripar. Kolym. (1878); Tr. Peterb. Bot. Sada, VI, 25.—S. integrifolius
(L.) Clairv. subsp. atropurpureus (Ldb.) Cuf. in Fedde, Repert. LXX,
I-III (1943) 43.—S. uralensis Gorodk. ex Kryl. Fl. Zap. Sib. XI (1949)
2342.—Cineraria atropurpurea Ldb. in Mém. Acad. Sc. Pétersb. V
(1815) 574.—C. frigida Richards, Bot. Append. to Frankl. narr. of
Journ. (1823) 748.—C. frigida a. genuina, a. arctica, b. taimyrensis,
c. uralensis Herd. in Bull. Soc. Nat. Mosc. XL (1867) 439.

Perennial. Rhizome short, horizontal or ascending. Stems solitary
or few, erect, 2-10(20) cm high, lanate from base or only in upper
part, with dark purple arachnoid hairs, finely sulcate, simple, very
rarely with 1-2 short branches. Basal and lower cauline leaves ovate,
oblong-ovate or lanceolate, 1-6 cm long and 0.5—2.0 cm wide, sessile
or petiolate, petioles almost as long as lamina or slightly longer,
subobtuse, narrowed toward base, entire or with remote teeth, arachnoid-
hairy or subglabrous; upper leaves lanceolate, sessile, smaller. Capitula
usually solitary at stem apex, less often 2—3. Involucre 7-8 mm long
and about 10 mm in dia, in a single whorl; involucral bracts lanceolate-
linear, acuminate, with dark brown, thin, entangled, multicellular hairs,
with narrow scarious margin. Ligulate florets yellow, 12—15, oblong,
about 10(15) mm long and 4(6) mm wide. Achenes terete, glabrous or
slightly pubescent, about 3 mm long. Flowering July to August. (Plate
XXXV, Fig. 2).

Tundra: mossy, patchy, polygonal, cotton-grass-lichen, sedge-
lichen; in thickets of low shrubs.—Arctic: Arctic Europe, Novaya
Zemlya, Arctic Siberia, Chukotka, Anadyr; Eastern Siberia: Lena-
Kolyma; Far East: Okhotsk (mountains), Kamchatka (mountains).
General distribution: Bering Strait area, Arctic America. Described
from Siberia. Type in Leningrad.

Note. Over its wide range, the plant varies in all respects.
Sometimes it is very small, 2-4 cm high, sometimes it reaches 20 and
even 30 cm (seldom) high; pubescence also varies, which apparently
depends partly on age: young plants, just beginning to flower, are

TS

Plate XXXVI.
1 — Senecio krascheninnikovii Schischk., habit, achene; 2— S. dubius Ldb.,
habit, achene; 3 — S. sosnovskii Sof., habit.

737

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2

775

738

usually strongly pubescent, sometimes almost tomentose (var.
tomentosus Trautv.); later they become glabrescent (var. glabratus
Trautv.). The color of the ligulate florets is sometimes light yellow,
sometimes dark yellow or even orange (var. aurantiacus Trautv.). The
ligulate florets sometimes do not develop (var. flosculosus Trautv. or
var. discoideus Trautv.), but are usually well developed (var. radiatus
Trautv.). The capitula usually are solitary (var. monocephalus Trautv.)
or two or three. The florets mostly are light yellow, but sometimes
reddish-orange (var. pyroglossus Trautv.).

83. S. tichomirovii Schischk. sp. nov. in Addenda, XXV, 886.

Perennial. Rhizome short, ascending. Stem solitary, erect, thick,
4-6 mm thick, 10-15 cm high, glabrous from base to middle, tomentose-
arachnoid-hairy above, with brownish hairs. Basal leaves 4—6, with
petioles 2-3 times as long as lamina, lamina narrowly ovate, 3-4 cm
long and 0.5-1.5 cm wide, glabrous, light green, subacute, toothed,
teeth remote and with callous thickening; cauline leaves like basal but
sessile and amplexicaul, glabrous; upper leaves smaller and with stem
densely arachnoid-hairy. Capitula at stem apex, solitary. Involucre 1.5
mm long and 2.5 mm in dia; involucral bracts in single whorl, as also
part of stem densely lanate tomentose with dark brownish hairs. Ligulate
florets 20-25, dark yellow, linear, 1.5—2 times as long as involucre.
Achenes not known. Flowering July.

Tundra.—Arctic: Chukotka. Endemic. Described from Chukotka.
Type in Leningrad.

84. S. pricei Simps. in Journ. Linn. Soc. XLI (1912-1913) 425.—
S. cryophilus Schischk. and Serg. in Sistem. Zam. Gerb. Tomsk. Univ.
1-2 (1949); Krylov, Fl. Zap. Sib. XI, 2841.—S. integrifolius (L.) Clairv.
subsp. atripurpureus (Ldb.) Cuf. var. robustus Cuf. in Fedde, Repert,
LII, I-III (1933) 43.—Ic.: Simps. op. cit. Pl. 22, fig. 8-12.

Perennial. Rhizome creeping, 2-5 mm thick, with long fibrous
root. Stems solitary or few, erect or basally ascending, 6-25 cm high, with
leaves arachnoid-hairy or floccose-arachnoid-hairy, glabrous below, lanate-
tomentose below capitulum; lower leaves on long expanded petioles
(up to 12 mm wide), 2—4 times as long as petiole, lamina ovate or
almost round, obtuse, at base sometimes almost cordate, with indistinct
obtuse teeth, 1.5-4.0 cm long and almost as wide, glabrous beneath,
floccose-tomentose above; middle cauline leaves like basai but smaller
and short-petiolate; upper leaves sessile, ovate-lanceolate or lanceolate,
acuminate, often arachnoid-woolly. Capitulum solitary, large, including
ligulate florets 4-5 cm wide. Involucre arachnoid-woolly; involucral
bracts in single whorl, linear, acuminate, sordid violet. Ligulate florets

7716

oo

20-25, yellow, linear-oblong, 12-20 mm long and 3.5—4.5 mm wide.
Achenes glabrous, 2.0-2.5 mm long. Flowering July.

Alpine zone, on rubbly tundra, rocks and stony slopes.—Western
Siberia: Altai; Eastern Siberia: Angara-Sayans. General distribution:
Mongolia. Described from Tannu-Ola Range. Type in London.

85. S. turczaninovii DC. Prodr. VI (1837) 360.—Cineraria frigida
Richards. 8. robusta Herd. a. atropurpurea Herd. ex p. a eriocephala
Herd. ex p. in Bull. Soc. Nat. Mosc. XL, 2 (1867) 439.—Senecio
integrifolius (L.) Clairv. ssp. atropurpureus (Ldb.) Cuf. var. taimyrensis
Cuf. in Fedde, Repert. LXX, I-III (1933) 45.—S. integrifolius (L.)
Clairv. ssp. atropurpureus (Ldb.) Cuf. var. robustus (Herd.) Cuf. op.
cit. (1933) 44.—S. tundricola Tolm. in Comt. Rend. Acad. Sci. URSS
(1928) 226, p. min. p.—S. aurantiacus y. tomentosus Turcz. FI. baic.-
dahur. II (1856) 97.—S. aurantiacus 6. capitatus Turcz. Ibid. (1856).—
S. frigidus Turcz. Ibid. (1856) 94, non Less.—Ic.: Popov, Fl. Sr. Sib.
II, Fig. 88.

Perennial. Rhizome short, densely covered with numerous roots.
Stems few, erect, simple, cylindrical, somewhat sulcate: young stems
woolly, old glabrous below and glandular-woolly above, mixed with
white dark purple hairs, 10-20 cm high. Leaves on both sides
subglabrous; basal leaves oblong-ovate, narrowed into petiole shorter
than or as long as lamina, entire or toothed; lower cauline leaves oblong-
lanceolate or lanceolate, semiamplexicaul, toothed or entire; uppermost
leaves sessile, with cordate and amplexicaul base, lanceolate, acuminate,
somewhat sinuate. Capitula at stem apex, 2-3, less often solitary.
Involucre 10-12 mm long and 15 mm in dia. Tubular florets intensely
egg-yellow, anthers dark purple. Ligulate florets about 15, orange-
yellow, with dark purple striations, more intensely colored on lower
side, sometimes ligulate florets absent (f. pseudofulvus Cuf.). Achenes
glabrous or with occasional hairs, about 4 mm long; pappus white.
Flowering July to August.

Marshy places near treeline, alpine and subalpine meadows, in
mountain sedge-hypnum moss tundra up to 3,000 m.—Eastern Siberia:
Angara-Sayans, Dauria. General distribution: Mongolia (mountains in
vicinity of Lake Kosogol). Described from Shebel Range. Type in
Geneva; cotype in Leningrad.

Note. The plants vary greatly in the color of the ligulate florets—
from yellow to orange: in the number of the capitula—1-—S, in the
degree of pubescence of the plant, and in the height of the stem. The
achenes sometimes are glabrous and at times pubescent.

As in the majority of the species of section Tephroseris, plants of
this species are found with capitula lacking ligulate florets (f.
pseudofulvus Cuf.).

Ua

740

86. S. tundricola Tolm. in Compt. Rend. Acad. Sci. URSS (1928)
266 p. p.; Hultén in Svensk. Vet. Akad. Handl. Ser. 3, VIII, No. 2,
205.—S. campester var. aurantiaca Trautv. in Tr. Peterb. Bot. Sada,
V, 2 (1878) 538.—S. campester var. fulva Trautv. in Tr. Peterb. Bot.
Sada, V, 2 (1878) 539, non Cineraria fulva Stev.—S. kamtschaticus
Kom. Fl. Kamch. III (1930) 166.—S. integrifolius ssp. atropurpureus
var. robustus (Herd.) Cuf. Repert. spec. nov. LXX, I-III (1933) 44.

Perennial. Rhizome short, apically covered with blackish-brown
remnants of dead leaves. Stem erect, simple, together with leaves densely
arachnoid-hairy, 12-40 cm high. Basal leaves 2-4, ovate or oblong,
their laminas 1-4 cm long and 1.8—2.0 cm wide, abruptly or gradually
narrowed into more or less long, expanded petiole; cauline leaves oblong
or lanceolate, sessile, semiamplexicaul, entire or toothed. Capitula 1-
5(10), at stem apex. Involucre in a single whorl, broadly campanulate,
7 mm long and 15 mm in dia; involucral bracts oblong-linear, acuminate,
greenish, pubescent, apically dark purple. Ligulate florets 12-15, orange,
2 cm long and 3 mm wide. Achenes glabrous or pubescent, 2.5—3.0
mm long; pappus white, slightly shorter than tubular florets. Flowering
July to August.

Hummocky and meadow tundra, stony mountain slopes, alpine
meadows.—Arctic: Arctic Europe (Gydansk tundra, Polar Urals), Arctic
Siberia, Anadyr, Chukotka; Eastern Siberia: Lena-Kolyma; Far East:
Okhotsk, Kamchatka, Endemic. Described from Gydansk tundra from
the lower reaches of the Yenisei and other rivers. Type in Leningrad.

87. S. subfrigidus Kom. Fl. Kamch. III (1930) 169.—S. malaisei
Hult. in Svensk. Vet.-Akad. Handl. Ser. 3, VIII, No. 2 (1930) 199.—
Ic.: Hult. Ibid. Pl. 6, fig. a and page 199, fig. 3 (fl.).

Perennial. Rhizome almost horizontal, not thick. Stem erect, sulcate,
sparsely pubescent, simple or somewhat branched, 10-20 cm high.
Basal and lower cauline leaves long-petiolate, glabrous, their laminas
almost spatulate, with a few teeth; upper leaves sessile or subsessile,
broadly lanceolate. Capitula solitary or 2-5. Involucre about 1 cm in
dia; involucral bracts lanceolate, acute. Ligulate florets about 15-20,
yellow, 8-10 mm long. Achenes glabrous, ribbed; pappus after flowering
longer than florets. Flowering July to August.

Mountain forests with dwarf pine, on sandy-clayey mountain
slopes.—Arctic: Anadyr; Far East: Okhotsk, Kamchatka. Endemic.
Described from vicinity of the village of Tigil. Type in Leningrad.

88. S. pseudoaurantiacus Kom. in Fl. Kam. III (1930) 168.—S.
bogdanoviczii Kom. Ibid. (1930) 170.

778

741

Perennial. Rhizome short, oblique, covered with dark brown rem-
nants of dead leaves. Stem sulcate, together with leaves arachnoid-
hairy, 20-35 cm high, simple. Basal leaves oblong-obovate, toothed,
3-10 cm long and 2-3 cm wide, narrowed into winged petiole; cauline
leaves like basal but sessile, semiamplexicaul, acuminate. Capitula 1-
3(5), in corymbose inflorescence; peduncles arachnoid-hairy. Involucre
10 mm long and 15 mm in dia; involucral bracts lanceolate-linear,
densely arachnoid-hairy in lower part, glabrous above or puberulent,
green or dark purple. Ligulate florets orange-yellow, 1.6—2.5 cm long
and 2-3 mm wide. Achenes sparsely pubescent. Flowering July to
August.

Grass-mixed herb meadows, mountain tundra, alpine meadows.—
Far East: Kamchatka, Okhotsk. Endemic. Described from Klyuchevaya
Sopka [mud volcano]. Type in Leningrad.

89. S. sumneviczii Schischk. and Serg. Sistem. Zam. Gerb. Tomsk.
Univ. 1-2 (1949); Kryl. Fl. Zap. Sib. XI, 2856.—S. aurantiacus Kry].
Fl. Alt. Tomsk. Gub. III, 679, non DC.—S. integrifolius (L.) Clairv.
ssp. atropurpureus (Ldb.) Cuf. var. taimyrensis Cuf. f. nekrassovii
Cuf. Repert. spec. nov. LXX, I-III (1933) 45.—Cineraria frigida
Richards. B. robusta a. atropurpurea Herd. in Bull. Soc. Nat. Mosc.
XL (1867) 44, 45.

Perennial. Rhizome obliquely ascending, densely covered with
slender roots. Stem erect, simple, 12-50 cm high, usually floccose-
arachnoid-hairy. Lower leaves ovate, apically round or subobtuse,
narrowed into petiole almost as long as lamina, lamina 2—9 cm long
and 1.0-3.5 cm wide; middle and upper leaves oblong or ovate-
lanceolate, sometimes lanceolate-linear, 3.5—6.0 cm long and 3-10 cm
wide, obtuse or acute; all leaves irregularly and finely toothed, weakly
arachnoid-hairy. Capitula at stem apex, 2-4 or even solitary, lacking
ligulate florets, 10 mm long and 6-10 mm wide. Outer involucral bracts
1-2 or even absent; inner bracts linear, more or less pubescent or
subglabrous, often violet. Ligulate florets light or orange-yellow, 2-3
times as long as capitula[sic., recte involucre]. Achenes terete, about
3-4 mm long, ribbed, glabrous or pubescent (mainly on ribs).

Rocks, stony slopes, in alpine zone and occasionally in alpine
tundra.—Western Siberia: Altai; Eastern Siberia: Angara-Sayans; Soviet
Central Asia: Dzhungaria-Tarbagatai (Saur Tarbagatai). Endemic.
Described from Altai. Type in Tomsk.

90. S. reverdattoi Sobolevsk. Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XIV (1951) 50.—S. fedorovianus Schischk. ibid. XVI
(1954) 435.—Ic.: Sobolevsk. loc. cit. Fig. 2.

779

742

Perennial. Rhizome short, vertical, with long and slender roots,
covered with blackish-brown remnants of dead leaves; whole plant
arachnoid-hairy. Stem solitary, erect, straight, more or less sulcate,
usually violet, 12-30 cm high. Basal leaves clustered into rosette,
narrowly ovate or oblong-ovate, gradually narrowed toward base into
flattened petiole, obtuse, with fine, remote, violet teeth, lamina 2.0—6.5
cm long, 0.7-2.0 cm wide; other leaves sessile, semiamplexicaul,
lanceolate or linear-lanceolate, acute, 6.0-7.5 cm long and 0.4—1.2 cm
wide, more or less adpressed to stem; upper leaves smaller and narrower.
Capitula 3-6, at stem apex, on arachnoid-hairy peduncles forming
umbellate-corymbose inflorescence, with 2—4 linear or almost filiform,
violet colored bracteal leaves at base. Involucre 5—8 cm long and almost
as much in dia, in single whorl; involucral bracts linear, dark violet,
long-acuminate, with scarious border, dorsally sparsely arachnoid-hairy.
Ligulate florets 8-10, egg-yellow, 10-12 mm long and 1-2 mm wide
or even absent (var. eligulatus Theod.); tubular florets with 5 teeth,
slightly violet in upper part. Achenes terete, ribbed, glabrous or with
occasional hairs, 4 mm long; pappus white, almost as long as florets.
Flowering July to August.

Alpine zone in alpine meadows.—Eastern Siberia: Angara-Sayans,
Lena-Kolyma. Endemic. Described from Tsagan-Shibetau Range. Type
in Tomsk; cotype in Leningrad.

Series 7. Jacutici Schischk.—Perennials with woody rhizome;
leaves white-tomentose beneath, small, entire; stem 8-10 cm high;
ligulate florets 8-10, egg-yellow.

91. S. jacuticus Schischk. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XVI (1954) 430.—Ic.: Schischkin, ibid., p. 431.

Perennial. Plant forming more or less dense tussock; rhizome
woody, ascending, 3-4 mm thick, strongly branched in upper part and
densely covered with white-tomentose dead leaves and few brownish,
dead stems of previous year. Flowering shoots[scapes], 8-10 cm high,
simple, violet, finely arachnoid-hairy. Basal leaves numerous, ovate or
broadly ovate, usually obtuse, entire, abruptly narrowed into petiole,
petioles shorter or slightly longer than lamina, lamina green and weakly
arachnoid-hairy above, densely white-tomentose beneath, 1-2 cm long,
0.7-1.3 cm wide; cauline leaves oblong-linear, 1-2 cm long and 1-3
mm wide; acute, arachnoid-hairy. Capitula solitary, at stem apex.
Involucre in a single whorl, 10 mm long and as much in dia; involucral
bracts lanceolate, acute, glandular-hairy. Ligulate florets 8-10, egg-
yellow, with 4 darker veins, about 6 mm long and 4.0-4.5 mm wide.
Achenes terete, glabrous; pappus white.

780

743

Rubbly, scrub-lichen mountain tundra.—Eastern Siberia: Lena-
Kolyma. Endemic. Described from Aldan highlands. Type in Leningrad.

Section 14. Obejacae DC. Prodr. VI (1837) 341; Boiss. Fl. or. III,
383; Rouy, Fl. Fr. VIII, 341.—Annuals, less often biennial; cauline
leaves more or less divided, auriculate or amplexicaul; ligulate florets
absent or slightly, or much longer than florets; involucre with a few
outer involucral bracts; florets yellow; achenes usually pubescent, less
often glabrous.

Series 1. Viscosi Schischk.—Plants annual, sticky; ligulate florets
yellow, slightly longer than involucre, achenes glabrous.

92. S. viscosus L. Sp. pl. (1753) 868; Ldb. Fl. Ross. II, 629;
Boiss. Fl. or. III, 386; Schmalh. Fl. II, 86; Grossh. Fl. Kavk. IV,
146.—Jacobaea viscosa Gilib. Plantae rar. et comm. Lithuaniae (1785)
30; Grossh. Fl. Kavk. IV, 146.—Ic.: Syreistsch. Ill. Fl. Mosk. Gub. III,
282; Fedtsch. and Fler. Fl. Evrop. Ross. 992.—Hegi, III. Fl. 791, 792;
Jav. and Csapody, Iconogr. Fl. Hung. 530 (1933).—Exs.: GRF No.
3497a, b; Herb. Fl. ingr. No. 330; Pl. Finl. exs. Nos. 393 and 983.—
Fl. Polon. exs. No. 376.

Annual. Whole plant glandular-hairy, with unpleasant odor, sticky.
Stem erect, usually branched from base or middle, less often simple,
5-50 cm high. Basal (early-withering) and lower cauline leaves more
or less long-petiolate, oblong or ovate, simple pinnate or almost twice
pinnate, 1-10 cm long and 0.5-3.0 cm wide, their lobes oblong,
irregularly coarsely toothed or, in turn, pinnatifid; upper leaves smaller,
lanceolate, toothed, amplexicaul, with large, entire or toothed auricles.
Capitula at apices of stem and branches, usually few aggregated in lax
corymb, less often solitary. Involucre campanulate, 6—8 mm long and
4—6 mm in dia; outer involucral bracts linear, 1-3(10), a third to a half
as long as inner; inner bracts linear, whitish along margin, acuminate
and blackish at apex. Ligulate florets yellow, linear, slightly longer
than involucre. Achenes terete, glabrous or sparsely pubescent, ribbed,
dark violet, about 4 mm long. Flowering July to August; fruiting
September.

Coastal and river sands, railroad embankments and near
dwellings.—European part: Ladoga-Ilmen, Baltic Region, Upper Volga,
Volga-Kama, Upper Dniester; Caucasus: Cis-caucasia, Eastern and
Western Transcaucasia. General distribution: Scandinavia, Central
Europe, Atlantic Europe, North America (introduced). Described from
Europe. Type in London.

781

744

93. S. calvertii Boiss. Diagm. Ser. II, 3 (1856) 32; Grossh. FI.
Kavk. IV, 146.—S. viscosus Boiss. Fl. or. III, 386, ex p.—B. viscosus
auct. cauc.; Trautv. in Tr. Peterb. Bot. Sada, IV (1877) 152, non L.

Annual. Like previous species but plant often less branched; leaves
simple pinnate; capitula somewhat larger; involucre about 10 mm long;
peduncles usually longer; achenes sparsely pubescent between ribs with
short and stiff hairs. Flowering June to July.

Herb slopes, along stream banks and gravel beds at 1,200-—2,100
m.—Caucasus: Ciscaucasia, Eastern, Western, (Karchkhal Mountain)
and Southern Transcaucasia. General distribution: Armenia and
Kurdistan. Described from vicinity of Zreerum [sic., Erzurum?]. Type
in Geneva.

Note. As mentioned above, the plant is difficult to distinguish
from S. viscosus L. and requires further study.

Series 2. Vulgares Schischk.—Plants annual, with simple hairs,
not sticky; ligulate florets absent; achenes pubescent.

94. S. vulgaris L. Sp. pl. (1753) 867; Ldb. Fl. Ross. II, 628;
Turcz. Fl. baic.-dahur. II, 88; Schmalh. Fl. II, 86; Grossh. Fl. Kavk.
IV, 146; Kryl. Fl. Zap. Sib. XI, 2838.—S. flosculosus Gilib. Fl. Lithuan.
III (1785-1787) 198.—Jacobaea vulgaris Claus in Goebel. Reise, II
(1838) 283.—Ic.: Fedtsch. and Fler. Fl. Evrop. Ross. 990; Syreistsch.
Ill. Fl. Mosk. Gub. III, 281; Hegi, Ill. Fl. VI, 2, 7; Javorka and Csapody,
Iconogr. Fl. Hung. (1933) tab. 530.—Exs.: Herb. Fl. Ingr. No. 323; FI.
Polon. exs. No. 154.

Biennial or annual. Whole plant more or less arachnoid-hairy or
subglabrous. Stem erect, 12—40(50) cm high. Basal and lower leaves
oblong-spatulate, narrowed into short petiole, or petiole almost as long
as lamina, toothed, withering early, 2-10 cm long and 0.5-2.5 cm
wide; middle cauline leaves oblong or lanceolate, deeply sinuate-
pinnately lobed, with irregularly toothed or entire lobes, sometimes
leaves only deeply toothed, acute, narrowed toward base, sessile, 3-11
cm long and 0.5—2.0 cm wide. Capitula campanulate, 6—8 mm long and
about 5 mm in dia, on often divergent, 1.0-3.5 cm long peduncles.
Outer involucral bracts 2-7, lanceolate, often blackish at apex, (1/6)
1/4-1/2 as long as inner bracts. Ligulate florets absent. Achenes
pubescent, about 2.5 mm long. Flowering July to August; fruiting August
to September.

Near dwellings, kitchen gardens, crops, gardens and parks, railroad
embankments, sandy banks of rivers and lakes, forest edges.—Euro-
pean part: Karelia-Lapland, Dvina-Pechora, Ladoga-Ilmen, Baltic
Region, Upper Dnieper, Upper Volga, Volga-Don, Volga-Kama, Middle

782

745

Dnieper, Upper Dniester, Crimea (mainly in vicinity of the Nikita
Botanical Garden); Caucasus: Ciscaucasia, Western Transcaucasia;
Western Siberia: Ob River Area, Irtysh, Upper Tobol; Eastern Sibe-
ria: Yenisei, Angara-Sayans, Dauria, Lena-Kolyma (very rarely city of
Yakutsk); Far East: Ussuri, Uda River Area, Kamchatka, Sakhalin,
Kuril Islands (Shimushir Island). General distribution: Scandinavia,
Central and Atlantic Europe, Mediterranean Region, northern Africa,
Asia Minor, Iran, Japan, northeastern China, Korea; introduced into
North America. Described from Europe. Type in London.

Economic Importance. The leaves of S. vulgaris contain alka-
loids, viz., senecionine and senecine, which are known antihemorrhagic
substances and are used as an extract and tincture in gynecological
practice.

95. S. dubius Ldb. Fl. Alt. IV (1833) 112; Ej. Fl. Ross. II, 628;
DC. Prodr. VI, 341; Fisch. and Mey. Ind. VII sem. horti Petropol.
(1840) 55; Kryl. Fl. Zap. Sib. XI, 2839.—S. vulgaris L. var. dubia
Trautv. in Bull. Soc. Nat. Mosc. XL (1866) 56; Kryl. Fl. Alt. III,
669.—S. vulgaris var. parvula Trautv. op. cit. (1866) 363.

Annual. Stem erect, sometimes slightly bent, 6-30 cm high,
branched, somewhat angular, glabrous or with occasional, rather long,
white hairs. Leaves oblong-lanceolate, semiamplexicaul, with remote
teeth or pinnatisect, with linear, indistinctly toothed or entire lobes, 2—
7 cm long and 3-10 cm wide, less often leaves linear and entire.
Inflorescence corymbose paniculate. Capitula on long (3.0-3.5 cm long)
peduncles with linear, entire, long-acuminate, bracteal leaves; capitula
narrowly campanulate, 6 mm long and 4 mm in dia in upper part. Outer
involucral bracts 1-8, lanceolate-linear, acuminate, adpressed to inner
bracts, a sixth to a fourth as long as inner bracts lacking black spot at
apex; inner bracts often dark at apex. Ligulate florets absent; tubular
florets yellow, slightly longer than involucre. Achenes sericeous, ribbed,
3.0-3.5 mm long. Flowering April to June; fruiting June to July. (Plate
XXXVI, Fig. 2).

Sandy, often damp places, alkaline meadows, salt marshes, gravel
beds of rivers, riparian woodlands, river sands, among chee grass.—
European part: Trans-Volga (reported from Kamyshin); Western Sibe-
ria: Irtysh; Eastern Siberia: Angara-Sayans (Tuva Autonomous Re-
gion); Soviet Central Asia: Aralo-Caspian Region, Lake Balkhash
Region, Syr-Darya, Dzhungaria-Tarbagatai, Tien Shan, Pamiro-Alai
Region. General distribution: Sinkiang, Mongolia, Kuldzha, Kashgaria,
Kwan Lun, Hansu, Shanghai. Described from northeastern Kazakhstan
(Chaganka River). Type in Leningrad.

783

746

Note. The plant is highly variable depending on the ecological
conditions. Under drought conditions, it barely measures 2-5 cm while
under adequately moist conditions, it may be 20 cm high and more.
Efforts have been made a number of times to segregate botanical
varieties, which, in my opinion, is purposeless.

Series 3. Sylvatica Schischk.—Ligulate florets 2-14, slightly
longer than involucre or as long; achenes 2-3 mm long, densely
pubescent.

96. S. sylvaticus L. Sp. pl. (1753) 868; Ldb. Fl. Ross. II, 629;
Schmalh. FI. II, str. 86; Grossh. Fl. Kavk. IV, 146; Syreistsch. Fl.
Mosk. Gub. III, 282; Fedtsch. and Fler. Fl. Evrop. Ross. 991.—Exs.:
GRF No. 319; Pl. Polon.iexs...No., 377;

Annual. Stem erect, branched, less often simple, sulcate, arachnoid-
hairy or pubescent, with simple hairs, sometimes mixed with glandular
hairs, 20-70 cm high. Basal leaves withering early; lower cauline leaves
short-petiolate; middle and upper leaves sessile or with auriculately
expanded amplexicaul base, their laminas ovate or oblong, 4-8 cm
long and 1.0—4.5 cm wide, pinnately parted, lobes oblong or oblong-
linear, deeply and irregularly pinnate or irregularly toothed with sharp
teeth, subglabrous and dark green above, lighter beneath sparsely
pubescent mainly along veins. Capitula numerous, at apices of stem
and branches; peduncles hairy, shorter than capitulum, forming lax
corymbose panicle. Involucre pubescent, 6 mm long and 3—4 mm in
dia, in 2 whorls; outer involucral bracts a sixth as long as inner bracts,
often with a blackish spot at apex; inner bracts linear, with scarious
margin, with a blackish spot at apex. Ligulate florets 8-14, very short
and twisted, as long as or longer than involucre, light yellow. Achenes
terete, ribbed, densely pubescent, 2-3 mm long; pappus white. Flowering
June to August; fruiting September.

Forest edges, pine woods, edges of marshes, sandbars, cut-over
areas and burned clearings in forests.—European part: Ladoga-Ilmen,
Baltic Region, Upper Dnieper, Upper Volga? Volga-Kama, Volga-Don,
Upper Dniester; Cauca-Caucasus: Western Transcaucasia (in vicinity
of Batumi, probably introduced). General distribution: Scandinavia,
Central Europe, Atlantic Europe, Mediterranean Region, North America
(introduced). Described from forests of northern Europe. Type in
London.

97. S. krascheninnikovii Schishck. in Bot. Mat. Gerb. Bot. Inst.
Akad. Nauk SSSR, XV (1953) 410.—S. pedunculatus Edge. in Trans.
Linn. Soc. XX (1851) non Sch. Bip. (1844); Boiss. Fl. or. Suppl.

784

747

301.—S. coronopifolius var. sphacelatus O. Hoffm. in Ova Paulsen
Olufsen’s second Pamir. Exped. (1903) 152.—S. coronopifolius auct.
non. fl. As. Med. non Desf.

Annual. Stem erect, 8-25 cm high, branched from base or above,
sparsely pubescent or subglabrous. Leaves obliquely upright, ovate-
oblong, 1.5—3.0 cm long and 0.4—1.5 cm wide, pinnatisect almost up
to midrib, with 2-4 pairs of linear, subobtuse lobes, toothed, with few
irregular teeth or almost entire, like stem sparsely pubescent. Capitula
campanulate, 5-6 mm long and 2.5—4 mm in dia in upper part, on
slender peduncles 1-5 cm long, with 2-4, linear, acuminate bracteal
leaves, usually numerous and forming lax, corymbose-paniculate or
paniculate inflorescence. Outer involucral bracts 1-5, narrowly linear,
a sixth to a fourth as long as inner bracts; all involucral bracts glabrous,
apically (inner ones) usually with blackish spot (var. sphacelatus O.
Hoffm.). Ligulate florets 2-7, slightly longer than involucre. Achenes
terete, 2.5-3.0 mm long, densely pubescent. Flowering July to August;
fruiting August to September. (Plate XXXVI, Fig. 1).

Mountain spruce forests, gravel beds of mountain streams, near
rocks, stony slopes, rubbly talus and occasionally in lucerne crops; up
to 3,800 m.—Soviet Central Asia: Tien Shan, Pamiro-Alai. General
distribution: Sinkiang, Tibet, Iran Region (Afghanistan), Indo-
Himalayas. Described from Dhawla Valley (Himalayas). Type (S.
pedunculatus Edg.) in London.

Series 4. Vernales Schischk.—Plants more or less densely
arachnoid-hairy; biennials, rarely perennials; leaves pinnatifid; achenes
densely pubescent.

98. S. vernalis Waldst. and Kir. Pl. rar. Hung. I (1802) 23; Ldb.
Fl. Ross. II, 630; Trautv. in Mélang. biol. II, 129; Boiss. Fl. or. III,
389; Schmalh. Fl. II, 86; Grossh. Fl. Kavk. IV, 147.—S. squalidus
MB. FI. taur.-cauc. II, 309, non L.—S. crassifolius Sibth. and Sm. FI.
Greeca IX (1837) 53 non Willd.—S. rapistroides DC. Prodr. VI (1837)
346.—S. chrysanthemifolius Ldb. Fl. Ross. Il, 630, non Lam.—S.
peduncularis Griseb. Spicil. II (1844) 224.—S. vernalis a. genuinus
and B. pauciflorus Lindm. Prodr. Fl. cherson. I (1872) ll; Fl. cherson.
I, 225.—S. euxinus Mind. in Ukr. Bot. Zhurn. XIII, 3 (1956) 60.—
Jacobaea squalida C.A.M. Verzeichn. Pfl. Cauc. (1831) 81.—Ic.:
Waldst. and Kit. op. cit. tab. 24; Sibth. and Sm. op. cit. 868; Hegi, III.
Fl. VI, 2, 781, 782, 785; Javorka and Csapody, Iconogr. Fl. Hung.
(1933) 530.—Exs.: GRF Nos. 976, 976a; Fl. Polon. exs. No. 46 and
No. 375; Bot. Mus. of University of Tartu, No. 179.

748

Biennial or annual. Stems solitary or 2—4(6), simple or branched
in upper half (sometimes from base), arachnoid-hairy and finely sulcate
in young stage, later subglabrous (var. glabrescens), 10-40 cm high.
Leaves oblong or broadly linear, arachnoid-hairy; basal and lower
cauline leaves oblong, more or less long-petiolate, withering early,
sinuate-lobate, their lobes horizontal, irregularly toothed with remote
and sharp teeth; other leaves sessile, with toothed, amplexicaul, auricles,
oblong-linear, pinnately divided, with oblong sharply toothed, 2-8 cm
long and 0.5—2.5 cm wide lobes. Capitula at apices of stem and branches
in corymbose inflorescence. Involucre 7 mm long and almost as much
in dia; inner involucral bracts linear, acuminate and with a black spot
at apex; outer bracts a sixth as long as inner bracts, narrowly linear,
with blackish spot at apex. Ligulate florets linear-oblong, yellow or
light yellow, less often almost white, 12-16, 1.5—2 times as long as
involucre, 2 mm wide. Achenes terete, pubescent, 2-3 mm long.
Flowering May to July.

Sandy meadows, stony and calcareous slopes, logged areas, scrubs,
railroad embankments, field borders.—European part: Ladoga-Ilmen,
Baltic Region, Upper Volga, Upper Dnieper, Middle Dnieper, Volga-
Kama (western), Volga-Don, Bessarabia, Lower Volga, Upper Dniester,
Crimea; Caucasus: All regions; Soviet Central Asia: Syr-Darya
(?introduced), mountainous Turkmenia; General distribution:
Scandinavia, Central Europe, Balkans-Asia Minor, Mediterranean
Region, northern Africa. Described from Balkans. Type in Vienna.

Note 1. I did not think it possible to recognize the species B. euxinus
Mind., recently split off by Minderova from S. vernalis Waldst. and Kir.
of southern Ukraine. The differences cited by the author between
the new species and S. vernalis, namely, a not very tall stem, more
branching, smaller leaves and capitula, and the absence of a blackish spot
on the involucral bracts, are hardly convincing and apparently result
from ecological conditions. On the basis of the last character, De
Candolle split off a separate species from S. vernalis from the vicinity
of Baku under the name S. rapistroides DC. Examination of much material
shows that the appearance or absence of blackish spot is not a reliable
character, and it is hardly possible to consider both S. rapistroides DC. and
S. euxinus Mind. as a separate species. For some reason S. rapistroides
DC. is not mentioned at all in the work of Minderova.

Note 2. Trautvetter (loc. cit.) reports that the plant becomes strongly
branched with age, and then the pubescence on the stem and branches
disappears.

99. S. sosnovskyi Sof. in Izv. Akad. Nauk Azerb. SSR, | (1957)
89.—S. vernalis B. caucasicus DC. Prodr. VI (1837) 345; Ldb. FI.

785

749

Ross. II, 630.—S. vernalis y. nanus Boiss. Fl. or. III (1875) 389, non
S. nanus Sch. Bip. (1847).—S. glacialis Marcov. in Tr. Yurevsk. Bot.
Sada, IV (1903) 239; Grossh. Fl. Kavk. IV, 146, non S. glacialis Sch.
Bip. (1845) and Wedd. (1855-—1857).—S. candolleanus Sosn. in Zhurn.
Russk. Bot. Obshch. XIV, 1 (1929) 86; Kolak. Fl. Abkh. IV, 248;
Mandenova, in Fl. Gruzii, VIII, 395; Grossh. Opred. Rast. Kavk. 466,
non Hook. and Arn. (1841).—S. vernalis var. leiocarpa Trautv. in Tr.
Peterb. Bot. Sada, X, | (1887) 116.

Biennial or annual. Root slender. Stem solitary or few, with leaves
arachnoid-hairy or glabrous, simple or branched, sulcate, 5-25 cm
high. Basal leaves withering early, with petioles as long as lamina,
lamina ovate or oblong, more or less toothed, 10-20 mm long and 3-—
7 mm wide; cauline leaves larger, lower ones narrowed into short
petiole, upper ones sessile, semiamplexicaul, 1-6 cm long and 3-10
mm wide, irregularly toothed, or shallow-pinnately divided, obtuse or
acute; uppermost leaves small, linear, acuminate. Capitula at apices of
stem and branches, 2-6, often forming corymbose inflorescence.
Involucre 6-7 mm long and almost as much in dia; outer involucral
bracts linear, about 10, a fourth to a third as long as inner ones; inner
bracts linear-lanceolate, dorsally keeled, acuminate and, like outer
involucral bracts, with a dark spot at their apices. Ligulate florets
yellow or sometimes pinkish-violet, ovate-oblong, 7-8 mm long and
3-5 mm wide. Achenes terete, weakly ribbed, pubescent (sometimes
glabrous), 2.5-3.0 mm long. Pappus almost as long as tubular florets.
Flowering July. (Plate XXXVI, Fig. 2).

Moraines, talus, edges of glaciers, alpine meadows, at 1,800-3,700
m.—Caucasus: Ciscaucasia, Dagestan, Eastern, Western and Southern
Transcaucasia (Mongokhlyu-Tapa). Endemic. Described from Glavnyi
[Main] Caucasian Range. Type in Geneva.

100. S. polycephalus Ldb. Fl. Ross. II (1844-1846) 630, non
Clarke (1872); B. Schischkin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XVI, 438.—S. vernalis auct. fl. cauc. ex parte.

Annual. Stem ascending at base, erect above, strongly branched
from middle, sulcate, with leaves sparsely arachnoid-hairy, 20-30 cm
high. Basal leaves withering early; cauline leaves sessile, oblong, 1.0—
3.5 cm long and 0.5-1.0 cm wide, upright, often deflexed downward,
pinnately cut up to midrib, their lobes linear-oblong, cartilaginously
thickened along margins, irregularly toothed with deltoid, sharp teeth;
uppermost leaves oblong or linear, almost entire. Capitula at apices of
stem and branches, numerous. Involucre 5 mm long and almost as
much in dia, glabrous or sparsely pubescent; outer involucral bracts
narrowly linear, a third to a half as long as inner ones; inner bracts

78

fo»)

750

linear-lanceolate, with narrow scarious margin, acute; all bracts with-
out blackish spot at their apices, less often with an inconspicuous spot.
Ligulate florets about 12, yellow, ovate-oblong, 4.0-4.5 mm long and
2 mm wide. Achenes terete, weakly ribbed, densely pubescent, 3 mm
long; pappus as long as tubular florets. Flowering June to July.

Southern slopes in scrubs.—Caucasus: Eastern and Southern
Transcaucasia (Zangezur). Endemic. Described from vicinity of Tbilisi.
Type in Leningrad.

Note. Ledebour in Fl. Ross. (loc. cit.) gives a brief but quite
accurate description of the species on the basis of Wilhelms’s speci-
mens collected from the vicinity of Tbilisi. Wilhelms’s collection was
repeated by Lipsky, but Lipsky’s specimens remained unidentified.
Boissier saw Wilhelm’s specimens and determined them as a variety
of Senecio vernalis but did not assign any name to the variety. I think
it is necessary to consider this plant as the separate species S.
polycephalus Ldb. All specimens collected were without roots, but, as
was supposed by Ledebour, it apparently is a perennial plant.

Series 5. Subdentati Schischk.—Plants subglabrous, annual, 10-
20 mm high, with pinnately divided or toothed, often entire leaves;
achenes densely short-pubescent.

101. S. subdentatus Ldb. FI. Alt. IV (1833) 110, non Turcz. (1837)
nec Cineraria subdentata Bge. Enum. pl. Chin. boreal. (1832) 39.—
S. subdentata &. ramosus Ldb. B. simplex Ldb. y. parvulus Ldb. Ic. pl.
Fl. Ross. V (1834) tab. 493, f. 7, f. 2, f. 3.—S. coronopifolius Desf.
var. subdentatus Boiss. Fl. III (1875) 890.—S. subdentatus Ldb. var.
fasciculatus Litw. in Spisok Rast. Gerb. Russk. Fl. VI (1908) No.
1876.—S. alabugensis Winkl. in Tr. Bot. Sada, XI (1890) 320.—Ic.:
Ldb. Ic. pl. Fl. Ross. V (1834) tab. 493, f. 1, f. 2, f. 3—Exs.: GRF No.
1876, 3496.

Annual. Stem branched from base or middle, including leaves
glabrous, 5—25 cm high. Leaves broadly linear or oblong. 2.5-—5.0 cm
long and about 2-10 mm wide, subobtuse, toothed, sometimes almost
entire, sessile, semiamplexicaui, with occasional hairs or glabrous; upper
leaves smaller; bracteal leaves linear, glabrous or sometimes ciliate.
Inflorescence corymbose-paniculate, lax; capitula upright, on long (1.5-
4.0 cm) peduncles. Involucre 6 mm long and 6 mm in dia in upper
part; outer involucral bracts few, linear or entirely lacking; inner bracts
oblong-linear, with scarious margin. Ligulate florets 1.5—2 times as
long as involucre. Achenes terete, densely pubescent, 3-5 mm long;
pappus white, 5-6 mm long. Flowering May to June.

787

751

Wormwood, sandy, and clayey steppes on dry and stony slopes, in
crops of common millet, wheat, and others, ascending mountains to
2,800 m.—Caucasus: Western Transcaucasia; Siberia: Altai, Irtysh;
Soviet Central Asia: Lake Balkhash Region, Syr-Darya, Amu-Darya,
Aralo-Caspian Region, Kara-Kum, Dzhungaria-Tarbagatai, Tien Shan,
Pamiro-Alai, mountainous Turkmenia. General distrubution: Mongolia,
Dzhungaria-Kashgaria. Described from Altai. Type in Leningrad.

Note. Cufodontis (loc. cit.) thinks that the name S. subdentatus
Ldb. cannot be retained for this plant because, in his opinion, it must
be assigned to a species of section Tephroseris on the basis of the
following considerations. In 1832, Biinge described a Chinese plant
from section Tephroseris under the name Cineraria subdentata which
in 1837 was transferred by Turczaninow (Bull. Soc. Nat. Mosc., X,
No. 7, 1837, 154) to the genus Senecio—S. subdentatus (Bge.) Turcz.
But Turczaninow overlooked the fact that, in 1833, Ledebour had
already described another plant under the name S. subdentatus Ldb.
and that this epithet must be retained for that plant, while the Chinese
perennial plant S. subdentatus from section Tephroseris should be
assigned another name. That name is S. pierotii Miquel. in Ann. Mus.
Bot. Ludg.-Batav, II (1886) 182 (cf.: Kitagawa, Lin. p. 470).

102. S. noéanus Rupr. in Bull. Phys.-Math. Acad. Pétersb. XIV
(1856) 231.—S. salsugionosus Schischk. in Spisok Rast. Gerb. FI.
SSSR, XI (1949) 164.—S. subdentatus var. pinnatipartitus Bge. Reliq.
Lehmann. (1851) 172; Trautvetter in Tr. Peterb. Bot. Sada, I, 274.—

* §. coronopifolius var. pinnatipartita Trautv. in Tr. Peterb. Bot. Sada,

V (1877) 444.—S. coronopifolius auct. Fl. As. Med. non Desf.—Exs:.:
GRF No. 3496.

Annual. Stem branched from base or almost simple, weakly
arachnoid-hairy or crisped-hairy, 10-20 cm high. Leaves pinnately
parted, broadly ovate or oblong, 2.0-3.5 cm long and 1-2 cm wide,
deeply lobed but not to midrib, like stem weakly arachnoid-hairy or
subglabrous, lobes lanceolate-linear, 1.0—-1.5 cm long and 1.0-2.5 mm
wide, acute or obtuse, entire or with few teeth, sessile, semiamplexicaul,
basally expanded like auricles or somewhat sagittate. Inflorescence lax
corymbose; peduncles 0.7-4.0 cm long, with few, small, entire bracteal
leaves. Capitula broadly campanulate. Involucre 5-6 mm long and, 5
mm in dia in upper part; inner involucral bracts linear; outer bracts S—
8, linear, acuminate, a sixth to a fourth as long as inner bracts, glabrous.
Ligulate florets 6-8, yellow, 1.5 times as long as involucre. Achenes
finely pubescent, terete, about 2 mm long. Flowering May.

Salt marshes and alkaline meadows.—European part: Lower Volga;
Caucasus: Ciscaucasia, Eastern, Western and Southern Transcaucasia,

788

752

Talysh; Soviet Central Asia: Aralo-Caspian Region, Kyzyl-Kum. Gen-
eral distribution: Iran, Asia Minor. Described from vicinity of Kotto.
Type in Leningrad.

Note. In many works on the Flora of Soviet Central Asia, this
plant is called S$. coronopifolius Desf. The latter was described from
North Africa. It is very close to S. noéanus Rupr., but differs by having
larger capitula, smaller achenes, and somewhat fleshy leaves. From
Africa S. coronopifolius Desf. was introduced to Sicily by ships (see:
A. Fiori, A. Beguinot, R. Pampanini. Flora Italica exsiccata, No. 674).

The following names are unexplained and doubtful, and also
erroneously reported for the USSR.

1. Senecio triangularis Hooker, Fl. bor. Am. I (1833) tab. 115;
Ldb. Fl. Ross. II, 642.—An American plant reported by Ledebour
from Sitka Island.

2. S. lugens Richard. in Bot. App. Frankl. Journ. ex. 2 (1823) 747;
Ldb. Fl. Ross. Il, 644.—An American arctic plant unknown in the
USSR.

3. S. gmelini Ldb. Fl. Ross. II (1844-1846) 645.—Apparently,
this plant does not belong to the genus Senecio L., as was noted also
by Ledebour.

4. S. aquaticus Hill. Veg. syst. II (1771) 120.—A western European
plant mistakenly reported by Ledebour from Kiev and the Far East.

5. S. rupestris Waldst. and Kit. Pl. rar. Hung. II (1805) 436; Ldb.
Fl. Ross. II, 635.—Mistakenly reported by Ledebour from the mouth
of the Volga River and Western Georgia.

6. S. chrysanthemifolium Poir. Encycl. meth. VII (1806) 96; Ldb.
Fl. Ross. Ii, 630.—Mistakenly reported by Ledebour from the southern
European part of the USSR, Crimea and the Caucasus.

7. S. kochii o. schkurii Ldb. op. cit. (1844-1846) 644.—Reported
by Ledebour from vicinity of Orenburg.

GENUS 1564. Ligularia Cass. Nom. Cons.!:?

Cass. in Bull. Soc. Philom. (1816) 198, non Duval (1809).—
Senecillis Gaertn. De fruct. et sem. II (1791) 453, tab. 173.—
Erythrochaete Sieb. and Zucc. in Abh. Akad. Miinch. IV, (1846) 188.—
Senecio sect. Ligularia Benth. in Benth. and Hook. f. Gen. pl. II (1873)
449 (excl. Farfugium).—Hoppea Reinchb. in Flora, VII (1824) 254,

'Treatment by A.I. Pojarkova.
*From the Latin ligula—tongue.

753

Wa ws S
: — —

=

fie
8
he
a Kf ctr ie
a 934 P
2 we
aa ern
ay + ae
A ae a
fe SD is \
oa \ to ee '
- 7 eS
me . Tes >
= Ss SE Seeere ie.
sett Ts ~
eg ae

Plate XXXVII.

Ithifolia Maxim.; 2 — L. correvoniana

789

(Alb.) Pojark.

1 — Ligularia ca

791

754

non Willd. (1801).—Cyathocephalum Nakai in Bot. Mag. Tokyo, XXIX
(1915) de

Capitula heterogamous; outer florets ligulate, in a single whorl,
fertile, sometimes with staminodes; disk florets tubular, with regular,
5 toothed, campanulate or tubular-campanulate corolla. Involucre
tubular, campanulate or goblet shaped, mostly with 1—3(5) small narrow
[supplementary] bracts at base. Involucral bracts 5-many, in 2 distinct
or indistinct whorls, but always dimorphic; inner bracts broader, with
wide scarious margin; outer bracts narrower, without or with very
narrow scarious margin. Anthers with very short, acuminate or obtuse
basal appendage; apices of stamen filaments (more probably sterile
lower part of connective) thickened and broadened forming so-called
antheropodia. Style branches slender, with a small conical or inside
flattened apical appendage, acute or roundish at apex, hairy on outer
side throughout or only in upper part, sometimes forming only a groove
at base of appendage, inside of style branches always glabrous, stigmatic
surface occupying entire inside of style branches or rarely as 2 raised,
wide, lateral stripes. Pappus of long, white, sordid white, brownish,
reddish-brown, or reddish, soft or stiff hairs, with very short barbs,
less often pappus hairs rather short (0.5—2.0 mm long) and then acicular-
toothed. Achenes always glabrous, finely ribbed-barbate. Common
receptacle glabrous, flat or weakly convex, finely alveolate. Perennial
herbs, usually with short thick rhizome, densely covered with fibers of
thick adventitious roots, less often with slender creeping rhizome and
slender adventitious roots. Leaves alternate, contorted in bud, mostly
cordate, reniform, ovate or ovate-oblong, less often oblong, mostly
sinuate-toothed, less often entire. Capitula of different size (small to
large), in racemose or corymbose, simple or compound inflorescences,
less often capitula solitary. Florets yellow.

Type of genus: L. sibirica (L.) Cass.

The genus consists of more than 150 species distributed mainly in
the temperate regions of Eastern and Central-Asia. A few species grow
in the Middle East, the Caucasus, and Europe.

Economic Importance. Almost all species of the genus Ligularia
are ornamental plants.

ki, Involucral bhai Trees: Biceps ikon e-dechat ss ddieenanen eee a etate dd Bae « ye
+ Involucral bracts fused into 3—4-toothed cup ...........:ccceeeeeee
05th. 3 Rig ee Pena tenn 34. L. schmidtii (Maxirn.) Maxim.
2. | Pappus hairs very short, 0.5—2.0 mm long, a tenth to a fourth as
long as tubular floret, acicular toothed \..:..Jt.....ctssscvcnscasanconcsenes 3
+ Pappus hairs 4.5-10.0 mm long, not more than a half as long as
tubular florets, SCADIOUS, sso <n-s-nappaasnae cirendttedslars> <aceensbetat dcammiline 4.

792

~ +

So +

10.

in

12:

755

Deaves always entire: .......0. 6200.08". 31. H. glauca (L.) O. Hoffm.
All or at least basal leaves sinuate-toothed ............ccccessceesseeseeeeeees
JU Aceeae ives 32. L. carpatica (Schott. Nym. and Kotschy) Pojark.
EPeapp Nes am INE HET. Bs TO AE NN ON lace sctes 3.
Pappus brownish or reddish-brown ..........:ccsscesessesseeseeeseeeeesneees 24.
Rhizome creeping, slender, with slender adventitious roots; capitula
large, solitary or 2—3(6); involucre cupulate, of numerous involucral
bracts; leaves reniform; plants of the Caucasus (subgenus

DIG CHGF PIE Rattle ha A oon TTR CA tede wcecesns 6.
Rhizome short, thick, with dense clusters of cord-like, fleshy,
AO VSILITIOUS TO ee LT Nt 8.

Stem leafless; capitula usually solitary .............cceceesssesseeeeseeeseeeees
tb Wa ree Serer otc, eens Land 38. L. correvoniana (Alb.) Pojark.
Stem leafy; capital Tey eave lene el eodiveedteenvoenes 7.
Cauline leaves 2, thin when dry, papery; peduncle below capitula
covered with thickish hairs; involucre of (7)9—14 flat (lacking thick
Hbsvon back) *iivalucral BYacle: Riis i ART eee
rneninteerténtetctmesrectecicice A 36. L. renifolia (C.A.M.) DC.
Cauline leaves (2)3—6, crowded; peduncle below capitula tomentose-
floccose; involucre of 18-22 bracts, dorsally with thick ribs in
LOWER at RR IN i teareeie 37. L. caucasica (MB.) C. Don.
Stem at base with several large, nonleafy, membranous bulging
sheaths; leaves oblong-elliptical, gradually narrowed into petiole,
not thick and not fleshy; capitula 1-3, with goblet-shaped involucre,
leDervin' dia 20k. ARI, gat 35. L. pavlovii (Lipsch.) Cretz.
Nonleafy membranous sheaths lacking at base of stem ............... 9.
Inflorescence simple raceme (sometimes only lower-most peduncles
bearing 2—3(4) capitula), or capitula 1-3 .........cecceseeeeeeeereeeeees 10.
Inflorescence compound, oblong-cylindrical, cylindrical-pyramidal,
or broadly corymbose panicle; capitula small, with 5—8(12)
MaVGlUCTal“bracis-AANe AR ti Le, ge, 18.
Leaves glaucous, somewhat thickish, fleshy or more or less
coriaceous, stiff, ovate to oblong; inflorescence dense, often spicate

HASSE A sak eee ees US. ER. RA CNRS. nce: Tt:
Leaves green, not fleshy, soft, cordate, sagittate or reniform;
inflorescence more or less lax raCEME .............cccesccccssseccoeesceoes i.

Involucre always more or less pubescent dorsally; leaves often
covered with short, few-celled hairs, but often toothed ................
TLR RUA ahi ail iv eR 29. L. alpigena Pojark.
Involucre always glabrous; leaves always entirely glabrous and entire
Stash ancaapee desdens conte KSA Meee ToC ATA 30. L. altaica DC.
Short, 6-14 cm high, alpine plant; basal leaves small, 1.5—3.5(4.5)
cm long, round; capitula 1(2), large, with bowl-shaped involucre ....
esd ite HMO NONE abe pvt abner itavcanausassareonaiin 12. L. schischkinii Rubtz.

793

794

756

14.

15.

16.

a7

18.

20.

Plants taller and with larger leaves; if not tall (1440 cm high), then
leaves elongate, oblong-cordate, or Sagittate ...........cccscesseeeeeeees 13:
Very large plant, up to 2 m high; basal leaves up to 37 cm long and
40 cm wide; capitula large, aggregated in narrow, long (18-50 cm)
LACEING |. ocnvocsenscarnsnsevesesehh@hepleireedsenisere ee een 20. L. jaluensis Rom.
Plants much shorter; leaves much smaller; shorter and broader,
capitula often in almost corymbose raceMes ...........cssceeeseeeeseeee 14.
Leaves elongated, oblong-cordate or sagittate; involucre narrow,
campanulate; capitula (1)3-6(10), aggregated in lax raceme
Beh «Leb ga ode EMRE: OE RT ea 23. L. knorringiana Poljak.
Leaves roundish-cordate or reniform; involucre wider, goblet- shaped
essasunssochesposznapeob avons sgektnehses dh sud aaaed zeebb «te age ep eens oo aes FRSA by Sana ee 15.
Capitula large, including ligulate florets (5.0)6.5-7.5 cm wide,
solitary or 2-5, on remote, stiff peduncles 8-16 cm long ...........
«seusisbial spot iis, i eaiey Ja be apelin cae 15. L. robusta (Ldb.) DC.
Capitula smaller, (1)3—10(14), aggregated in dense or lax, sometimes
more or less corymbose raceme (sometimes with lower peduncles
bearing 2-4 capitula); peduncles soft ...........ceesscesseeeeneeeeeeeenees 16.
Leaves reniform, strongly broadened; hairs erect and very lax,
tomentose-arachnoid-hairy; corolla tube of tubular florets 2/5 as
longras limibiaiey. WANN 8... 18. L. karataviensis (Lipsch.) Poljark.
Leaves roundish ovate-cordate; pubescence more dense, floccose,
appressed tomentum; corolla tube of tubular florets 1/2—2/3 as long

as bina A schestt das sdde Biel canarias Dee real ces ei ae Ly.
Capitula (1)3-4(6), aggregated at stem apex on short peduncles 1-
5. cmiong is. sas ig har eee 16. L. kareliniana Stschegl.

Capitula 4-10, aggregated in lax raceme; middle and lower peduncles
up to 810 cm long? cists es cls RS. anlar
terry rece eer ey 17. L. narynensis (Winkl.) O. and B. Fedtsch.
Inflorescence compound, narrow, dense, cylindrical or cylindrical-
pyramidal panicle,with short lateral branches; leaves glaucescent,
thick and; sueculentoblong “adi dieetneslaaitagidaaed ae ee 19.
Inflorescence lax panicles, with more or less long lateral branches;
leaves not glaucous and with one exception (L. altissima) neither
thickynorisuicculent![sscc....i3a.. ¢dadlhs desis eee Bee Oe 21.
Involucre campanulate or bowl-shaped, involucral bracts 8-12, (8)9-
12 mm long; capitula with (5)6-8 ligulate and 12-20 tubular florets
aneandadadadistde bathe ddch obidod onaptedtaee «batekb aba ene «see 28. L. talassica Pojark.
Involucre narrower, tubular or tubular-campanulate; involucral bracts
3.5-8.0(9.0) mm long; florets in capitulum fewer, ligulate florets
146, tubular florets 4—T4C1G) oe bs ott eee eee eas cas 28.
General inflorescence oblong-cylindrical, in upper part a narrow,
dense, spicate, oblong raceme or panicle, usually considerably longer

735

23.

24(4).

757

than lower part; lower part with remotely branched peduncles bear-
ing racemose or paniculate inflorescence; involucre tubular-cam-
panulate, with 6-8 involucral bracts; ligulate florets (3)4—5; tubular
PUTS TNO RAG) sith teddies stscccscne 27. L. heterophylla Rupr.
General inflorescence more or less pyramidal, upper part shorter,
broader, usually ovoid; much shorter than lower branched part;
involucre narrowly tubular or obconical, with 5—6(8) involucral
bracts; capitula with 1-3 ligulate and 4~7 tubular florets ...........
CRE ot a). Se waa DasLidvicwediansent 26. L. macrophylla (Ldb.) DC.
Basal leaves broad, roundish-cordate or deltoid, often expanded ...
sorte ce ek tod thy PRUNE I, RIPE Os Fs cat satiaac ss batons docu tv aR IR ts 22.
Basal leaves elongate, oblong-ovate, or sagittate .........ccssee 23.
General inflorescence pyramidal panicle with numerous branches
bearing cylindrical or pyramidal individual inflorescences in upper
part; involucre tubular-campanulate, green; basal leaves broadly
cordate, light green, somewhat thick and succulent, glabrous or
sometimes weakly arachnoid-hairy beneath in younger leaves ....
UIE ect Te INDE Eiechadanssscrsnes 25. L. altissima Pojark.
General inflorescence corymbose-panicle, with lateral branches
bearing at their apices corymbose inflorescences; involucre narrow,
tubular-cylindrical, quickly turning golden-brown; basal leaves dark
green above, soft, roundish-ovate-deltoid or broadened, at least
young (leaves tomentose, beneath isi sscisicnnasid. dap atlaheineentbh.
WUReH a Cee, ese ath cbc 24. L. thomsonii (Clarke) Pojark.
Inflorescence with (30)60—300 capitula, elongated, often a more or
less corymbose raceme, with long branches, bearing capitula in
raceme or panicle; involucre very narrow, narrowly cylindrical
RM REET. Cb eal cds 22. L. songarica (Fisch.) Ling.
Inflorescence with (10)12-60 capitula aggregated in narrow raceme
with short branches bearing (1)4(5) capitula; involucre broader,
campanulate or almost goblet-shaped .............c:csssscssseseseessrseeeeeseeees
Ob. m8. 2001 Slit. cee Bok 21. L. thyrsoidea (Ldb.) DC.
Capitula aggregated in oblong, simple, dense raceme; involucre
natrow, cylindricalior campanitlate® 5. ;..iciiisccdciinessopissesscrsedecveceusevece 27.
Capitula 1-2 or more, usually aggregated in lax, broad, corymbose,
sometimes elongated raceme; involucre wide, goblet-shaped-
campanulate-or gobbet-sHApe se. isiisscdsdocssestsscuedesaissribionsobaetecensee oak
Involucral bracts densely covered with thickish, dark brown hairs
LS aca Baste AE REE anette hes ih abies ddvcban'vidnaderntaasedcoenarns 25.
Involucral bracts lacking dark brown hairs .........:ccsssceseseesees
oe, RANA Nek RIED a Pee eben tics aiGietitde ndcko ois 1. L. hodgsonii Hook.
Capitula (2)3-11, more or less in corymbose raceme; involucral
bowl-shaped-campanulate, 10-12 mm long and 0-12 mm in dia;

796

758

27.

28.

29.

30.

I

32.

basal leaves reniform or roundish-reniform, usually broadened, with
palmate! venation s2isenies: aaa! 2. L. trichocephala Pojark.
Capitula usually 1-2, less often 3-4; involucre goblet-shaped, 12-
14 mm long and 13-18 mm in dia; basal leaves usually ovate-
sagittate or roundish-cordate, elongated or more or less as long as
wide, with pinnate venation ..............ccee 3. L. calthifolia Maxim.
Leaves fleshy and succulent, light glaucous, ovate or oblong, with
keeled [thickened] truncate or weakly sinuate base; pappus hairs
Ste PSCLACEOUIS ANE...) tbh 33. L. mongolica (Turcz.) DC.
Leaves neither fleshy nor succulent, with deeply sinuate, cordate or
sagittate base; pappus hairs soft, not setaceOusS ...........cceeee 28.
Pappus shorter than tubular florets, as long as corolla tube or very
shehtly longer (Series Speciosae) xn. Axes. RE... 29.
Pappus as long as tubular floret or very slightly shorter .......... aL.
All bracteal leaves broad, ovate; involucral bracts glabrous on back
or with occasional appressed hairs; stem glabrous from base to
inflorescence or weakly arachnoid-hairy .............ssesseseeeeerseteeceneeseeens
» AIR Ge ait acter pune 5. L. splendens (Levl. and Vaniot) Nakai.
Only lower bracteal leaves broad, ovate or ovate-lanceolate, others
linear, narrow (1-2 mm wide); involucral bracts pubescent on back
with erect, brown, curly hairs; stem and leaves beneath more or less
densely. covered. withisimilar hairs icc... onl oe Ak 30.
Only 2-5 lowermost bracteal leaves ovate or lanceolate, rest very
narrow, linear; involucral bracts 6-10 mm long; ligulate florets
with 4—-5(7) veins, tubular florets 9-10 mm long ........... ee
AG SUSHI. BATS Cl. eerie. 6. L. sachalinensis Nakai
Bracteal leaves in lower half of inflorescence broad; involucral
bracts 10—13(14) mm long; ligulate florets with 7-10 veins, tubular
florets 11-12 mm long .............. eee 4. L. fischeri (Ldb.) Turcz.
Plants small, not more than 70-75 cm high, with slender stem and
rather thin (0.5—2.0 mm thick), mostly very long, petioles of bracteal
leaves; lamina of bracteal leaves small, 1.5—8.0(10.0) cm long, less
often solitary, somewhat larger, with strongly developed lobes at
base, often sagittate; inflorescence small, with (1)3-15 capitula
(Series Lon sipedes) 252 71D.Gi. XN I TO ed AR Rene a2)
Plants large, with thick stem; basal leaves large, up to 25 cm long,
with thicker (to 7-10 mm) petioles; inflorescence with numerous
capitula, much longer (Series RACEMOSAE) ..........:ccssseecenseeeeneeeees 34.
Basal leaves usually elongate, sagittate or triangular-sagittate, with
long (often as long as remaining part of lamina) and usually
acuminate lobes at base, petiole 2.5-5 times as long as lamina;
inflorescence with (1)2—15 capitula, usually with 1—2(3) sessile

797

33.

34.

35.

=

759

scale leaves below resembling bracteal leaves, but lacking peduncles
in their axils; ligulate florets with very short oblong ligule (11-15
mm long and 3-4 mm wide); tubular florets with rather broad,

campanulate limb, as long as tube ................ 12. L. longipes Pojark.
Basal leaves mostly broadly deltoid-cordate, cordate-sagittate or
Pies BRR Ope no onne We eee onemeermer, UUmtcos ia gee kits | em 35.

Basal leaves toothed, broad or sometimes only as small, cartilaginous
mucro, their petioles mostly 1.5—2(3) times as long as lamina; ligulate
florets very narrow (linear or oblong); tubular florets with broadly
campanulate limb as long as corolla tube; scale leaves below
inflorescence usually absent, rarely one ........ 13. L. arctica Pojark.
Basal leaves sharply toothed to serrate, mostly with sagittate base
and long basal lobes at base, narrowed downward and usually
acuminate, their petiole 2.5-4 times as long as lamina; ligulate
florets wider; tubular florets with narrowly campanulate limb, much
shorter than corolla tube; scale leaves below inflorescence usually
FS. A 0... A naa 14. L. bucovinensis Nakai
Leaves glabrous beneath or hairy only on veins, less often sparsely
Baan yee tae eget. kA Sa ak La ba scepead die 35:
Leaves somewhat densely or densely silbiacent to tomentose beneath

Leaves (in herbarium) thin, up to 24 cm long and 22 cm wide;
inflorescence up to 32 cm long, with many, usually 30—80 capitula;
ligulate florets 7—8(11); tubular florets (18)23-32 with narrow,
tubular-campanulate, limb 1.5 times as long as tube ................:006
sp TBR AI As LAB IIR Ba AIO Ow Ads 7. L. sibirica (L.) Cass.
Leaves thick, almost coriaceous, up to 11 cm long and as wide;
inflorescence 3—12 cm long, with (1—-3)5—10 capitula; ligulate florets
4-7; tubular florets up to 20, with campanulate limb, as long as
fhe BAL. WLI PARRA ld 10. L. sichotensis Pojark.
Plants up to 130 cm high, stem 3-10 cm thick; leaves in herbarium
thin, basal leaves 22 cm wide, mostly expanded deltoid-cordate and
deltoid reniform, with deeply cordate or hastate base and strongly
divergent lobes at base, somewhat densely or densely pubescent
beneath, soft, grayish-tomentose; inflorescence 6-32 cm long, with
10-36 capitula; involucre cylindrical-campanulate ...................+
2 DRO Sd ORG. OE RO 6. L. lydiae Minder.
Plants not as large stem up to 70 cm or up to 90 cm high and 3.5-
8.0 mm thick; basal leaves with erect, brownish hairs (not soft-
tomentose) and smaller, up to 15 cm wide; inflorescence with fewer
Capieiia AGES KAA UR. RERUN WANE A UR le eend 37:
All bracteal leaves (at base of peduncles) broadly ovate; ligule of
ligulate florets obovate or oblong, with (5)7—10 veins; limb of tu-

798

760

bular florets 1.5—-1.75 times as long as tube; scale leaves below
inflorescence 1-3, like bracteal leaves but lacking peduncles in
their axils; basal leaves broad, mostly roundish-cordate deltoid-
cordate, or cordate-reniform, 6.5—12.0 cm long and 7-10 cm wide,
usually densely hairy beneath, less often somewhat densely hairy
With Dre wants hha airs ais issstiscennesansnannssneperernncdbess 9. L. abakanica Pojark.
+ Bracteal leaves lanceolate; ligule of ligulate florets linear-oblong
and oblong-elliptical, with 4—5 veins; limb of tubular florets as long
as tube or somewhat longer, (1)3-6 scale leaves below inflorescence;
basal leaves narrower, up to 1.5(2) times as long as wide, 5—18(24)
cm long and 4.5—15.0 cm wide, mostly cordate or cordate-sagittate
less often deltoid-sagittate, somewhat densely or sparsely pubescent
beneath with brownish haifs ................. 11. L. subsagittata Pojark.

Subgenus |. Ligularia.—Senecio (sect.) Ligularia Benth. in Benth.
and Hook. f. Gen. pl. II (1876) 449.—Senecio B. Eu-ligularia Fr. in
Bull. Soc. bot. Fr. 39 (1892) 284.—Ligularia O. Hoffm. in Pflanzen-
fam. IV, 5 (1894) 301 p. p. (excl. Farfugium).—Ligularis sect.
Cyathocephalum (Nakai) Kitam. Compos. Japan. (1942) 185; Ligularis
sect. Senecillis (Gaertn.) Kitam. ibid. Ligularia sect. Euligularia Kitam.
ibid.—Rhizome short, thick; densely covered with fibers of filiform
adventitious roots. Capitula small, medium or large; involucre with 5
or more involucral bracts, usually free but sometimes (subsection
Stereochaete, series Schmidtianae) fused into cup, with 3-4 teeth in
upper part, style branches narrowed toward tip or with conical, or flat
short appendage, more or less acuminate or obtuse; stigma surface
usually occupying entire inner side of style branches, less often as 2
lateral raised stripes; antheropodia somewhat thicker than filament;
pappus mostly of long (more or less as long as achene), soft or stiff
scabrous hairs or (subsection Senecillis) much shorter than achene.

Section 1. Corymbosae (Fr.) Hand.-Mazz. in Bot. Journ. 69 (1939)
107 p. p.—Senecio groupe Ligularia B. Eu-ligularia 3. Corymbosi Fr.
in Bull. Soc. bot. Fr. 39 (1892) 290.—Erythrochaeta Sieb. and Zucc.
in Abh. Akad. Miinch. IV, 3 (1846) 188.—Ligularia sect. Euligularia
series Corymbosae Kitam. Compos. Japan. III (1942) 197.—Capitula
large or medium, aggregated in broad, mostly corymbose raceme,
sometimes 1-3; peduncles sometimes lacking bracteal leaves at base
or mostly arising from axils of large sessile scale leaves, at least lower
of these leaves keeled, enclosing peduncles and often involucre as
well; involucre broadly campanulate or goblet-shaped with (5)8—-12
involucral bracts; pappus reddish-brown or reddish, nearly as long as
tubular florets; antheropodia somewhat thicker than filament; style

799

761

branches long, slender, apically attenuate, with stigma surface occupy-
ing entire inner side.

Lectotype of section: L. calthifolia Maxim.

Species of eastern and southeastern Asia.

Our species of this section constitute the phylogenetic ‘series
Calthifoliae Pojark., characterized by the presence of large bracteal
leaves at the base of the peduncles as well as at the base of the involucre.

One other series of geographical races from Western and Central
China, described as varieties of L. hodgsonii and L. yesoensis, should
be referred here.

1. L. hogdsonii Hook. in Curtis Bot. Mag. XX (1863) subtab.
5417; Koidz. in Bot. Mag. Tokyo, XXIV (1910) 264 p. p. (excl.
synom.); Kitam. Compos. Japon. III (1942) 197 p. p. (excl. var.
sachalinensis.—L. clivorum Maxim. in Mélang. biol. VII (1870) 555
p- p- (quoad pl. e Hakodate).—Senecio yesoensis Fr. in Bull. Soc. bot.
Fr. 39 (1892) 306.—L. hodgsonii var. calthaefolia Koidz. op. cit. (excl.
synon.).—L. calthaefolia auct. non Maxim. (1870): Matsum. Ind., pl.
Japon. II, 2 (1912) 656; 1857; Vorobiev in Tr. Dalnevost. Fil. Akad.
Nauk SSSR, Ser. Bot. III (V) (1956) 76.—Senecillis hodgsonii Kitam.
in Acta Phyt. Geobot. VIII (1939) 82.—Ic.: Hook. loc. cit. tab. 5417.

Perennial. Plant with short, thick, fleshy rhizome covered with
numerous fibers of thick cord-like roots, but with stiff fibrous remnants
of dead leaves in upper part. Stem 25-100 cm high and 2-10 mm
thick, succulent, sulcate, purple below, entirely glabrous or pubescent
above with coarse, brown and white arachnoid hairs. Leaves bright
green, lighter beneath, on both sides glabrous, glaucous; basal leaves
2-3, fleshy, not winged but with semiamplexicaul sheath only at base,
petioles glabrous, 1.5—4 times as long as lamina, lamina 6—27 cm long
and 8-35 cm wide, reniform, broadly notched at base or cordate-
roundish, with deep and narrow notch, and obtuse lobes at base, usually
obtuse, sometimes abruptly attenuated into acute apex, with sharp and
often irregular teeth, usually with well developed palmate or pinnate
venation; caulines leaves 1—3(4), ones from lower part of stem of same
shape as basal but most often smaller and with shorter and thin petiole
expanded into long sheath, upper leaf (or leaves) with small or almost
reduced, deeply toothed lamina and short-winged petiole, greater part
of petiole enlarged in sheath. Inflorescence 2.5—13.0(26.0) cm long,
elongate or often corymbose raceme, with (4)5-10(15) capitula;
peduncles at anthesis slender, soft, later thickened, covered with
somewhat thick brownish hairs, upper ones 6—25 2.5(4) cm long, lower
up to 5-8 cm, at fruiting up to 14 cm long, all peduncles white-
arachnoid-hairy and with large, broad, keeled and glabrous bracteal

800

762

leaves at base, enclosing peduncle, arachnoid-hairy and toothed along
margin, attenuate at apex; lower bracteal leaves 3.5—-5.0 cm long, upper
2-3 cm, often longer than corresponding peduncles. Involucre at base
densely arachnoid-hairy, bowl-shaped, 8-16 mm long and at flowering
10—18(20) mm in dia in middle, with 2 apical bracteal leaves at base
longer than it, broad, keeled or sometimes narrow to lanceolate; in-
volucral bracts 5-12, always lacking dark thick hairs, glabrous or weakly
arachnoid-hairy, sometimes purple; inner bracts broadly ovate or ob-
long-ovate, with wide scarious border; outer bracts lanceolate to al-
most linear, with or without narrowly scarious margin; all involucral
bracts acuminate and arachnoid-hairy at tip. Capitula including ligulate
florets 4—7 cm in dia; ligulate florets 7-14, ligule oblong, 15-25 mm
long, 3-6 mm wide, with 6-10 veins, tube 5-7 mm long, sometimes
filiform staminodes present. Tubular florets numerous, 8—11 mm long,
with narrowly campanulate limb 4.5-6.0 mm long, gradually narrowed
into 3.5-5.5 mm-long tube; basal appendage of anthers very short,
subobtuse; style branches narrowed toward apex and obtuse, outside
approximately up to middle covered with flattened hairs. Pappus pale
chestnut-colored, at fruiting reddish-chestnut colored, distinctly shorter
than corolla; achenes 6-8 mm long, with almost as long pappus.
Flowering July to first half of August; fruiting September.

Herb meadows, open birch groves, herb and stony slopes of
mountains up to 700 m.—Far East: Sakhalin (south Kuril Islands).
General distribution: Japan. Described from Hakodate on Hokkaido
Island. Type in Kew.

2. L. trichocephala Pojark. sp. nova in Addenda, XXV, 886.—L.
hodgsonii var. sachalinensis Naki in Bot. Mag. Tokyo, XXIV (1910)
264; Kitam. Compos. Japon. III, 198.

Perennial. Differs from the previous species by having peduncles
and involucral bracts that are always densely covered with thickish,
dark brown hairs. The capitula are usually smaller and the involucres
usually 10-15 mm long and 10-17 mm wide.

Wet mixed-herb meadows, birch grooves.—Far East: Sakhalin
(southern part of Sakhalin and Moneron islands). Endemic. Described
from vicinity of Yuzhno-Sakhalinsk. Type in Leningrad.

3. L. calthifolia Maxim. in Bull. Acad. Sc. Pétersb. XV (1871)
374 in Mélang. biol. VII, 4-5 (1870) 554; Kom. Fl. Manchzh. III,
696; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1053.—Senecio
calthaefolius Maxim. in Mélang. biol. VIII, 1-2 (1870) 14; Franch. in
Bull. Soc. Bot. Fr. 39, 306.

80

—_

763

Perennial. Rhizome short, thick, with dense clusters of slender,
cord-like roots, covered with somewhat stiff fibrous remnants of dead
petioles in upper part. Stem erect, strong, 30-80 cm high and 2.5—5.0
cm thick, white-arachnoidhairy only in upper part (like peduncles with
involucre), and also covered, often very densely, with short, erect,
dark hairs, entirely glabrous below. Leaves dry, light green above,
much paler beneath, on both sides glabrous or with occasional, short,
dark hairs on veins beneath and with thin, pinnately divergent, quite
distinct secondary veins; basal leaves 3-6, with long (2—4 times as
long as lamina), thin, glabrous, petioles with sheath-like expanded base,
lamina cordate-ovate or cordate-roundish, 8-16 cm long and 6-12 cm
wide, usually strongly reduced (2—3 cm long and up to 7 cm wide),
cordate-reniform only in outermost leaves, with obtuse lobes at base
and obtuse or short, acuminate, evenly or unevenly crenate, toothed
with sharp, remote teeth; cauline leaves usually 2(1), lower like basal
leaves; but much smaller and with shorter petiole; upper, sometimes
both leaves, with small, deltoid-hastate lamina and very short, broadly
winged, amplexicaul petiole. Capitula 1-2, less often 3—4, including
ligulate florets 5.5—-7.0 cm in dia, aggregated at stem apex in corymbs,
lower capitula on long, up to 7-9 cm, slender peduncles, with large
bracteal leaf at base; bracteal leaves up to 3-4 cm long, sessile, carinate,
enclosing peduncle and involucre, lanceolate to ovate in unfolded form,
thin, acuminate, white-arachnoid-hairy and sometimes with short, dark
hairs, upper of them usually longer than peduncles. Involucre broadly
campanulate or bowl-shaped, 12-13 mm long, 13-18 mm in dia, in
middle at flowering with 2 large (mostly slightly longer than involucre),
wide, entirely folded bracteal leaves at base; involucral bracts (5-6)
8-12, leafy, more or less densely white-arachnoid-hairy; inner bracts
covered with somewhat thick, dark brown hairs, oblong-ovate or broadly
lanceolate with wide scarious margin; outer bracts lanceolate or
lanceolate-linear, with narrow scarious margin on both or on one side.
Ligulate florets with 20-30 mm long and 4-6 mm wide, ligule with 8—
12 veins, tube 7-9 mm long, tubular florets 12-13 mm long, with
narrow, campanulate-tubular limb and narrow tube approximately as
long; stamens with very thin and short basal appendages of anthers and
narrow antheropodia; style branches slender and long, contorted,
attenuate, with long hairs on outer side up to middle and below. Pappus
light brownish-white, later darker, reddish-brown; mature achenes
unknown. Flowering July. (Plate XXXVII, Fig. 1).

Dry, herb and stony slopes, from seacoasts to alpine bald zone (up
to 1,600 m).—Far East: Ussuri (only southern part of Sikhote-Alin
Mountains). Endemic. Described from coast of Olga Bay. Lectotype in
Leningrad.

802

764

Section 2. Ligularia.—Hoppea Reichnb. in Flora, VII (1824)
254.—Senecio groupe Ligularia B. Eu-ligularia 1. Racemosi Fr. in
Bull. Soc. bot. Fr. 39 (1892) 289 p. p.—Ligularia II. Racemosae Hand.-
Mazz. in Bot. Jahrb. 69 (1939) 117 p. p.—Ligularia sect. Euligularia
series Racemosae Kitam. Compos. Japon. III (1942) 190.

Capitula aggregated in simple oblong raceme (sometimes lower
peduncles bearing 2-3 capitula); peduncles arising from axils of rather
large scaly leaves, of which at least lower leaves carinately folded,
enclosing peduncle and involucre; pappus of brownish or reddish-brown
less often white hairs, as long as tubular floret or its narrow tube;
antheropodia somewhat or distinctly thicker than stamen filament; style
branches with acute or obtuse apical appendages; stigma surface
occupying entire inner side. Capitula medium-sized, with narrowly
cylindrical or campanulate involucre.

Type of section: L. sibirica (L.) Cass.

The species of this section are distributed in southeastern, eastern,
and northern Asia, the Caucasus and Europe.

Series 1. Speciosae Poljak.—Pappus distinctly shorter than tubular
floret or slightly longer than its tube; leaves usually serrate; achenes
blackish.

Besides the species distributed within our boundaries, a few, still
undescribed races from China should be referred here.

4. L. fischeri (Ldb.) Turcz. Catal. pl. baic. (1838) 11; Kitag.
Lineam. fl. Manch. 457 (excl. f.).—Cineraria fischeri Ldb. Ind. sem.
hort. Dorpat. (1820) 17.—Cineraria maxima Turcz. in sched.—
Cineraria speciosa Schrad. ex Link. Enum. pl. hort. Berol. II (1822)
334.—Hoppea speciosa Reichb. in Flora, VII (1824) 245.—L. sibirica
speciosa DC. Prodr. VI (1837) 315; Ldb. Fl. Ross. II, 620; Korsh. in
Tr. SPb. Bot. Sada, XII, 357.—L. speciosa Fisch. and Mey. Ind. sem.
hort. Petrop. V (1838) 38; Turcz. Fl. baic.-dahur. II, 84; Maxim. Prim.
fl. amur. 164; Kom. Fl. Manchzh. III, 693, p. max. p.; Kom. and Allis.
Opred. Rast. Dalnevost. Kr. II, 1053 p. p.; Ling in Contrib. Inst. Bot.
nation. Acad. Peip. II (1934) 535 p. p. (quoad pl. prov. Kirin).—
Senecio cacaliaefolius auct. non Sch. Bip. (1845); Maxim. in Mélang.
biol. VIII (1872) 14, p. min. p. (quoad synon. L. speciosa).—Senecio
cacaliaefolius 8. speciosa Franch. in Bull. Soc. bot. Fr. 39 (1892) 296
p- p. (quoad pl. dahur. amur.).—Senecillis fischeri Kitam. in Acta
Phyt. Geobot. VIII (1939) 82.

Perennial. Rhizome short, thick, glabrous, densely covered with
fibrous cord-like adventitious roots and with fibrous remnants of dead
leaves in upper part. Stem (30)50-150 cm high, sulcate, like petiole

803

765

covered with entangled, yellowish but later turning brownish, multicel-
lular, crisped hairs. Basal leaves with slender, long, petiole, up to 2.5—
3 times as long as lamina, with lamina 13-22 cm long and 22-30 cm
wide, cordate or cordate-reniform, sometimes almost hastate, obtuse
or short-acuminate, with broad or rather deeply lobed base, with short,
roundish or rather long, narrow, divergent basal lobes; cauline leaves
2-4, lowermost like basal, but with shorter petiole; next 1-2 leaves
smaller, usually acuminate, with short, broadly winged, amplexicaul
petiole; upper leaf small, ovate or lanceolate, sessile, with amplexicaul
base. Capitula 5-45, aggregated usually in dense or sometimes lax
simple raceme, less often lower peduncles branched, bearing 2-4 ca-
pitula; peduncles initially straight, upward spreading, later arcuately
deflexed, 5-30 mm long, lower ones sometimes up to 6, less often,
particularly when branched, up to 10-12 cm long, densely covered
with thick, multi-cellular, crisped hairs; bracteal leaves in lower part
of raceme large, ovate or ovate-lanceolate, long-acuminate, toothed,
often carinate, enclosing peduncle or capitulum, lowermost 3-5 cm
long, next smaller, as long as capitulum, with tubular florets, upper
ones narrow, lanceolate-linear, entire, all with pubescence like on
peduncles and often also with sparse, white, arachnoid hairs. Capitula,
including ligulate florets, (2.5) 3-4 mm in dia and with tubular florets
exserted from involucre over entire limb. Involucre with 2-3 filiform-
linear bracts at base, as long as involucre or shorter, tubular-campanu-
late, later bowl-shaped; involucral bracts 7-12 dimorphic, 10—13(14)
mm long, broader short-atuminate, with wide scarious border, nar-
rower long-acuminate and with narrow scarious border, back and apex
of involucral bracts densely covered with brownish, thick, crisped hairs.
Ligulate florets (5)6—9, with bright yellow, oblong or oblong-obovate, 7—
10-veined ligule, 15-25 mm long and 2.5-5.5 mm wide, and tube 6.0—7.5
mm long, often with filiform staminodes or few with abortive anthers;
tubular florets 18-35, with corolla (10)11—12(13) mm long, limb ob-
long-campanulate, 4-5 mm long, with narrow deltoid, 1.5-2.0 mm
long teeth; corolla tube somewhat longer, 5—6(6.5) mm long; stamen
tube considerably exserted from corolla; anthers with narrow, acute,
apical appendages and very short, acute, basal appendages, filaments
with rather broad antheropodia; style branches long, slender, narrowed
toward tip, on outside up to base covered with thick short hairs. Pa-
ppus brownish, somewhat longer than corolla tube; achenes immature
(probably, they do not mature) blackish-brown, narrowly terete, 6—9
mm long, ribbed; pappus slightly longer than achene. Flowering June
to July.

Wet meadows, scrub thickets and cut-over mixed, deciduous, and
rarely larch forests on moist soil.—Eastern Siberia: Angara-Sayans

766

(vicinity of Irkutsk and the village of Cheremkhovo, Irkutsk Region);
Dauria; Far East: Zeya-Bureya, Uda River area, Ussuri (only along
Amur and Ussuri rivers, in south up to Lake Khanka. General distri-
bution: Mongolia (northeastern part), China (northeastern part). De-
scribed from cultivated specimen. Type not preserved.

5. L. splendens (Lévl. and Vaniot) Nakai in Bull. Nat. Sc. Mus. 31
(1952) 117.—Senecio cacaliaefolius auct. non Sch. Bip. (1845); Maxim.
in Mélang. biol. VIII (1872) 14 p. p. quoad pl. e Kiusiu et excl. synom.—
Senecio cacaliaefolius var. speciosa Franch. in Bull. Soc. bot. Fr. 39
(1892) 296, p. min. p. (quoad pl. jap.).—L. sibirica auct. non Cass. (1823):
Matsum. and Koidz. in Bot. Mag. Tokyo, XXIV (1910) 149 (excl. syn.);
Koidz. in Bot. Mag. Tokyo, XXIV, 265 (quoad aream Japon. et excl. syn.
et var.).—L. speciosa auct. non Fisch. and Mey. (1838): Kom. Fl. Manchzh.
III (1907) 693 p. p.; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1053
p. p-—Senecio splendens Lévl. and Vaniot in Fedde, Repert. sp. nov. VIII
(1910) 139.—L. sibirica var. fallax Nakai, Reg. veg. ins. Quelp. (1914)
90.—L. sibirica var. araneosa Nakai in Bot. Mag. Tokyo, XXXI (1917)
117, non DC. (1837) nec Kom. (1907).—L. fischeri auct. non Turcz.
(1838); Koidz. Fl. symb. orient.-asiat. 88 p. p. (quoad pl. ex Yezo); Kitam.
Compos. Japon. III, 193 p. p. (quoad pl. Korean. japon. et excl. synon.);
Nakai in Bull, Nat. Sc. Mus. 31, 117; Hara, Enum. Spermatoph. Japon. II,
227 p. p. (quoad pl. japon. et excl. synon. praeter Lévl. and Vaniot and
Nakai).—L. sibirica ssp. fischeri Kitam. op. cit. (1942) 194, in synom.

Perennial. Rhizome as in two previous species, Stem 50-250 cm
high and up to 12 mm thick, ribbed-sulcate, glabrous from beginning
or more or less densely arachnoid-hairy ,less often, particularly below
capitulum, covered with short brown hairs. Leaves bright green and
glabrous above, lighter beneath, glabrous or more or less densely
arachnoid-hairy, less often sparsely, pubescent with occasional brownish
hairs; basal leaves 2, less often 3, with petioles 1.5-2 or more times
as long as lamina, hairy like stem or glabrous, slender, basally forming
narrow sheath, lamina 10-30 cm long, 12-40 cm wide, roundish-
reniform or cordate, usually obtuse, with recurved, roundish, usually
short lobes at base, sinuate-toothed or sinuate-serrate; cauline leaves
2—4, mostly 3, lower 1-2 leaves near base, like basal leaves but on
shorter petiole, next leaf much above, with small reniform or deltoid,
expanded lamina, broadly winged petiole modified into swollen sheath,
sometimes one more small, sessile, elongated leaf above, identical to
bracteal leaves. Inflorescence of (15)20-75 capitula, aggregated in
simple, 10-75 cm-long raceme; peduncles 3-12 mm long, lower ones
up to 3-5 cm, thick, arachnoid-hairy and sometimes also with very

805

767

short, brown, thickish hairs; all bracteal leaves broad, ovate, sessile,
sometimes lower ones pushed to lower part of peduncle, carinately
folded and enclosing peduncle and involucre, sometimes slightly longer
than involucre, glabrous or arachnoid-hairy, toothed or upper ones
entire. Capitula, including ligulate florets, 3—4 mm in dia, erect, later
only slightly deflexed. Involucre tubular-campanulate, later bowl-
shaped, with 2-3 narrowly linear bracts at base, as long as involucre
or much shorter; involucral bracts 8-9, in 2 whorls not distinct; inner
bracts broad- to oblong-elliptical, short- or long-acuminate, with wide
scarious border; outer bracts almost linear, usually long-acuminate,
with narrow scarious margin, enclosed in longitudinal axis formed by
spine and margin of neighboring involucral bract, all bracts glabrous
on back, only at apex tomentose-floccose and with very small, brown,
thickish hairs. Ligulate florets 6-8, with 7—10-veined ligule 15-24 mm
long and (3.5)4.5 mm wide, and 7-9 mm long-tube; staminodes observed
rarely; tubular florets 22-30, with corolla (12.5)13.5—16.0 mm long, their
limbs campanulate or tubular-campanulate, 6.5—7.5 mm long with nar-
rowly deltoid, 1.5—2.0 mm-long teeth, tube (5)6—8 mm long, stamens and
styles like previous species. Pappus as long as corolla tube or reaching up
to middle of limb, reddish-brown; achenes immature, 7-9 mm long, very
narrow, linear. Flowering second half of July to August.

Wet meadows and scrubs on marshy soil.—Far East: Ussuri (only
in extreme southwest in Posjet District along Suifun and on Russkoe
and Popov (Rynda) islands). General distribution: China (entire Korea
and Quelpart Island and, apparently, part of northeastern part of China
bordering Korea), Japan (Hokkaido, Honshu, Kyushu islands).
Described from Hallaisan on Quelpart (Cheju) Island. Type in France.

6. L. sachalinensis Nakai in Journ. Jap. Bot. XX (1944) 137.—
L. sibirica var. speciosa auct. non DC. (1837) Koidz. in Bot. Mag.
Tokyo, XXIV (1910) 265 (excl. synon.).—L. speciosa auct. non Fisch.
and Mey. (1838): Miyabe and Miyake, FI. Saghal. (1915) 270; Kom.
Fl. Manchzh. III, 694 p. p.; Kom. and Alis. Opred. Rast. Dalnevost.
Kr. II, 1053 p. p.; Sugaw. Ill. fl. Sagl. IV, 1858 (excl. synon. Trautv. and
Mey.).—L. fischeri auct. non Turcz. (1838): Koidz. Fl. Symb. orient.-
asiat. (1930) 48 p. p. (quoad pl. sachal.); Kitam. Compos. Japon. III,
193 p. p. (quoad aream Saghal. et excl. synon.); Sugaw. op. cit. 1859
(excl. synon.).—L. fischeri f. diabolica Kitam. in Acta Phyt. et Geobot.
IIT (1904) 170 (p. p.?); Hara, Enum. spermatoph. Japon. (1952) 227.—
L. speciosa var. araneosa Kom. Fl. Manchzh. III (1907) 695, non DC.
(1937).—Ic.: Sugaw. Illustr. fl. Saghal. TV (1940) tab. 853.

Perennial. Rhizome as in L. fischeri. Stem sulcate, usually sparsely
pubescent, with brownish hairs or subglabrous from beginning,

806

768

sometimes arachnoid-hairy below inflorescence. Leaves bright green
above, light glaucescent beneath, sparsely or densely covered with
erect, short, thickish, brownish hairs, sometimes mixed with white
tomentum; basal leaf usually 1 (sometimes 2), withering early, with
long, slender petiole up to 4 times as long as lamina, forming narrow
sheath at base, lamina cordate or reniform, up to 27 cm long and up
to 42 cm wide, mostly round at apex and with very short mucro, sinuate-
toothed, with deltoid, short, acute teeth, less often serrate; cauline
leaves 2-3, lower of them like basal leaf but with shorter petiole,
1/2—2/3 as long as lamina or as long, next leaf with much smaller
lamina and with short, broadly winged, often sheath-like expanded
petiole, upper leaf small, sessile, amplexicaul, sometimes one more leaf
above, like bracteal leaves in shape but larger. Inflorescence of (10)15—
55 capitula, often dense, up to 35 cm long, as simple raceme, less
often lower elongated peduncles bearing 2(3) capitula; peduncles 2—12
mm long, shorter than involucre or as long, less often slightly longer,
only lowermost peduncles up to 2—3(5) cm long; bracteal leaves, like
peduncles, covered with crisped, brown, long arachnoid-hairs, as long
as peduncles, with involucre including tubular florets exserted from it,
only lowermost 2-5, broad, ovate-lanceolate or lanceolate, toothed,
other bracteal leaves narrow, almost linear, entire. Capitula including
ligulate florets 2.5—3.0 cm in dia, with limb of tubular florets exserted
from involucre, initially erect, later drooping and adpressed to stem.
Involucre with 2(3) narrow, linear bracts at base, as long as or shorter
than involucre, tubular-campanulate; involucral bracts 6—8, dimorphic,
6-10 mm long, compactly imbricate because scarious margins of outer
bracts entirely enclosed in longitudinal axis formed by spine and wide
scarious margins of inner bracts; all bracts long-acuminate or broad,
(inner bracts) short-acuminate, on back densely covered with crisped
brownish hairs. Ligulate florets 2—6(8), ligule oblong, 10-15(20) mm
long and 2-4 mm wide, sometimes deeply lobed, with 4—5(7) veins,
tube 6-7 mm long; style with thin, long stylodes; staminodes often
present; tubular florets 17-24, with brownish pappus slightly longer
than corolla tube; corolla 9-10 mm long, limb oblong-campanulate,
4.5-5.5 mm long, with oblong-triangular teeth 1.25-1.5 mm long, and
tube 4-5 mm long, stamens and styles as in previous species. Achenes
blackish-brown, terete-fusiform, narrow, 5-7 mm long; pappus as long
as achene. Flowering July to August; fruiting from end of July.

Marsh meadows, scrubs, in forests on grassy areas and in cut-over
places with moist soil_—Far East: Uda River Area (basin of lower reaches
of Amur River and occasionally in Amur valley), Ussuri (southern and
eastern parts of district: from Vladivostok and Suchen, along eastern slope
of Sikhote-Alin), Sakhalin (Sakhalin Island). Endemic. Described from
Serulako on Sakhalin Island. Type in Tokyo University.

807

769

Series 2. Racemosae Kitam. Compos. Japon. III (1942) 190 p. p.
typ.—Pappus as long as tubular floret or somewhat shorter. Plants
usually large, with large leaves; stem thick; inflorescence usually with
several capitula; achenes brown.

Besides the species described here, the following European species
should also be referred to this series: L. cacaliiformis (Reichb.) Nakai
(1944) (Senecio cacaliiformis Reichb. fil. Icon. Fl. Germ. XVI (1854)
43 tab. 86, excl. syn. and area: Sibir. and Pyren.) and L. sebennensis
Rouy.

7. L. sibirica (L.) Cass. in Dict. Sc. Nat. XXVI (1823) 402; Ldb.
Fl. Ross. II, 620, p. p.; Turcz. Fl. baic.-dahur. II, 83; Schmalh. FI. II,
84 p. p.; Fedtsch. and Fler. Fl. 996 p. p.; Krasch. in Fl. Yugo-Vost.
Evrop. Ch. SSSR, VI, 373; Kryl. Fl. Zap. Sib. XI p. p. (saltem quoad
var. glaberrima Serg. and var. gigantea Serg.); Majevski, Fl. 593, p.
min. p.; Minder. in Ukr. Bot. Zhurn. XIV, 2, 45 p. p. and excl. fig.
1.—Cineraria sibirica L. Sp. pl. ed. 2 (1763) 1242 (quad aream Sibir.
and excl. synon. praeter Gmel.); Ldb. Fl. Alt. IV, 102, p. p.—Senecio
cacaliaefolius Sch. Bip., in Flora, XXVIII (1845) 50.—Hoppea sibirica
Reichnb. in Flora, VII (1824) 245; idem. Fl. Germ. exc. I (1830) 240
(quoad typ., excl. specim.).—Ic. Fedtsch. and Fler. loc. cit. Fig. 996;
Krascheninn. loc. cit. Fig. 692.

Perennial. Rhizome short, densely covered with thick or slender
adventitious roots, with fibrous remnants of dead leaves at apex. Stem
30-125 cm high and up to 10 mm thick, ribbed-sulcate, green or
reddish-violet below, up to inflorescence usually entirely glabrous, or
less often sparsely pubescent with few-celled brownish hairs. Leaves
thin when dry, less often thick, bright green above, on both sides
glabrous or sparsely hairy beneath along veins, less often entirely;
basal leaves (2)3, with mostly rather thick petioles, expanded into
short sheath, 2—2.5(3) times as long as lamina, lamina of different
shapes, mostly cordate or ovate-cordate, less often deltoid-cordate,
with length somewhat exceeding width, with deep and narrow, or more
or less broad incision, basal lobes somewhat divergent or recurved,
usually obtuse, sometimes reniform with strongly divergent, obtuse or
short-pointed-subacute lobes, (5.5)9-24 cm long and 7-22 cm wide,
sinuate-toothed, with deltoid, mostly broad, short-acuminate teeth; 1-
2 lower cauline leaves with slightly smaller lamina and much shorter
petiole narrowed into much longer sheath; upper cauline leaves with
small deltoid or deltoid-cordate, finely toothed lamina with short petiole,
greater part of petiole modified into very broad expanded sheath, very
rarely 1, brown or reddish, sessile, amplexicaul, scale-like leaf above,
like lower bracteal leaf but larger. Inflorescence of (5)10-—52 capitula,

808

770

simple, racemose, mostly oblong, up to 12 cm long, stem below inflores-
cence and peduncles sparsely or densely pubescent, with brownish hair
and often also with white arachnoid hairs; bracteal leaves on outer side
slightly tomentose, lower leaves ovate-lanceolate, long-acuminate,
sometimes in upper part toothed, 2.5-6.0 cm long, carinate, upper
ones ovate-lanceolate to linear, somewhat longer than involucre;
peduncles thick, lower ones up to 5—10 cm long, usually with 1, very
rarely 2 capitula, upper 5-12 mm long. Capitula including ligulate
florets 2.5—4.5 cm in dia, obliquely upright, later drooping. Involucre
at base with 2 lanceolate-linear or linear bracts as long as or slightly
shorter than involucral bracts; involucre campanulate or cylindrical-
campanulate, glabrous, often even at base or somewhat arachnoid-
hairy, indistinctly biseriate, consisting of 11-12 bracts 9-11 mm long,
inner lanceolate and outer lanceolate-linear or linear. Ligulate florets
7-8(11), with oblong-obovate, (10)14—17(20) mm long and 3-5 mm
wide ligule with 4-9 veins, tube 5-6 mm long. Tubular florets (18)23-
32, with 8.5-9.5 mm-long corolla, their limbs narrow, tubular-
campanulate, 4.5-5.8 mm long, with 1.0-1.5 mm-long teeth, but tube
usually shorter, 3.0-4.5 mm long; anthers with ovate-lanceolate apical
appendages and very short obtuse basal appendages, style branches
slender, at apices slightly flattened (spatulately or gradually) into short,
flat, obtuse, appendage, outside up to bifurcation covered with short hairs.
Achenes 5-6 mm long; pappus sordid brown, approximately as long as
achene. Flowering July to August; fruiting from second half of August.

Wet meadows, herb marshes, scrubs along riverbanks, alder
groves.—European part: Dvina-Pechora, Volga-Kama, Trans-Volga
(western part); Western Siberia: Ob River Area (southern half of dis-
trict), Irtysh (eastern part), Altai; Eastern Siberia: Lena-Kolyma (south),
Angara-Sayans, Dauria (western part); Far East: Okhotsk (southern,
part of the district: Ayan), Uda River Area (including Faklistov, Belichii,
Bolshoi Shantar islands); Soviet Central Asia: Lake Balkhash Region
(only at eastern border along Kokpektinka River). Described from
Siberia. Type in London.

8. L. lydiae Minder in Ukr. Bot. Zhurn. XIV, 2 (1957) 48.—
Senecio cacaliiformis auct. non Reichnb. f.: Meinsh. Fl. ingr. (1878)
173 (sphalm. “cacaliaefolius Reichb.”).—Cineraria sibirica Sobol.?
Sanktpeterb. Flora (1802) 163.—an Cineraria glauca Sobol.? Ibid.
163, non L. (1763).—Ligularia sibirica auct. non Cass. (1823): Rupr.
Fl. ingr. (1860) 602; Fedtsch. and Fler. Fl. Evrop. Ross. 996 p. p.;
Syreistsch. Fl. Mosk. Gub. III, 279; Majevski, Fl. 593 p. p.—Senecio
cacaliaefolius auct. non Sch. Bip. (1845): Schmalh. Fl. II, 86.—L.
sibirica var. bohemica Rupr. in Bull. phys.-math. Acad. Sc. Pétersb.

809

771

XII (1854) 213.—Ic.: Syreistsch. op. cit. Fig. on p. 279; Minder. loc.
cit, Figw3:

Perennial. Rhizome as in previous species. Stems 1-2, 50-130 cm
high, 3-10 mm thick, erect, mostly reddish-violet, particularly in lower
part, glabrous up to inflorescence, less often pubescent, with few-
celled, brownish hairs, densely below inflorescence and on peduncles
with similar hairs often mixed with white arachnoid hairs. Leaves bright
green and glabrous above, lighter beneath, scatteredly or somewhat
densely, often densely covered with soft, almost floccose, grayish,
multicellular, entangled hairs, along margin unevenly, rather often also
sinuate-toothed and with short, thickish hairs; basal leaves 1-3, with
glabrous, relatively slender petiole 2.5 times as long as lamina and
terminating into, elongated, narrow sheath, lamina 7-17 cm long and
7-22 cm wide, of rather different shapes: mainly deltoid-cordate,
deltoid-ovate or deltoid-reniform, with deeply cordate or broad,
sometimes weakly hastate base, more of less with divergent obtuse or
short-acuminate basal lobes, obtuse or short-acuminate; cauline leaves
2-3, lower cauline leaf like basal but with shorter petiole. Next leaf
near middle of stem or higher, deltoid-cordate or reniform, with short
petiole expanded into large inflated sheath, upper leaf small with petiole
modified into inflated sheath, less often 1—2 scale-like, sessile, carinately
(folded), amplexicaul leaves above it below inflorescence, like bracteal
leaves but larger. General inflorescence simple, racemose (less often
1-2 lower peduncles bearing 2-5 capitula), 6-32 cm long, of 10-36
capitula; peduncles very densely covered with erect, brownish hairs,
lower peduncles 2—6(7) cm long, others 4-12 mm long; bracteal leaves
brownish-purple, glabrous or weakly arachnoid-hairy, particularly along
margin, lower lanceolate, carinately folded, long-acuminate, with few acute
teeth in upper part along margin, others narrow, mostly linear, as long as
peduncle with capitula. Involucre with 1-2 linear bracts at base, often
slightly longer than involucre, cylindrical-campanulate in 2 indistinct
whorls; involucral bracts 7-11, dimorphic, 9-11(12) mm long; inner
involucral bracts wide, ovate or ovate-lanceolate, with wide scarious
border, short- or long-acuminate; outer bracts narrow, lanceolate-linear,
long-acuminate, with very narrow scarious border along one or both margins;
all involucral bracts green and glabrous on back, apically floccose.
Peripheral florets 5—8, with oblong or oblong-linear, 9-17 mm-long
and 2.5-4.0 mm-wide, usually seven-veined ligule, tube 3.5-6.0 mm
long; tubular florets 19-28, with corolla 8.0-9.5 mm long, its limb
3.25—4.0 mm long, rather narrow, tubular-campanulate, with 1.25-2.25
mm-long narrow-deltoid teeth, tube somewhat shorter, 3.0-3.5 mm
long; anthers and style branches identical with those of previous species.
Achenes terete, 5-7 mm long; pappus sordid yellow, scarcely longer
than achene. Flowering July to August; fruiting August to September.

810

772

Marshy meadows, herb marshes, wet scrubs and alder groves.—
European part: Karelia-Lapland (Karelia: materials from its eastern
part only), Dvina-Pechora (near western border of river), Baltic Region,
Ladoga-Ilmen, Upper Volga, Volga-Don, Upper Dnieper (northeastern
part). Endemic. Described from Leningrad Region, from Suida River
in vicinity of the village of Voskresenskoe southward from Gatchina.
Type in Kiev.

9. L. abakanica Poljark. sp. nov. in Addenda, XXV, 887.—Cin-
eraria sibirica auct. non L. (1763): Ldb. Fl. Alt. IV (1833) 102 p. p.
and excl. syn.—L. sibirica var. hirsuta Serg. in Kryl. Fl. Zap. Sib. XI
(1949) 2861.

Perennial. Rhizome short, with thick, filiform adventitious roots,
in upper part covered with fibrous remnants of dead leaves. Stems 1-
2, 20-70 cm high and 3.5-8.0 cm thick, erect, hard, deeply sulcate,
woolly or densely floccose throughout, with brown, entangled, flat,
multicellular hairs. Leaves thick when dry, light green above, sometimes
slightly colored from anthocyanin, somewhat lighter and densely or
almost densely pubescent beneath like stem, but hairs shorter, more or
less uniformly toothed with sharp teeth; basal leaves 1-3, usually 2,
mainly broadly or roundish-cordate or deltoid-cordate, elongated, less
often cordate-reniform, 6.5-12.0 cm long and 7-10 cm wide, short-
acuminate or obtuse, at base mostly with narrow, deep, less often with
broad notch, with thickish petiole, usually 1.5—2 times as long as lamina,
but sometimes as long or up to 2/3 as long, covered with erect and
dense or tomentose, brownish hairs; cauline leaves 1-3, most often 2,
lowermost (or 2 lower leaves) identical with basal leaves, usually (but
not always) with shorter petiole expanded like basal leaves, into narrow
sheath, upper leaf with small or very small deltoid lamina and short
petiole, major part of petioles (or entire petiole) modified into broad
inflated sheath; usually 1-3, sessile, amplexicaul cauline leaves above
it, identical to bracteal leaves in form consistency and color, but lacking
axillary peduncles. Inflorescence of 7-21 capitula, racemose, simple
(very rarely lower peduncle bearing 2-3 capitula), elliptical or oblong,
6.5—20.0 cm long, usually dense, peduncles thick, lower ones 10-12
mm long, less often up to 3.5(6.0) cm long, upper ones 2—8 mm long,
densely covered with brownish multicellular hairs, often like stem below
inflorescence, also with white arachnoid hairs; all bracteal leaves broad,
ovate, weakly white-tomentose on back particularly along margin, thin,
often colored from anthocyanin, often sharply toothed near apex, lower
ones carinately folded, 2-4 cm long, upper somewhat longer than
involucre. Capitula including peripheral florets 3.5-4.5 cm in dia,
obliquely upright, later almost horizontal. Involucre broadly campanulate,

831

—_

773

indistinctly in 2 whorls, basally pubescent; with 2(3) linear or linear-
lanceolate bracts at base; involucral bracts (7)9-11, dimorphic, glabrous
or weakly arachnoid-hairy, 9.5-11 mm long; inner bracts 3-4 mm
wide, with wide scarious margin, long- or short-acuminate, outer bracts
a half as wide as inner, lanceolate-linear, with very narrow scarious
border along one or both margins. Ligulate florets 7-12, with 15-20
mm long and 4—6 mm wide, (5)7—10-veined, oblong or obovate-oblong
ligule, their tubes 5.0-6.5 mm long. Disk florets 26-42, with 7.5-9.0
mm-long corolla, limb campanulate or less often narrowly campanulate,
4.5-5.0 mm long, with oblong-deltoid, 1.5 mm-long teeth, tube 2.75-
3.5 mm long; pappus whitish-brown, slightly shorter than corolla or
slightly short of reaching base of its teeth; anthers with short, ovate-
lanceolate, apical appendages and very short basal appendages,
antheropodia narrow; style lobes slightly enlarged at tip, capitate,
bearing fine papillae, uniformly hairy outside up to base. Achenes
narrow, terete, brownish, 5 mm long; pappus approximately as long.
Flowering July to August; fruiting second half of August.

Wet meadows, marshy banks of rivers and lakes.—Western Siberia:
Altai; Eastern Siberia: Angara-Sayans (western part of district;
Kuznetsk and Abakan ranges and their foothills; no material from
western Sayans). Endemic. Described from Uibat River valley in
northern part of Abakan Range. Type in Leningrad.

10. L. sichotensis Pojark. sp. nov. in Addenda, XXV, 888.

Perennial. Rhizome as in previous species. Stem 18-80 cm high,
2.0—3.5 cm thick, more or less purple, as also petiole, glabrous up to
inflorescence or (only in plants from Svetlaya River) covered with
erect, entangled, brown hairs. Leaves in herbarium specimens thick,
almost coriaceous, light green above, whitish glaucescent beneath
glabrous on both sides or sparsely hairy beneath along veins; basal
leaves with petioles a half to two-thirds as long as lamina or as long,
slender or (less often) somewhat thick, lamina 5-11 cm long and as
wide, cordate or roundish-cordate, with short obtuse lobes at base,
roundish or obtuse at apex, sinuate-toothed, with wide, sometimes
weakly developed teeth, quite often entire; cauline leaves absent or
one usually in lower half of stem, with small, 1.5—-2.0 cm long (less
often up to 4.5 cm) lamina and usually short petiole, major part of
petioles modified into sheath. Inflorescence 3-12 cm long, with (1-
3)5-10 capitula; all peduncles with single capitulum, short, usually
shorter than involucre (3-10 mm long), rarely lowermost peduncles
2.0—3.5 cm long, more or less densely white tomentose like stem below
inflorescence and covered with short erect hairs; all bracteal leaves

812 narrow, almost linear, or lowermost leaves narrowly lanceolate, green

774

Plate XXXVIII.
Ligularia narynensis (Winkl.) O. and B. Fedtsch.; 2 — L. pavlovii (Lipsch.)

813

1

Cretz.

815

775

or slightly purple and weakly arachnoid-hairy, usually all leaves longer
than corresponding peduncles; less often lowermost leaves shorter than
peduncles; usually 1-4 such leaves below inflorescence, but lacking
axillary peduncles. Capitula 3.5—4.0 cm in dia. Involucre campanulate
or campanulate-cylindrical with 2 long bracts at base, usually as long
as involucre; involucral bracts 7-8, (8)9-10 mm long, acuminate,
dimorphic; inner bracts ovate, with wide scarious margin; outer
lanceolate, with narrow scarious margin, glabrous on back, sometimes
with occasional thickish hairs or sparsely white-arachnoid-hairy.
Ligulate florets (4)5-7, with oblong-elliptical, 10-17 mm long ligule
and 5(7) mm long tube. Tubular florets about 20, with 8-9 mm long
corolla; their limb narrowly campanulate, as long as tube; style branches
spatulately enlarged into short roundish appendage. Pappus somewhat
shorter than corolla, reddish-brown; achenes (immature) 5—6 mm long.
Flowering: end of July to August.

Dry stony and mountain-meadow slopes, edges of forests, edges
of marshes, from seacoasts to alpine bald zone.—Far East: Ussuri
(only eastern slope of Sikhote-Alin) Endemic. Described from Botcha
River valley. Holotype in Leningrad; isotypes in Leningrad and
Vladivostok.

11. L. subsagittata Pojark. sp. nova in Addenda, XXV, 889.—
Cineraria sibirica auct. non L. (1763): MB. FI. taur.-cauc. II (1808)
313; C.A. Meyer. Verzeichn. 82.—Ligularia sibirica auct. non. Cass.
(1823): Ldb. Fl. Ross. II, 2 (1845-1846) 620, p. min. p. (quoad pl.
cauc.); Grossh. Fl. Kavk. IV, 155.—Senecio calcaliaefolius auct. non
Sch. Bip. (1845): Schmalh. Fl. II (1897) 84, p. min. p. (quoad pl.
cauc.).

Perennial. Rhizome short, densely covered with adventitious roots.
Stem at base with slender fibrous remnants of dead leaves, erect, strong,
sulcate-ribbed, glabrous up to inflorescence, purple at base and below
inflorescence, sometimes entirely, 25-90 cm high and 4—7 mm thick.
Leaves, when dry (herbarium material), rather thick, less often thin,
bright green and glabrous above, lighter beneath, glaucescent, entirely
covered with dense, less often sparse, short, simple, flat, white, later
usually brown hairs; basal leaves 2—3, with glabrous petioles 1.5—2.5
times as long as lamina, expanded into semiamplexicaul short and
narrow sheath, lamina more or less cordate or sagittate, less often
deltoid-sagittate, usually up to 1.5(2) times as long as wide, 5.5—18(24)
cm long, 4.5-15 cm wide, short-acuminate or obtuse, with deep a
notch and large lobes at base, acuminate or obtuse, recurved or more
or less divergent, toothed, with uneven, large, deltoid, acuminate or
sometimes obtuse teeth, with involute glabrous margin; lower cauline

816

776

leaf like basal leaves, but with shorter petiole and longer sheath, next
leaf with smaller lamina; and petiole usually not longer than lamina,
larger part of petiole modified into flat sheath; upper cauline leaf with
very small lamina and broadly inflated sheath, usually (1)2-6 sessile,
amplexicaul, scale-like, violet leaves above it like bracteal but larger.
Inflorescence simple, recemose, usually oblong. 4.5-30(34) cm long,
with 7-35 capitula, horizontal, later downwardly deflexed; stem below
inflorescence and peduncles sparsely or densely covered with erect,
thickish, simple, brown hairs, sometimes also with white arachnoid
hairs. Bracteal leaves large, brownish-violet, sessile, glabrous or weakly
arachnoid-hairy; all bracteal leaves lanceolate, finely, thin, long-
acuminate or upper leaves almost linear, lower 22-35 mm long,
carinately folded; upper leaves usually somewhat longer than involucre,
lower peduncles up to 3.5 cm long, upper 0.3—1.0 cm long. Involucre
campanulate-cylindrical or almost cylindrical, in 2 indistinct whorls,
with 2 bracts at base, almost as long as involucre; involucral bracts 8—
10, thin, papery, pale, greenish or more or less anthocyanin-colored,
10-12(14) mm long, glabrous or with occasional thickish, erect or
appressed arachnoid hairs; inner bracts lanceolate, with wide scarious
border; outer bracts linear, lacking scarious border and with very narrow
border along one or both margins; all bracts long-acuminate. Ligulate
florets 7-12, with oblong-elliptical or linear-oblong ligule, slightly
longer than involucral bracts, 8-15 mm long and 2.5-3.5 mm wide,
with 4-5 veins; tube 4.0-5.5(7.0) mm long. Tubular florets 17-32,
with 8-11 mm-long corolla; limb narrowly campanulate, 3.5-5.0 mm
long, with narrowly deltoid, 1.0-1.5 mm-long teeth; tube 4.0-5.5 mm
long; style branches curved, slender, slightly spatulately enlarged at
apex, roundish at apex, and covered with very small papillae, densely
covered on outside up to base, with short pointed hairs, inner side
almost entirely occupied by stigma surface. Achenes narrowly terete,
5-6 mm long; pappus whitish-reddish-brown, as long as achene.
Flowering July to August; fruiting September.

Wet alpine meadows.—Caucasus: Ciscaucasia, Glavnyi [Main]
Caucasus Range, Dagestan, Eastern and Western Transcaucasia,
reported also from Southern Transcaucasia. General distribution:
Armenia and Kurdistan (Kars Region). Described from vicinity of the
village of Kazbek. Type in Leningrad.

Series 3. Longipedes Pojark.—Plants occasionally more than 70
cm high, with slender stem and rather thin, and mostly very long petioles
of basal leaves; lamina of basal leaves small, 1.5(8) 10 cm long, mastly
with long lobes at base. Inflorescence of small number of capitula.

The following three species are included in this series.

817

777

12. L. longipes Pojark. sp. nova in Addenda, XXV, 890.—L.
sibirica auct. fl. sib. non Cass. (1823): Turcz. Fl. baic.-dahur. II (1846)
83 (and var. alpestris minor...); DC. Prodr. VI, 315, p. min. p. (quoad
pl. Sibir. baic.); Ldb. Fl. Ross. II, 620, p. min. p.; M. Popov, FI. Sr.
Sib. II, 742, p. max. p.—L. sibirica var. gracilis Serg. in Kryl. Fl. Zap.
Sib. XI, 2860 p. p.

Perennial. Rhizome short, thick, densely covered with filiform
adventitious roots, and in upper part more or less densely covered with
fibrous remnants of dead leaves. Stem 30-65 cm high and 1.5—2.5 mm
thick, slender, sparsely leafy, finely sulcate, glabrous, hairy only below
inflorescence, in lower part, but sometimes throughout, more or less
purple. Leaves thin when dry (herbarium specimens) but relatively
thick, light green above, paler beneath, glaucescent, on both sides
entirely glabrous, rather uniformly sinuate small-toothed, with broadly
deltoid teeth, abruptly narrowed into short callous acute tips; basal
leaves 2—3(4), petiolate, petiole glabrous, very thin, 0.75—2.0 mm thick,
and 2.5-5 times as long as lamina, often bright purple, lamina 2.3-
8.0(10.0) cm long and as broad, often elongated, sagittate, deltoid-
sagittate, or deltoid, less often reniform-sagittate or reniform-cordate,
mostly acute, less often obtuse, deeply lobed, with large lobes at base,
often as long as other part of lamina or up to two-fifths to two-thirds
as long, acute or roundish at apex; cauline leaves mostly 2, less often
3, lower leaf identical to basal but sometimes with shorter petiole and
broader amplexicaul base, upper leaf distant from it, with very small
lamina and petiole modified into broad, inflated sheath, sometimes one
more leaf with intermediate shape in between them in upper part of
stem; often 1—2(3) scale-like leaves below inflorescence like bracteal
leaves but usually larger. Inflorescence 1-2 (“‘L. sibirica var. alpestris
Turcz.”); capitula 3—15(20) aggregated in lax, always simple raceme,
3-10(20) cm long; capitula including ligulate florets 2.0-3.5 cm in
dia, on short peduncles; upper peduncles 2 mm long, lower 8-30 mm
long (in certain cases lowermost peduncle up to 4—7 cm long), obliquely
upright, later strongly drooping, parallel to stem; stem below
inflorescence as well as peduncles sparsely covered with thickish,
simple, white hairs, later turning brown. Involucre cylindrical and
narrowly campanulate with 1-2 filiform or lanceolate-linear bracts at
base; involucral bracts 7-8, 8-11 mm long, glabrous except floccose
apices, but more or less anthocyanin-colored; inner bracts lanceolate-
ovate with wide scarious margin, short-acuminate; outer bracts narrower,
linear or lanceolate-linear, finely long-acuminate. Ligulate florets 6-7,
usually with oblong 11-15 mm-long and 3-4 mm-wide ligule; tube
4.5-6.0 mm long. Disk florets 15-20, with 7.0-9.5 mm-long corolla;
limb rather broadly campanulate, 3-4 mm long, with about 1 mm-long

778

teeth, but tube 3-4 mm-long; anthers with narrow and long apical
appendage and very short, subobtuse, basal appendage, antheropodia
long; style branches slender with slightly enlarged roundish appendage,
on outer side up to base covered with short conical hairs. Achenes
light-colored, 5—6 mm long; pappus pale, weakly reddish-brown, slightly
longer than achene. Flowering July to August; fruiting September.

Marshy and saline meadows, river valleys, herb marshes, taiga and
steppe as well as alpine bald zone.—Western Siberia: Ob River Area
(southeastern part), Altai; Eastern Siberia: Lena-Kolyma (southern part
of river), Angara-Sayans, Dauria; Far East: Zeya-Bureya (except north-
ern part), Ussuri, including Pacific Coastal Region (evidently very
rarely). General distribution: Mongolia (northern part). Described from
Trans-Baikal Region from the village of Konstantinovka in Eravo Lakes
region. Type in Leningrad.

13. L. arctica Poljak. sp. nova in Addenda, XXV, 891.—L. sibirica
var. minor Nym. Consp. fl. Europ. (1879) 350.—L. sibirica var. gracilis
Serg. in Kryl. Fl. Zap. Sib. XI (1954) 2861 p. p.—L. sibirica auct. non
Cass. (1823); Fedtsch. and Fler. Fl. (1910) 996 p. p.; Hilt. Suom.
Kasv. (1933) 709 p. p.

Perennial. Rhizome as in previous species; plant wholly glabrous
excluding upper part of stem, peduncles, and base of involucre covered
with thick, brownish, simple, hairs. Stem 15-70 cm high, slender, 1.5—
3.0 mm thick, sparsely leafy, often entirely violet-purple, less often
green in middle. Leaves in herbarium specimens thin, but sometime
rather thick, bright green above, light green beneath; basal leaves usually
2, cordate, small, (1.7)2—7(11) cm long and 2.2-—7.0(10.0) cm wide,
deltoid-cordate, cordate, or cordate-sagittate, obtuse or short-acuminate,
with divergent obtuse or less often short-acuminate lobes at base, mostly
short, irregularly toothed, with deltoid, mostly broad teeth, with short
callous acute tip, sometimes with inconspicuous, remote teeth with
only | short cusp, their petioles thin, 1.5—2(3) times as long as lamina;
lower cauline leaf identical with basal leaves but with shorter petiole
and longer sheath, sometimes one more leaf in upper part with very
small lamina and petiole entirely or almost entirely modified into
amplexicaul sheath; sometimes | scale-like sessile, lanceolate leaf below
inflorescence like bracteal leaves but larger. Inflorescence simple
raceme, lax, with few, 2—7(12) capitula; lower peduncles 12—35(50)
mm long, upper 6—10 mm long; bracteal leaves brownish-violet, lower
ovate or more often lanceolate, long-acuminate, usually entire,
enveloping peduncle and involucre, upper lanceolate or linear. Capitula
including ligulate florets 2-3 cm wide, later drooping. Involucre
cylindrical or cylindrical-campanulate, with 2 linear or filiform bracts

819

779

at base; involucral bracts 6—-8(9), 6—10(12) mm long, glabrous, more
or less entirely purple; inner bracts lanceolate or almost ovate, with
wide scarious margin; outer bracts lacking scarious border or with
very narrow membranous border on one or both sides. Ligulate floret
5—6(7), with linear or oblong ligule 8-10 mm long and 1.25-4.0 mm
wide, with 3-6 veins; tube 3-4 mm long. Tubular florets 17—26, with
7-8 mm-long corolla; limb campanulate, often wide, 3.0-3.5 mm long,
with deltoid teeth, about 1.5 mm long usually abruptly narrowed into
narrow tube 3 mm long; anthers and style branches like previous species.
Achenes narrowly terete, 5.0-6.5 mm long; pappus whitish-reddish
brown, slightly longer than achene. Flowering from end of June to
August; fruiting August to September.

Marshy meadows, herb marshes, marshy and stony banks of rivers,
less often scrubby tundras and forest glades.—Arctic Europe; Euro-
pean part: Karelia-Lapland (Kola Peninsula, eastern Karelia up to Lake
Onega, Urals (along ridge as far as Southern Urals inclusively); Western
Siberia: Ob River Area (only northwestern). Endemic. Described from
Pae-Khoi Range on Kanin Peninsula. Type in Leningrad.

14. L. bucovinensis Nakai in Journ. Jap. Bot. XX (1944) 135
(sphalm. “bucoviensis”).—Cinenaria sibirica auct. non L. (1763): Bess.
Enum. pl. Volhyn. (1822) 33.—Senecio cacaliaefolius auct. non Sch.
Bip. (1845): Knapp. Pfl. Galiz. Bukow. (1872) 132 (sphalm.
“cacaliformis Sch. Bip.”), excl. synon.; Schmalh. Fl. IJ, 84 p. min.
p.—L. sibirica auct. non Cass. (1823): Ldb. Fl. Ross. II (1844-1846)
620, p. min. p. (quoad specim. volhyn.); Grecescu Consp. fl. Roman.
132; Prodan, Fl. Roman. I, 2, 1029; Katina in Vizn. Ros]. URSR, 352;
Szafer. Kulcz. Pawl. Rosl. Polsk. 698 p. p.? (saltem quoad aream
Wyz. Lub.).—L. ucrainica Minder. in Ukr. Bot. Zhurn. XIV (1957)
46.—Ic.: Grecescu, loc. cit.; Minder. op. cit. Fig. 2.

Perennial. Rhizome strongly shortened, covered with slender,
filiform, adventitious roots and above with thin fibrous remnants of
dead leaves. Stem erect, 30—75(100) cm high, 2.5—5.0 cm thick, sulcate,
purple in lower part or less often entirely, glabrous up to inflorescence
or with occasional, erect, brown, multicellular, thickish hairs, but below
inflorescence always densely covered with similar hairs and often also
with long, thin, white, arachnoid hairs. Leaves on both sides glabrous
or with occasional, thickish, short hairs beneath on prominent veins
and margin; leaves of sterile rosettes and basal leaves with long (2.5-
4 times as long as lamina), very thin, glabrous, often purple, petiole
and deltoid cordate, often almost sagittate lamina, with long and usually
acuminate, divergent lobes at base, mostly acuminate, not quite
uniformly sinuate-toothed with acute, almost serrate teeth, 4-9 cm long

820

780

and 5—10(14) cm wide; cauline leaves 2(3), with shorter petiole ex-
panded into long, inflated sheath, upper cauline leaf sometimes sessile;
upper 1-4 scale-like leaves sessile, amplexicaul, carinately folded like
lower bracteal leaves. Capitula (5)7—20, usually in simple, short, and
lax panicle; peduncles 3-8 mm long, only lowermost up to 3.0-4.5 cm
long, white-arachnoid-hairy, and with erect, brownish hairs, erect, later
drooping in upper part; bracteal leaves more or less violet- purple,
lower and sometimes almost all lanceolate, long-acuminate, sometimes
partially toothed, lower carinately folded and enveloping peduncle,
upper often almost linear, usually somewhat longer than involucre.
Involucre with 2—3 narrowly linear bracts at base, approximately as
long as involucre, narrowly campanulate; involucral bracts 7—9 dimor-
phic, in 2 indistinct whorls; inner bracts narrowly lanceolate, with very
narrow scarious border on both or one side; all bracts long-acuminate,
glabrous on back, apically floccose. Ligulate florets 6-8 with oblong,
13-17 mm long and 2.5-4.0 mm wide, 7—8-veined ligule; tube 5-7
mm long. Tubular florets about 20-24, with 9.0-10.5 mm-long co-
rolla; limb 3.0-3.5 mm long, oblong-campanulate, with 1.25-1.5 mm-
long narrow-deltoid teeth; tube 4-5 mm long;
anthers with long, narrowly lanceolate apical appendages and very
short basal appendages, antheropodia short; style branches narrow,
with somewhat enlarged appendages, capitate or acute, on outside up
to base covered with very short, thickish hairs. Achenes (immature) 4—
5 mm long, narrow, almost terete; pappus brownish, slightly longer
than achene. Flowering July to August.

Marshy meadows, mossy marshes, wet forests of Alnus
glutinosa.[black, common or European alder]. European part: Upper
Dnieper and Upper Dniester (very rarely). General distribution: Central
Europe (Polish highland and Eastern Carpathians and, apparently,
Transylvania). Described from western Bukovina (Romania). Type in
Tokyo University.

Section 3. Stenostegia Pojark. sect. nova in Addenda, XXV, 892.—
Bracteal leaves at base of peduncles narrow, linear or subulate, mostly
shorter than lower peduncles; pappus of white, soft hairs; stigma surface
occupying whole of inside of style branches or excluding narrow middle
groove; antheropodia broader than filament; leaves green, mostly soft,
not fleshy.

Type of section: L. narynensis (Winkl.) O. and B. Fedtsch.

Species mainly of Soviet Central Asia and Central Asia.

Subsection 1. Oligocephalae Pojark. subsect. nova 1. c. 892.—Capitula
large or medium, solitary or up to 10, in racemose inflorescence;
involucre goblet-shaped; rhizome densely tomentose.

82

_

781

Type of subsection: L. narynensis (Winkl.) O. and B. Fedtsch.
This subsection includes Soviet Central Asian species that form a
very natural group, which may be considered as a series.

15. L. robusta (Ldb.) DC. Prodr. VI (1837) 316; Ldb. Fl. Ross.
II, 621; Kryl. Fl. Zap. Sib. XI, 2863.—Cineraria robusta Ldb. FI. Alt.
IV (1833) 106.—Senecio robustus Sch. Bip. in Flora, XXVIII (1845)
50.—S. robustus &. typica auct. non Trautv. (1866): Franch. in Bull.
Soc. bot. Fr. XXXIX (1892) 300 p. p. (quoad pl. alt.).—Ic.: Ldb. Ic.
Fl. Ross. Alt. IV, tab. 354.

Perennial. Rhizome short, densely covered with numerous filiform
adventitious roots; stem solitary, simple, 25-80 cm high, basally up to
7 mm thick, strong, erect, ribbed, densely covered with fibrous remnants
of dead petioles and thick layer of brownish tomentum at base, initially,
like leaves, sparsely floccose or tomentose above. Leaves glabrescent,
lower cauline leaves with long (almost 1.5, less often almost about 2
times as long as lamina) petiole, somewhat enlarged toward base, lamina
ovate-cordate, wide, less often oblong, acuminate, 3-13 cm long and
3-10 cm wide; middle cauline leaves smaller than lower ones and
narrower, ovate or oblong-ovate, usually with shallow notched base,
long-acuminate, petioles usually as long as lamina; margins of lower
and middle leaves toothed with coarse, sharp teeth; upper leaves strongly
reduced, sessile, narrow, rhombic-lanceolate or lanceolate, undivided
or partly toothed. Capitula more or less drooping, large, including
ligulate florets (5) 6-7.5 cm in dia, solitary or 2-5, borne singly at
apices of long, 3 cm (upper ones), 8-16 cm-long, upright, very (lower
ones on 5-9 cm) remote thick peduncles. Involucre hemispherical,
with 3—4 very narrow bracts at base, green or less often slightly
anthocyanin-colored, initially arachnoid-hairy, later glabrous; involucral
bracts (10)15-17 mm long, inner bracts broadly elliptical, with wide
scarious margin; outer bracts narrower, oblong-elliptical or lanceolate,
with or without very narrow membranous border. Ligulate florets 8—
15, linear and oblong-elliptical to (less often) narrow-obovate, with
ligule (2)3 cm long and (3.5)5—8(13) mm wide. Corolla of tubular
florets about 9 mm long; tube a half as long as limb, limb lobed
approximately up to a fourth in narrowly deltoid acute teeth; basal
appendage of anthers short; style branches subacute or obtuse, with
hairs on outer side only near apex or up to middle, usually subglabrous
in ligulate florets. Achenes terete, somewhat flat, 6.5—7.0 mm long and
about 1.5 mm wide, light brown, ribbed-sulcate; pappus snow-white,
as long as corolla of tubular florets. Flowering June to July; fruiting
from August.

822

782

Stony and rubbly slopes of steppe.—Altai (western part). Described
from material from western Altai from Kerlyk and Kan rivers. Type in
London.

16. L. Kareliniana Stschegl. in Bull. Soc. Nat. Mosc. XXVII, 1
(1854) 176.—L. robusta var. kareliniana O. and B. Fedtsch. Perech.
Rast. Turk. III (1909) 212 p. p.

Perennial. Rhizome strongly shortened, densely covered with
slender, filiform adventitious roots and, like base of stem, with brownish
tomentum and stiff fibrous remnants of old dead petioles. Stem solitary,
35—46 cm high, basally 3.5—4.0 m thick, erect, ribbed-sulcate, as also
petiole, floccose-tomentose and more or less purple. Leaves grayish-
tomentose-floccose beneath; subglabrous above; basal and lower cauline
leaves on thin, long, semi-amplexicual petioles, expanded toward base,
(1.5)2—4(8) times as long as lamina; lamina of lower cauline leaves
4.5-7.0 cm long and as much wide (basal leaves smaller), cordate or
roundish-cordate, obtuse, less often some mucronate, sinuate-toothed
with small and often remote teeth or sinuate; middle cauline leaves 2—
3, with shorter petiole, as long as lamina or a half as long, narrower,
oblong-ovate and rhombic-ovate to ovate-lanceolate, upper leaves
sessile, rhombic-oblong to lanceolate and (uppermost leaves) linear,
with sharp, small, remote teeth to entire. Capitula 5.0-6.5 cm in dia
(including ligulate florets), less often solitary, usually 3—4(6), solitary,
on short, slender, more or less drooping peduncles 1.0—2.5 cm long,
narrowed at apices of stem, lower capitula sometimes slightly remote
and with longer, peduncles up to 3.5 cm. Involucre hemispherical,
1.5—2.0 cm in dia, up to 2.5 cm in upper capitulum, sparsely arachnoid-
hairy, with 2-4 bracts at base; involucral bracts 8-12, oblong-elliptical;
inner bracts much broader (sometimes broadly elliptical), with wide
membranous, usually purple margin; outer bracts with narrow
membranous margin. Ligulate florets 12-15, with 1.6-—2.5 cm-long and
2.5-6.0(9.0) mm-wide, narrowly-linear to oblong-elliptical ligule,
tubular florets (5.5)6.0-6.5 mm long; tube 1/2—3/7(1/3) as long as
lamina, incised up to a fourth (a fifth) length into acute, narrow deltoid
teeth; basal appendage of anthers short; style branches apically
narrowed, obtuse or subacute with hairs on outer side only below
appendage or slightly lower, glabrous or with occasional hairs on outer
side in ligulate florets. Achenes immature, about 6 mm long, ribbed-
sulcate. Flowering first half of June to first half of August.

Meadows and stony slopes of subalpine zone.—Soviet Central Asia:
Dzhungaria-Tarbagatai (Saur and Tarbagatai mountains). Endemic.
Described from Tarbagatai. Type (if not lost) in Moscow.

823

783

Note. In the description of this, presently forgotten species,
Stschegleev quite correctly noted its main differences from the Altaian
L. robusta (Ldb.) DC.: smaller capitula, with narrower, usually purple
involucres, on short peduncles aggregated at the apices of the stems;
lower and middle cauline leaves broader, obtuse or only mucronate,
their petioles longer than lamina; besides the leaves have smaller teeth
and throughout remain grayish-tomentose beneath. In its morphological
characteristics, this narrowly endemic species occupies an intermediate
position between L. robusta (Ldb.) DC. and L. narynensis Winkl.

17. L. narynensis (Winkl.) O. and B. Fedtsch. Perech. Rast. Turk.
III (1909) 212.—Senecio robustus var. typica Trautv. in Bull. Soc.
Nat. Mosc. XXXIX, 2 (1866) 362; Franch. in Bull. Soc. Nat. Fr.
XXXIX, 300 p. p. (quoad pl. alatav.).—Senecio robustus var.
kareliniana Trautv. loc. cit.; Franch. loc. cit—Senecio narynensis
Winkl. in Tr. SPb. Bot. Sada, XI (1890) 319; Franch. loc. cit.—L.
robusta var. typica O. and B. Fedtsch. loc. cit.—L. robusta var.
kareliniana O. and B. Fedtsch. loc. cit. p. p.—L. robusta auct. fl.
turkest. non DC. (1837).—L. robusta f. turkestanica Krasch. in sched.

Perennial. Rhizome short, covered with slender, filiform,
adventitious roots and together with stem base, densely covered with
lax brownish tomentum and stiff fibrous remnants of dead petioles.
Stem solitary or less often 2-4, 30-70 cm high, at higher altitudes
sometimes 14-17 cm high, usually somewhat flexuous, soft, sulcate,
floccose-tomentose, sometimes subglabrous below apex. Leaves
glabrous above or only partially, mainly in middle, discontinuously
floccose-tomentose, grayish beneath up to end of vegetative growth,
usually densely arachnoid-hairy; leaves in basal rosette and lower
cauline leaves crowded, roundish and roundish-cordate to ovate-cordate,
(1.7)3-12 cm long and up to 10 cm wide, obtuse or mucronate usually
with deeply cordate, less often almost hastate or truncate base, with
petiole as long as lamina or 1.5—2 times as long; middle cauline leaves
usually remote, lower leaves 1(2) among them, like upper leaf but with
shorter petiole, others transitional to upper leaf, narrower, oblong-
elliptical to lanceolate, short-petiolate, with small remote teeth, upper
cauline and bracteal leaves small, narrow, lanceolate, entire, acuminate,
sessile. Capitula aggregated at stem apex, usually 4-10(14), in racemes
to almost corymbose inflorescence, elongated in lower part, sometimes
with branched peduncles; upper 3-4 capitula usually on short (1-4 cm
long) peduncles, more or less crowded, lower ones remote on 3-—8(11)
cm and longer peduncles up to 8(10) cm, sometimes bearing 2—-3(4)
capitula, in short, particularly high-altitude specimens, capitula in all
1-2. Capitula drooping, including ligulate florets capitula at apex

784

(4.5)5-6 cm in dia. Involucre campanulate, 1-2 cm in dia, usually
arachnoid-tomentose, particularly at base, with 1-2 lanceolate bracts
at base; involucral bracts (8)10—15 in upper capitula and 8-10 in others,
green or purple; inner bracts narrowly elliptical to broadly elliptical,
with wide membranous margin, upper ones narrowly elliptical, with
scarcely membranous margin; all bracts abruptly narrowed into mucro,
short-ciliate and covered with lax, tomentum. Ligulate florets 8-14(17),
with linear-oblong to elliptical ligule, 1.4-2.5 cm long and 2-7 mm
wide, sometimes filiform staminodes also present; style branches hairy
outside only near apex or slightly below. Tubular florets 7.5-9.0 mm
long; corolla tube 1/2-—2/3 as long as limb lobed up to (1/5)1/4—1/3
into oblong- or narrow-deltoid teeth; appendage of style branches obtuse
or subacute, hairy outside, sometimes up to middle; anthers with ovate-
lanceolate, acute apical appendage and very short deltoid basal
appendage. Achenes 5-6 mm long and 1.0-1.5 mm wide, narrowed
toward base, ribbed-sulcate. Flowering June to July; fruiting end of
August to September. (Plate XXXVIII, Fig. 1).

Mountains from forest to alpine zone (up to 3,600 m), forest,
subalpine and alpine meadows, mountain steppes, open stony and rocky
slopes, talus and gravel beds.—Soviet Central Asia: Dhungaria-
Tarbagatai (only Dzhungarian Alatau), Tien Shan (entire Eastern and
Central Tien Shan, Trans-Ili Range, eastern part of Kirgiz Range,
western slope of Fergana Range), Pamiro-Alai Region (Alai and Trans-
Alai ranges). General distribution: Dzhungaria-Kashgaria (southwestern
Dzhungaria and western Kashgaria). Described from Central Tien Shan
(Kirgizia), from Naryn River. Type in Leningrad.

18. L. karataviensis (Lipsch.) Pojark. comb. nova.—Senecio
karataviensis Lipsch. in Fedde, Repert. sp. nov. XLII (1937) 160.

Perennial. Rhizome covered with filiform adventitious roots and
with fibrous remnants of dead leaves at root collar, densely covered
with brownish woolly tomentum. Stem erect, 15-35 cm high, sulcate,
densely pubescent below, sparsely above. Leaves thick, almost
coriaceous; basal and lower cauline leaves with long (4-14 cm) petiole
2.5~3 times as long as lamina, densely arachnoid-tomentose with erect
hairs, their lamina reniform or deltoid-reniform, basally cuneate, (2)3-
6 cm long and 2.5(4—5) cm wide, unevenly toothed with coarse, obtuse
or acute teeth; middle cauline leaves 3—4, gradually reduced upward,
deltoid-ovate, upper leaves among them lanceolate, sinuate-toothed or
entire, short-petiolate or sessile; upper cauline leaves small, linear,
mostly entire. Capitula including ligulate florets 1.5—2.0(2.5) cm in
dia, 3-7, aggregated at stem apex in racemose or somewhat corymbose
inflorescence. Involucre goblet-shaped, with few, lanceolate, small, 2—

825

785

4 mm long bracts at base; involucral bracts 8-11 mm long, dimorphic:
inner bracts oblong-obovate, rather short-acuminate, with wide scarious
margin; outer bracts narrowly lanceolate, long-acuminate, on both or
only one side with narrow scarious fringe; all bracts glabrous outside,
in upper part along margin rather short-ciliate and floccose at tip.
Ligulate florets about 10, with yellow, oblong-elliptical, 10-14 mm
long and 3-4 mm wide ligule with (7)9 veins; tube 5 mm long. Tubular
florets about 30, with narrow-campanulate almost tubular, 5 mm long
limb; tube 2 mm long; stamens with very short basal appendages of
anthers and weakly enlarged antheropodia; style branches short, with
conical, short, acute appendage, with a tuft of long hairs outside below
appendage and occasional short hairs, almost up to bifurcation of style.
Achenes brownish, 5-6 mm long, ribbed; pappus white. Flowering
second half of May to June.

Gravel beds and stone fields.—Soviet Central Asia: Tien Shan
(western part of Karatau Mountains and also, according to personal
communication of O.V. Czerneva, Akatau and Akbashtau mountains,
Chatkal and Talass ranges). Endemic. Described from Kazakhstan from
Bayaldyr Gorge and Karatau Range. Type in herbarium of Moscow
State University.

19. L. schischkinii Rubtz. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, VII (1938) 138.—Senecio schischkinii Lipsch. in sched.

Perennial. Plant short with small leaves; rhizome short, covered
like base of stem with brownish, soft tomentum and densely covered
with filiform adventitious roots. Stem solitary, simple, 6-14 cm high,
slender, at base 1.5—2(3) mm thick, soft, mostly bent, as also petiole,
entirely but later, only in upper part, white-arachnoid-floccose-
tomentose basally covered with fibrous remnants of dead petioles.
Leaves sparsely floccose to glabrous above, grayish-arachnoid-hairy
beneath, later usually glabrous, often purple, particularly along veins;
leaves of sterile rosette and basal leaves 1.5-3.5(4.5) cm long and as
wide, almost round or broadly cordate, rarely almost reniform, entire
or sinuate-toothed with small remote teeth, with petiole 1.5—2 times as
long as lamina, but sometimes as long or, in lowermost leaves up to
4—6(8) times as long as lamina; middle cauline leaves 1-2, sometimes
like lower leaves but more often like upper leaves, i.e., sessile or
short-petiolate and with small, lanceolate, acuminate, entire lamina.
Capitula drooping, solitary at apex of stem (very rarely second
capitulum developing nearby on very short, 0.7 mm-long peduncle),
including ligulate florets 3.5-4.0 cm in dia. Involucre broadly
campanulate, purple, 10-15 mm in dia, with 1-4 short, narrow subulate
bracts at base; involucral bracts 10-12; inner bracts oblong to quite

826

786

broadly elliptical, with wide scarious margin; outer bracts oblong,
almost lacking scarious margin; all bracts arachnoid-hairy at least at
base and on acuminate tip. Ligulate florets 10-14, with oblong, less
often rather broadly elliptical, ligule; style branches subglabrous outside,
obtuse. Corolla of tubular florets 8.5-10.5 mm long; tube about 2/3 as
long as limb; limb lobed up to (1/4) 1/5 in narrowly deltoid teeth; style
branches with short-pointed, conical tip, rarely obtuse, with tuft of
hairs only outside, subterminal or almost up to middle of branch.
Achenes (immature) 6.0—-6.5 mm long, narrowly terete, ribbed-sulcate.
Flowering July.

Alpine zone on stony mountain slopes.—Soviet Central Asia:
Dzhungaria-Tarbagatai (Saur, Tarbagatai and Dzhungarian Alatau
Mountains), Tien Shan (only northeastern part, Ketmen Mountains).
Endemic. Described from Tarbagatai. Lectotype and its duplicate in
Leningrad.

Note. L. schischkinii is an alpine species very similar to L.
kareliniana, from which it is distinguished by, besides low stature,
small, roundish, rather obtuse, lower cauline leaves that are usually
glabrous at anthesis; smaller, usually solitary capitula with narrower
involucres; and also a tubular floret corolla with relatively longer tube
and less deeply lobed limb.

Subsection 2. Racemiferae Poljak. subsect. nova in Addenda,
XXV, 893.—Capitula rather large (4.0-4.5 cm in dia), aggregated in
large number into long oblong raceme; basal leaves reniform-deltoid-
ovate; rhizome glabrous (lacking pubescence).

There are two very closely related species in Korea, one of which
occurs in Ussuri Territory.

Type of subsection: L. jaluensis Kom.

20. L. jaluensis Kom. in Tr. SPb. Bot. Sada, XVIII (1901) 420;
Fl. Manchzh. III, 695; Nakai, Fl. Korean. II, 38; Kom. and Alis. Opred.
Rast. Dalnevost. Kr. II, 1053.—Senecillis jaluensis Kitam. in Acta
Phyt. et Geobot. VIII (1932) 82.—L. pulchra Nakai in Bot. Mag.
Tokyo, XXXI (1917) 126; Bull. Nat. Sc. Mus. XXXI, 117.

Perennial. Rhizome short, covered with numerous, strong filiform
adventitious roots, in upper part with scaly blackish remnants of dead
petioles. Stem 65.200,cm high, 6-10 mm thick at base, erect, sulcate,
usually densely covered with brown, flexuous (glandular?) hairs and
white-arachnoid-hairy mainly below inflorescence. Leaves in dry
condition, rather thick, bright green and glabrous above, lighter and
finely tomentose or glabrous beneath; basal leaves usually 2, 18-37
cm long and 17—35(40) cm wide, reniform-deltoid or deltoid-ovate,

827

787

broadly or deeply cordate or almost hastate at base, obtuse or subacute,
with long winged petiole, 1.5(2.5) times as long as lamina, often
expanded, up to 6 cm wide, like stem entirely hairy or only in upper
part, or only near sheath-like expanded base, uniformly toothed with
sharp teeth; cauline leaves also 2, much smaller and narrower, 10-22
cm long, 10.5 cm wide, cordate-hastate to lanceolate, on short, broadly
winged petiole or sessile, amplexicaul; bracteal leaves strongly reduced,
narrowly lanceolate or linear, entire. Inflorescence 18-50 cm long, as
narrow, long, simple raceme or with lateral branches in lower part
bearing racemes of more or less 5 capitula; peduncles of capitula 7—
50 mm long, like stem usually densely covered with dark, reddish,
short hairs and sometimes with occasional white-arachnoid hairs.
Capitula including ligulate florets 4.0-4.5 cm in dia. Involucre
campanulate, with truncate base, 11-12(13) mm long and up to 15 mm
in dia, with 1-3 small, almost filiform bracts at base or lacking them;
involucral bracts 8—10(12); inner bracts oblong elliptical or lanceolate,
with rather broad scarious margin covering narrowly lanceolate outer
bracts with narrow scarious margin; all bracts glabrous on back,
acuminate. Ligulate florets 5-8, with 16-20 mm long and 4-6 mm
wide ligule, with 6-8 veins, 3.5—4 times as long as tube; tubular florets
numerous (30 and more), with 8.0—9.5 mm-long corolla; limb tubular-
campanulate, tube 2.5-3.0 mm long; anthers with very short, acute,
basal appendages, filaments with narrow flat antheropodia; style
branches long, twisted, obtuse, covered on outside from middle with
long, flat hairs, particularly dense below appendage. Achenes S—6 mm
long, narrow, ribbed, sordid stramineous; pappus snow-white, slightly
longer than achene. Flowering August to first half of September; fruit-
ing from end of September.

Wet, mainly valley meadows, in forest zone among tall herbs.—
Far East: Ussuri (southernmost area). General distribution: China
(northeast, near border with Korea), Korea. Described from Mukden
Region (northeast) from Yalu River valley above Sesanpo. Type in
Leningrad.

Note. L. deltoidea Nakai (loc. cit. 1917), is a closely related spe-
cies not known to me. Kitamura (I. c.) considers it identical with L.
jaluensis, along with L. pulchra Nakai (loc. cit. 1917). The author of
this species himself considered the latter species to be identical with
L. jaluensis in his more recent work (Nakai, 1952), where, however,
he retained the separate status of L. deltoidea.

Subsection 3. Paniculatae Pojark. subsect. nova in Addenda, XXV,
893.—Capitula small, mostly in compound paniculate inflorescence,
rhizome covered with dense tomentum.

828

788

Series 1. Thyrsoideae Pojark. 1. c. 893.—Involucre campanulate
or goblet-shaped; lateral branches of panicles usually much shorter
than main axis; capitula in racemes; leaves oblong-ovate, with cordate
or sagittate base.

A monotypic series.

21. L. thyrsoidea (Ldb.) DC. Prodr. VI (1837) 315; Kar. and Kir.
in Bull. Soc. Nat. Mosc. XIV, 3 (1841) 444; Ibid. XV, 2 (1842) 385;
Pojarkova in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XII, 298.—
Senecio sibiricus L. fil. Suppl. (1791) 370; Lepech. in Nov. Acta
Petropol. XI, 400; Ldb. Fl. Ross. II, 638; Trautv. in Bull. Soc. Nat.
Mosc. XXXII, 2, 58; Franch. in Bull. Soc. bot. Fr. XXXIX, 302; O.
and B. Fedtsch. Perech. Rast. Turk. III, 219; Kryl. Fl. Zap. Sib. XI,
2847.—Cineraria thyrsoidea Ldb. Ic. pl. Fl. Ross. II (1830) 18; FI.
Alt. IV, 107.—Ic.: Lepech. op. cit. tab. 9; Ldb. Ic. pl. Fl. Ross. II, tab.
157.—Exs.: Fl. alt. exc. No. 93 (sub Senecio sibiricus Lepech.).

Perennial. Rhizome short, with numerous, rather slender, filiform
adventitious roots, densely covered, like stem base, with brownish
tomentum and stiff fibrous remnants of dead leaves. Stem 40-125 cm
high, 3-10 mm thick in lower part, simple, straight or usually flexuous
above, green, sulcate, glabrous or more or less arachnoid-tomentose
when young, particularly in upper part. Leaves somewhat thick,
concolorous on both sides, glabrous from beginning or initially beneath,
uniformly arachnoid-tomentose, usually glabrescent, more or less
persistent as scattered floccose tomentum; basal and lower cauline
leaves oblong-ovate, less often deltoid, with deeply cordate or almost
hastate at base, gradually attenuate upward, acute or sometimes obtuse,
sinuate-toothed, with teeth of different forms, from broadly deltoid to
lanceolate, long-petiolate, petiole 1.5—2 times as long as lamina; middle
cauline leaves with shorter petiole, approximately as long as lamina,
usually lanceolate, long-acuminate, with a shallow notch or truncate at
base; upper cauline leaves strongly reduced in size, lanceolate-linear
to narrowly linear, mostly entire, short petiolate, upper-most leaves
sessile. Capitula including ligulate florets 2.5-3.5 mm in dia, at stem
apex, (10)12-—60, borne in narrow, rather dense panicle, with branches
usually much shorter than main axis, or less often, almost as long,
bearing 1—4(5); capitula; peduncles (5)7—-45 mm long. Involucre
campanulate or goblet-shaped, 7-12 mm long and 7-14 mm in dia,
with or without 1-2 small bracts at base; involucral bracts, as also
peduncles, often floccose-tomentose, later glabrous, only at apex
tomentose-floccose; inner bracts ovate or broadly elliptical, with wide
scarious border; outer bracts a half as wide as inner, oblong-elliptical,
with narrow scarious margin on both or one side. Ligulate florets

789

Plate XXXIX.
1 — Ligularia altaica DC.; 2—L. glauca (L.) O. Hoffm.

829

831

790

5—10(13), with elliptical or oblong,.sometimes linear, ligule, 8-14 mm
long and (2.5)3-5(6) mm wide; staminodes lacking; style branches
long, acute, with short, sparse papilliform hairs on outer side. Tubular
florets numerous, with campanulate-tubular limb, 1.5—2.5 times as long
as tube and with oblong-deltoid teeth; style branches hairy outside
almost from base, and with dense tuft below acute tip. Achenes
brownish-yellow, oblong; pappus snow-white, as long as achene.
Flowering end of June to July; fruiting from end of August.

Wet floodplain and saline meadows, river valleys in shrub thickets,
along edges of marshes.—Western Siberia: Irtysh, Altai; Soviet Central
Asia: Lake Balkhash Region, Dzhungaria-Tarbagatai. Endemic.
Described from Siberia (from Altai?). Type in London?

Series 2. Songaricae Pojark. op. cit. 893.—Involucre narrow,
cylindrical or obconical, green; general inflorescence compound panicle
or simple raceme; leaves oblong-deltoid, acute, with cordate base. Two
Soviet Central Asian species are included in this series.

22. L. songarica (Fisch.) Ling in Contrib. Bot. Inst. Nat. Akad.
Peip. II (1934) 532 (excel. synon.: Senecio bungei Fr. and L. thyrsoidea
var. alpina Bge.); Pojarkova in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XII (1950) 297.—Senecio songaricus Fisch. in Fisch. and Mey.
Enum. pl. Nov. Schrenk. (1841) 52; Ldb. Fl Ross. II, 637
(“songoricus”); O. and B. Fedtsch. Perech. Rast. Turk. III, 218 (excl.
syn. 1, 2, 4 and var. arachnoideus;, Franch. in Bull. Soc. bot. Fr.
XXXIX, 304, p. min. p.; Kryl. Fl. Zap. Sib. XI, 2848.—S. turkestanicus
Winkl. in Tr. SPb. Bot. Sada, XI (1890) 279.—S. songaricus var.
hastifolius Winkl. in sched.; O. and B. Fedtsch. op. cit. 219, nom.
nud.—Ic.: Pojarkova, loc. cit. Fig. 2.

Perennial. Rhizome short, densely covered with thick, adventitious
roots, in upper part tomentose. Stem 40-80 cm high, straight, in upper
part sometimes flexuous, sulcate, densely arachnoid-tomentose, later
glabrous or floccose-tomentose and below, often reddish. Leaves when
young, as also petiole, arachnoid-to mentose, later glabrous; basal and
lower cauline leaves with long petiole, at base expanded like sheath,
1.5-2 times as long as lamina, lamina oblong deltoid or oblong-ovate
to lanceolate, less often broadly deltoid, with deeply notched, more or
less hastate base, acute or less often obtuse, 7-35 cm long and 3.5—25
cm wide, sinuate, irregularly sharp-toothed; middle leaves smaller, with
shorter petiole, upper ones strongly reduced, from broadly angular
with base truncate to narrowly lanceolate, gradually narrowed into
short petiole, partly sessile, sometimes with amplexicaul auricle, entire
or with more or less remote teeth, uppermost leaves sometimes narrowly

832

791

linear, identical to bracteal leaves. Capitula upright, (30) 60-300, in
paniculate inflorescence, (6)10-40 mm long and 5-25 mm in dia,
pyramidal, elongated, with short lateral branches or more or less
corymbose if upper branches reach level of main axis; capitula on
inflorescence branches in panicle or simple racemes only in their upper
part; peduncles of capitula 2—-10(17) mm long. Involucre narrow,
tubular-cylindrical, unchanged in shape at fruiting, 7.5-9.0 mm long
and 4—5 mm wide in dia with 1-3 very small, appressed bracts at base;
involucral bracts 5—8, light green, pubescent on back; inner bracts
usually oblong-elliptical, with broadly scarious margin; all bracts acute
and apically short-ciliate along margin. Ligulate florets bright yellow,
2-4(5), with oblong, 7-10 cm long and 2.5—3.5 mm wide ligule, 2—2.5
times as long as tube, with 4—7 veins, lacking staminodes. Tubular
florets 7-12, with 7.5-10.0 mm long corolla; limb narrowly
campanulate, about 1.5 times as long as tube, lobed up to a fourth to
a third into narrow-deltoid teeth, style branches short, acute, outside
covered with somewhat long hairs up to base, longer below apex and
in a tuft. Achenes pale yellow, oblong, 4.5-5.0 mm long, somewhat
flat; pappus as long as or up to 1.5 times as long as achene, snow-
white, very fine hairs, with very short barbs. Flowering second half of
June to first half of August; fruiting from second half of August.

Floodplain meadows, scrubs and tall herb thickets along banks of
rivers, lakes, and edges of marshes, thickets of cheegrass and reeds,
wet saline meadows, mainly in foothills and in steppe Zone.—Soviet
Central Asia: Lake Balkhash Region, Dzhungaria-Tarbagatai, Tien Shan
(only eastern part of Kirgiz Range). General distribution: Dzhungaria-
Kashgaria (Dzhungaria). Described from Tentek River, in Dzhungarian
Alatau foothills. Lectotype and its duplicates in Leningrad.

23. L. knorringiana Pojark. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XXI (1961) 360, Fig. 1.

Perennial. Rhizome strongly shortened, densely covered with cord-
like, thickish, adventitious roots, its upper part, as also stem base,
densely covered with slender fibrous remnants of dead petioles and
sparse brown tomentum. Stem solitary, erect, 14-40 cm high, slender;
about 1.5 mm (less often more) thick in lower part, longitudinally
ribbed, rather densely arachnoid-floccose only below inflorescence,
but sparsely and irregularly below; often more or less with anthocyanin
coloration. Leaves at flowering glabrous on both sides, finely coriaceous
when dry, dull, light green, paler beneath, with thin lateral veins distinct
on both sides; lowermost leaves crowded in basal rosette and mostly
long-petiole, petioles 1.5—2 times as long as lamina, less often as long
or slightly shorter, thin, often reddish, sulcate, toward base expanded

833

792

into thick, veined, coriaceous sheath; lamina 3—8(10) cm long and
2.7-5.0 cm wide, cordate, oblong-cordate, or almost sagittate, short-
acuminate or more often long-acuminate, sinuate-small-toothed, only
near base coarsely toothed; middle cauline leaves 1-3, lower l—2 like
basal but with shorter petiole; upper cauline leaves 4—9, strongly
reduced, narrow, 3.5 cm long and 1.5-2.0 mm wide, oblong or often
narrowly linear, entire, finely long-acuminate, lowermost of them
petiolate, rest sessile. General inflorescence 3-10 cm long, with (1)3-
6(10) capitula, usually in more or less elongated, lax recame, less
often in weakly branched panicle; peduncles slender, arachnoid-hairy,
upper 0.8—2.5 cm long, with solitary capitulum, lower up to 5 cm long,
sometimes with 2-3 capitula. Capitula upright, including ligulate florets
1.5-2.5 cm in dia. Involucre narrowly campanulate to narrow-obconical,
7-10 mm long and as much in dia, with 1-2 very small setaceous
bracts at base; involucral bracts 7-8 (9), more or less densely floccose-
tomentose to subglabrous; inner bracts rather broadly elliptical or
oblong-obovate, with wide, transparent scarious border; outer bracts
narrowly linear or lanceolate-linear, lacking or with very narrow border;
all bracts acuminate, short-ciliate and tomentose at apex. Ligulate florets
5-7, with oblong, 11-13 mm-long and 3.0-4.5 mm-wide ligule, with
4-5 veins; tube a half as long as ligule; staminodes usually present.
Tubular florets 7-30, with 6.4-7.5 mm-long corolla; tube approximately
two-thirds as long as limb; limb narrowly campanulate, lobed up to a
third into oblong-deltoid teeth; style branches with a barb of long
thickish hairs outside below appendage, but densely covered with
thinner and shorter hairs below up to base; basal appendage of anthers
very short. Mature achenes unknown; pappus hairs snow-white, as long
as corolla of tubular florets. Flowering second half of July to August.

Mountains from forest to alpine zone, up to 3,600 m, in forests,
subalpine and alpine meadows, mountain-steppes, open stony and rocky
slopes, talus and gravel beds.—Soviet Central Asia: Tien Shan (eastern
and central parts, Ketmen and Terskei Alatau ranges). Described from
eastern Tien Shan from Terskei Alatau Range and in vicinity of the
village of Tekes in Tekes River valley. Endemic. Holotype and isotypes
in Leningrad.

Note. The closest species to L. knorringiana is L. songarica (Fisch.)
Ling. which, however, is quite sharply distinguished by a number of
characteristics: a narrower tubular-obconical (not campanulate-
obconical) involucre, with 5—6 (very rarely 7) involucral bracts and
not 7-10 bracts; the receptacle with smaller alveolae separated by
broad septae; in all, 2—4(5) and not 5-7, relatively short ligulate flo-
rets; a larger and more compound, many-headed inflorescence (with
up to 300 capitula); and also larger leaves and a taller stem.

834

793

Series 3. Thomsonianae Pojark.—Involucre narrowly cylindrical,
golden-brown; capitula in compound corymbose panicle, its lateral
branches bearing corymbose inflorescences at their ends; leaves
roundish-ovate or roundish-deltoid, soft, dark green above.

Besides L. thomsonii, L. trigonophylla Rech. f. from Afghanistan
also belongs to this series.

24. L. thomsonii (Clarke) Pojark. in Spisok. Rast. Gerb. Fl. SSSR,
XI (1949) 165 and in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR,
XII (1950) 293.—L. thyrsoidea var. alpina Bge. Rel. Lehm. in Mém.
sav. étr. VIII (1854) 347; Boiss. Fl. or. III, 383.—Senecio thomsonii
Clarke, Compos. indic. (1876) 205; Hook. f. Fl. Brit. Ind. III (1882)
348; Franch. in Ann. Sc. Nat. 6-me sér. XVI (1883) 313 “thompsoni’.—
S. bungei Franch. Ibid. 313.—S. songaricus auct. non Fisch. (1841):
Franch. in Bull. Soc. bot. Fr. 39 (1892) 304, p. max. p.: O. and B.
Fedtsch. Perech. Rast. Turk. III, 218 p. p. (excl. syn. S. turkestanicus,
S. songoricus var. hastifolius and partim area).—S. songaricus var.
arachnoideus Winkl. in sched.; O. and B. Fedtsch. ibid., 219, nom.
nud.—L. bungei M. Pop. in Tr. Uzb. Univ. Nov. Ser. 27, Biol. 14
(1941) 74.—Ic.: Pojarkova, loc. cit. Fig. 1.—Exs.: G. Fl SSSR, No.
3498.

Perennial. Rhizome short, with few approximate tiers of cord-like
adventitious roots, in upper part covered with thick layer of brownish
tomentum. Stem 30-200 cm high, up to 18 mm thick in lower part,
sulcate-ribbed, often reddish, in younger stage arachnoid-tomentose,
later glabrous or floccose-tomentose. Leaves somewhat thick, soft, dark
green and lustrous above, lighter beneath, initially entirely grayish,
arachnoid-tomentose, very often persisting till end, rarely entirely
glabrous; basal leaves mostly 2-3, petiolate, petiole 1.5—2.5 times as
long as lamina, basally enlarged into semiamplexicaul sheath, lamina
(5) 7-25 cm long and 6-35 cm wide, broadly ovate or roundish ovate
or deltoid to strongly expanded-reniform, with sagittate hastate, less
often deeply cordate base, with large, mostly roundish lobe at base and
with round, less often acute apex, sinuate-sharply toothed to finely
toothed, teeth resembling cusp, less often only sinuate or entire; lower
1-2 cauline leaves like basal but usually much smaller and with shorter
petiole, upper leaves strongly reduced, of same shape or strongly
narrowed, lanceolate, often toothed only in lower part, or all leaves
entire, short-petiolate. General inflorescence compound, corymbose
panicle, 11-35 cm long and 6-30 cm in dia, arachnoid-tomentose,
later often entirely glabrous; branches usually long, mostly bearing
compound corymbs at their tips, consisting of few simple, dense
corymbs at ends of branches of third order, containing a few capitula
crowded on short 2—10(15) mm-long peduncles. Involucre narrowly

835

794

cylindrical, 8-12 mm long and 3.5—4.0 mm in dia, with 1-3 appressed,
narrowly lanceolate, long-acuminate, 3—5 mm long bracts; similar one
or few bracts usually also found on peduncles; involucral bracts soon
turning brown, lustrous, glabrous, with a tuft of somewhat long hairs
only at apex and white-ciliate along margin in upper part; inner bracts
oblong- or lanceolate-elliptical, with wide scarious margin; outer bracts
two-thirds as wide to almost linear, with very narrow membranous
margin; all bracts acuminate; at maturity of fruits bracts stellately
squarrose. Ligulate florets 1-3 very rarely 4(5), bright yellow; ligule
oblong, less often oblong-elliptical, acute, 12-20 mm long and 3-4
mm wide, with (4)6—8 veins, 3—4 times as long as tube. Tubular florets
7-12, with 7.5-8.0 mm-long corolla; limb narrowly campanulate or
almost cylindrical-campanulate, about 1.5 times as long as tube and
lobed up to 1/4—2/5 into narrowly deltoid teeth; style branches acute,
with dense tuft of hairs outside below apex, and with very short
papilliform hairs up to slightly below middle. Achenes 5—6 mm long,
narrowly terete, light yellow; pappus snow-white, as long as achene.
Flowering first half of June to July; fruiting from end of August.

Upper part of arboreal-scrub vegetation zone in forests, particularly
in walnut and maple forests, and in tall herb thickets and also in
meadows in subalpine zone where, in places, it is the landscape plant.—
Soviet Central Asia: Dzhungaria-Tarbagatai (Dzhungarian, Alatau),
Pamiro-Alai Region (except eastern and western Pamir) and whole of
Tien Shan. General distribution: Dzhungaria-Kashgaria (Kashgaria).
Described from Kashgaria. Type in Kew.

Series 4. Altissimae Pojark.— Involucre tubular-campanulate;
capitula aggregated in pyramidal panicle, panicle branches bear capitula
at their tips in racemes or small panicles; leaves oblong-ovate, light
green, fleshy and thick.

One species.

25. L. altissima Pojark. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XII (1950) 316.—Ic.: Pojarkova, ibid. Fig. 6.

Perennial. Rhizome short, covered with numerous filiform
adventitious roots and with fibrous remnants of dead leaves in upper
part. Stem about 2 m high and up to 1.5 cm thick, erect, sulcate-
ribbed, glabrous or sparsely floccose-arachnoid-hairy only below.
Leaves light green, fleshy, lustrous, glabrous, but initially weakly
arachnoid-hairy beneath; basal leaves 2 with thick, up to 0.8 cm thick,
not winged petiole expanded into long sheath, their laminas as long as
petiole or slightly shorter, 25-39 cm long and 25-30 cm wide, cordate
or ovate, with more or less broadly cordate base, somewhat decurrent

836

837

795

on petiole, obtuse, sinuate-toothed, teeth narrowed into short cusp;
cauline leaves 3, reducing towards stem apex, lower among them with
ovate-deltoid lamina and narrowly winged petiole as long or up to a
half as lamina; next leaf with short, broadly winged petiole, third leaf
small, approximately as long as lower bracteal leaf, sessile, entire.
Inflorescence compound pyramidal panicle up to 100 cm long, with
numerous lateral branches having bracteal leaves at base, like upper
cauline leaf in lower branches but smaller, upper branches with narrow,
lanceolate-linear or linear bracteal leaves, bearing capitula in dense
pyramidal, elliptical, or capitate panicles only in upper part; lower
branches 20-45 cm long, widely spaced, generally reduced toward
apex of inflorescence, coming closer to each other. Capitula erect,
after anthesis drooping, on peduncles shorter than involucre. Involucre
tubular-campanulate, 9-10 (11) mm long and as much in dia with 1-
3 short linear bracts at base; involucral bracts 5-8, dimorphic: inner
bracts ovate or obovate, with broadly scarious margin; outer bracts
narrow, lanceolate or almost linear, acuminate and white tomentose-
ciliate at apex. Ligulate florets (2)3—4, with narrowly obovate, about
12 mm-long and 4.5-5.0 mm-wide ligule, with 5—7 veins; staminodes
absent. Tubular florets 10-14, with 9-10 mm-long, corolla; limb
narrowly campanulate, 5.5-6.5 cm long; anthers with very short basal
appendages, antheropodia weakly enlarged; style branches conical,
acute, with dense tuft of somewhat long hairs outside below apex and
shorter hairs up to middle of branches or slightly below; in ligulate
florets stylodes only with tuft of hairs below apex. Immature achenes
brownish, ribbed, 5 mm long; pappus white, slightly shorter than corolla
of disk florets. Flowering second half of June.

Upper part of tree-scrub zone, on slopes among tall herbs and also
in subalpine zone.—Soviet Central Asia: Pamiro-Alai Region (southern
slope of Hissar Range, where it is known only from basin of Kondara
River, tributary of Varzob River). Endemic. Described from Kvak in
basin of Kondara River. Type in Leningrad.

Section 4. Senecillis (Gaertn.) Kitam. Compos. Japon. III (1942)
187.—Senecillis Gaertn. De fruct. et Sem. II (1791) 453.—Sect.
Cyathocephalum (Nakai) Kitam. op. cit. 185 p. p.; Addenda XXV,
893.—Capitula mostly small, less often medium-sized, in simple raceme
or narrow, oblong, compound panicle. Involucre tubular or almost bowIl-
shaped; involucral bracts free or connate in 3-or 4-toothed bowl; pappus
hairs white, soft, long or short, stiff or brown setaceous; antheropodia
distinctly broader than filament. Style branches with conical appendage.
Leaves glaucescent, fleshy, succulent or more or less coriaceous.

Type of section: L. glauca (L.) O. Hoffm.

83

oo

796

Species of eastern Soviet Central Asia and Central Asia, Iran,
Siberia and Europe.

Subsection 1. Sericochaetae Pojark. subsect. nova in Addenda,
XXV, 894.—Pappus hairs snow-white, soft, long; involucral bracts
free; stem hollow; rhizome glabrous.

Type of subsection: L. macrophylla (Ldb.) DC.

Series 1. Macrophyllae Pojark. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XII (1950) 310.—Capitula small, numerous, few-flowered,
in oblong-cylindrical or cylindrical-pyramidal panicles; ligulate florets
(1)2-6; involucre narrow; involucral bracts 5-8.

This series includes three [sic] Central Asian species. L. persica
Boiss. (Elburz Range in northern Iran) and L. afghanica Pojark.
(Afghanistan).

26. L. macrophylla (Ldb.) DC. Prodr. VI (1837) 316; Kar. and
Kir. in Bull. Soc. Nat. Mosc. XIV, 3 (1841) 444; Ibid. XV, 2 (1842)
461; Ldb. Fl. Ross. III, 621; O. and B. Fedtsch. Perech. Rast. Turk.
III, 212 (excl. area Tien-schan and Pam.-Al.); Krylov, Fl. Zap. Sib.
XI, 2863; Pojarkova in Bot. Mat. Gerb. Bot. Inst. AN SSSR, XII,
310.—Cineraria macrophylla Ldb. Fl. Alt. IV (1833) 108.—Senecio
ledebourii Sch. Bip. in Flora, XXVIII (1845) 50; Franch. in Bull. Soc.
bot. Fr. 39 (1892) 302 p. p. typ.—Ic.: Ldb. Ic. pl. Ross. Alt. IV (1833)
tab. 367.

Perennial. Rhizome short, densely covered with thick, cord-like,
brown or yellowish-brown adventitious roots, and in upper part with
stiff fibrous remnants of dead leaves. Stem (35)100—180 cm high, up
to 1.0-1.5 cm thick in lower part, erect, strong, sulcate, covered with
yellowish and thickish short hairs below inflorescence and often some-
what below. Leaves light glaucous, thickish, on both sides glabrous or
hairy beneath (sometimes rather densely) with hairs as on stem; basal
leaves usually 2, on thick petiole, purple like stem in lower part, as
long as or 1.5 times as long as lamina or shorter, lamina 9-45 cm long
and 4.5-20 cm wide, broadly ovate or oblong-ovate, obtuse or short-
acuminate, mostly with truncate or roundish-cuneate base, decurrent
on petiole more or less as wide wings, sinuate-toothed; cauline leaves
2-3, lower leaf often like basal or much narrower, with relatively
shorter, winged petiole, others strongly reduced, sessile, with amplexicaul
base sometimes expanded into auricles. General inflorescence 4.5-40 cm
long, compound paniculate, rather dense and pyramidal in upper part,
with usually widely spaced, short (2.5—5.5 cm long, less often up to 8-
9 cm) branches in lower part, bearing at their ends a panicle or less

839

797

often a raceme of densely clustered capitula, only in weaker plants branches
absent and inflorescence short, 4.5-7.0 cm long, as dense, cylindrical-
pyramidal panicle; bracteal leaves only at base of lower-most branches of
inflorescence, 1.0-1.5 cm long, ovate, others small, linear or filiform;
peduncles of capitula 1-3 mm long. Involucre 3.5—6.0(8.0) mm long and
2.5—3.0 mm in dia, narrow, tubular or obconical, narrowed toward base
with 1-3 small bracteal leaves at base; involucral bracts 5—6(8), firmly
contiguous at anthesis and up to dehiscence of fruits; inner bracts obo-
vate, obtuse,with wide scarious margin; outer bracts oblanceolate, obtuse
or lanceolately acute, with narrow scarious margin; all bracts glabrous or
hairy on back and with ciliate margin in upper part. Ligulate florets (1)2-
3, with elliptical, 5.5-9.0 mm-long ligule with 4—6 veins; tube 3-5 mm
long. Disk florets 4—7, with oblong-campanulate limb; tube as long as
limb or slightly shorter; anthers with short basal appendages, filaments
with weakly enlarged antheropodia; style branches short-pointed, usually
only with tuft of short hairs outside at base. Achenes yellowish-brown,
3.5-4.0(5.5) mm long, angular, finely ribbed; pappus white, almost as
long as corolla of tubular floret. Flowering second half of June to July;
fruiting second half of July to August.

Wet, including marshy and saline meadows, herb thickets around
sources of springs, shrub thickets in valleys, edges of forests and glades
in lower and middle mountain zones and particularly in Kazakh low
volcanic hills.—Western Siberia: Irtysh (extreme south), Altai
(southwest Chingistal steppe); Soviet Central Asia: Lake Balkhash
Region (in the lower mountain volcanic cones), Dzhungaria-Tarbagatai,
Tien Shan (Karatau Range, foothills of Talass Alatau and Tersk Range
in western Tien Shan). General distribution: Dzhungaria-Kashgaria
(Dzhungaria). Described from Chingistai steppe (northeastern part of
Kazakhstan). Type in Leningrad.

27. L. heterophylla Rupr. in Mém. Acad. Sc. Pétersb. VII, ser.
XIV, 4 (1869) 53 p. p. (saltem quoad var. thyrsoidea and excl. pl.
Schrenk.); O. and B. Fedtsch. Perech. Rast. Turk. III, 212 p. p. (quoad
pl. tiench.).—Senecio ledebourii auct. non Sch. Bip (1837): Franch. in
Ann. Sc. Nat. 6-me ser. Bot. XVI (1883) 313 (excl. synon.); in Bull.
Soc. bot. Fr. 39, 302 p. p. (quoad pl. turkest.).—L. persica auct. fl.
turkest. non Boiss. (1845): Lipsk. Gorn. Bukhara 1 (1902) 128, 129,
142, II (1902) 420, 441, 480, 531 p. p.; O. and B. Fedtsch. op. cit.
212; Pojarkova in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XII,
311 p. p.—L. macrophylla auct. fl. turkest. non DC. (1837); O. and B.
Fedtsch. op. cit. 212 p. p.—Ic.: Pojarkova, loc. cit. Fig. 3.

Perennial. Plant from 22 (in alpine zone) to 200 cm high; rhizome
like in previous species. Leaves thick, succulent, glaucous, on both

840

798

sides glabrous or with thickish hairs beneath; basal leaves 2, usually
with elliptical or oblong lamina, 7-52 cm long, pointed at apex and
with cuneate base, decurrent as wings on petiole, less often ovate, with
truncate or notched base, petiole a half as long as lamina or almost
twice times as long, sinuate-toothed, often with narrow sharp teeth;
lower cauline leaf like basal but with shorter petiole, upper leaf
(sometimes 2) much smaller, elliptical or ovate, sessile, often entire.
Inflorescence compound, mostly cylindrical, less often narrow-
pyramidal panicle, more or less dense in upper part, spicate, cylindrical
and often longer than lower part, with more or less numerous, widely
spaced, 1.5-5.0(8) cm-long branches, bearing capitula from middle or
less often almost from base in dense, sparsely branched panicles or
spicate racemes, size and degree of branching of inflorescence strongly
variable; in short plants from higher altitudes inflorescence 4-7 cm
long, weakly branched, almost racemose (L. heterophylla B. subramosa
Rupr. p. p. 1.c.—L. macrophylla var. minor Winkl. in sched.—Senecio
ledebourii 8. minor Franch.), in robust tall plants of forest zone up to
60 cm long with numerous remote branches (L. heterophylla y.
thyrsoidea Rupr.); stem below inflorescence, its branches and peduncles
usually more or less very densely covered with thickish yellow hairs;
bracteal leaves at base of lower branches of inflorescence small and
narrow, narrowly lanceolate, long-acuminate, up to 30 cm long, others
very small, more or less linear to filiform; involucre campanulate, 6—
9 mm long and 5-8 mm in dia with 1-3 filiform bracteal leaves at
base; involucral bracts 6-8; inner bracts broadly elliptical or ovate,
obtuse or short-accuminate, with wide scarious margin; outer bracts
narrow, elliptical or lanceolate, with narrow scarious margin, usually
long-acuminate; all bracts floccose in upper part along margin, glabrous
to densely hairy on back; after flowering involucral bracts distinctly
squarrose, becoming broader. Ligulate florets (3)4—5, with elliptical or
oblong 8-10 mm long ligule; tube 3.0—-5.5 mm long. Tubular florets
(7)10—14(16), 12-14 mm long, with narrowly campanulate or narrowly
conical limb, more or less gradually narrowed into short (2-3 mm
long) tube; style branches with subacute apical appendage, with tuft of
long hairs outside at base and shorter hairs below, sometimes up to
base of branches. Achenes 5-7 mm long and 1.0-1.5 mm wide, light
brown. Flowering May to first half of August; fruiting July to September.

Upper part of forest zone in cut-over forests, edges in forest glades,
scrubs with tall herbs and subalpine and alpine zones under adequate
moisture conditions.—Soviet Central Asia: Dzhungaria-Tarbagatai
(Dzhungarian Alatau), Pamiro-Alai (not throughout Pamir, in Trans-
Alai, Nuratau, Kuramin and apparently Turkestan ranges), Tien Shan
(Chatkal, Fergana ranges and outer, central and eastern Tien Shan

841

799

mountains). General distribution: Dzhungaria-Kashgaria. Described
from central Tien Shan. Type not preserved.

Note. The Central Asian plants of this species were described by
Ruprecht under the name L. heterophylla var. thyrsoidea Rupr. from
the materials of Osten-Sacken. At the same time, Ruprecht described
two more varieties of his species: var. alpina, which, apparently, should
be referred in part to L. alpigena Pojark., and var. subramosa Rupr.,
whose taxonomic status is not quite clear to me; from their brief
descriptions, they can be referred both to L. heterophylla Rupr. s. str.
and to L. alpigena. Using the precedent of the author who, for the first
time, split off the species, I selected var. thyrsoidea as the type of L.
heterophylla, considering it as a Tien Shan-Pamiro-Alai race of the
series Macrophyllae.

L. heterophylla should be distinguished from L. persica Boiss.,
with which I earlier combined it. L. persica has denser pubescence and
a denser inflorescence with very short branches (the lower branches
mostly are shorter than the middle ones) bearing rather tightly clustered
capitula from the base itself.

28. L. talassica Pojark. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XII (1950) 311.

Perennial. Rhizome short, densely covered with brown fibrous roots,
with fibrous remnants of dead leaves in upper part. Stem 45-100 cm
high, 3-8 mm thick, erect, sulcate-ribbed, usually more or less purple
below, glandular-hairy only below inflorescence, glabrous below.
Leaves glaucous, succulent, glabrous; basal leaves three (four),
outermost [lowermost] considerably shorter than other 2-3 leaves,
uppermost with 14-32 cm-long and 9-15 cm-wide lamina, oblong-
ovate or elliptical, sinuate or sometimes indistinctly toothed, obtuse
or, less often acute, with roundish-cuneate or cuneate base, more or
less decurrent, petiole often winged almost entirely, up to 2/3 as long
or as long as lamina, expanded at base, semiamplexicaul; cauline leaves
usually 3, lower of them slightly above base of stem, sometimes as
long as inner [upper] basal leaves but with shorter petiole, usually 2
lower cauline leaves oblong or oblong-elliptical, narrowed in lower
third and then expanded below, broader part being winged petiole,
whose base decurrent over short distance, and often auriculate below;
upper cauline leaf strongly reduced, ovate or oblong, with or without
constriction, sessile, often entire; upper bracteal leaf in shape and size
often like upper cauline leaf, others small, narrowly lanceolate to
filiform. Capitula in elliptical, narrowly pyramidal or cylindrical panicle,
11-30 cm long, almost spicate-racemose in upper part, with short, 1-
5(6) cm long-branches below bearing spicate racemes; peduncles of

800

capitula 2-5(7) mm long; capitula including ligulate florets 22—30(35)
mm in dia. Involucre campanulate and campanulate-bowl-shaped lower
8 mm long, others 9-12 mm long, 8-12 mm wide at flowering with 3-
5 linear or filiform bracteal leaves at base; involucral bracts (5)6—-8;
inner bracts ovate or obovate, with wide scarious margin; outer bracts
lanceolate or linear, with or without narrow scarious margin; all bracts
more or less glandular-hairy on back and along margin, apically
tomentose. Ligulate florets 4—6, fertile, usually with staminodes, with
oblong or linear ligule, (1.5)2—3 times as long as tube. Disk florets
12-20, with 8.0-9.5 mm-long corolla, limb narrow-campanulate, tube
2-3 mm-long; anthers with subacute basal appendages, antheropodia
weakly enlarged; style branches conical, acute, with a dense tuft of
long hairs outside below apex and scattered hairs below almost up to
middle. Achenes 6-7 mm long, light-colored; pappus white, almost as
long as achene. Flowering July to first half of August.

Meadow slopes, banks of mountain streams from upper part of
tree-scrub zone to beginning of alpine zone.—Soviet Central Asia:
Tien Shan (western part—Talass Alatau). Endemic. Described from
the upper reaches of Ak-Su River in Talass Alatau. Type in Leningrad.

Series 2. Alpigenae Pojark. in Bot. Mat. Gerb. Bot. Inst. Akad.
Nauk SSSR, XII (1950) 313.—Capitula in simple, usually dense, often
spicate racemes, less often lower peduncles bearing 2-3 capitula;
involucre campanulate-goblet-shaped or almost goblet-shaped, with 7—
12(14) involucral bracts.

Besides our species, this series also includes L. tangutorum Pojark.
(L. potanii Pojark. non Ling (1937) from central China.

29. L. alpigena Pojark. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk
SSSR, XII (1950) 313.—L. heterophylla «. alpina Rupr. and B.
subramosa Rupr. in Mém. Acad. Sc. Pétersb. VII, sér. 4 (1869) 59 p.
p.—L. altaica auct. fl. turkest. non DC. (1837); O. and B. Fedtsch.
Perech. Rast. Turk. III, 211 p. p. (quoad pl. tiansch. et pamiroalaic.)

Perennial. Rhizome strongly reduced, densely covered with cord-
like, rather slender or thickish, adventitious roots, with dense thin
fibrous remnants of dead leaves in upper part. Stem 10—40(75) cm
high, erect, sulcate, basally purple, glabrous, only below inflorescence
(as also peduncles) more or less densely covered with yellowish, flat,
multi-cellular, flexuous hairs and often also with white, fine, arachnoid
hairs. Lowermost 1-2 leaves scaly, others dark glaucous or green,
lighter beneath, somewhat thick, succulent, glabrous on both sides or
rather densely hairy beneath, with short, thickish, multicellular hairs;
basal and lower cauline leaves 5—25 cm long (excluding petiole) and

843

801

2-10 cm wide, with lamina of different forms from broadly ovate and
broadly elliptical to lanceolate, obtuse or more or less acuminate, with
truncate or often cuneate base, decurrent on petiole, entire or often, at
least some, with small, mostly remote teeth, often reduced to small
callous cusp, petiole a third to a half as long as lamina, glabrous,
mostly purple, winged entirely or only in upper part; middle and upper
cauline leaves sessile, amplexicual, elliptical or oblong-obovate, lower
leaves large, upper strongly reduced. Inflorescence 2-11 cm long, with
5-20 capitula, usually in rather dense, spicate, ovoid raceme, often
lower peduncles branched bearing 2—3(5) capitula; peduncles of capitula
2.5-7.0 mm long. Capitula including ligulate florets 1.0—2.2(3.0) cm
in dia, drooping after anthesis. Involucre bowl-shaped or campanulate-
bowl-shaped, less often campanulate, with 1-3 linear or almost filiform
bracts at base; involucral bracts 8—12(7—14), on back more or less
densely covered with yellowish flexuous hairs, short-ciliate in upper
part, along margin, (7)8—11 mm long, distinctly dimorphic: inner bracts
much broader than outer ones, ovate and with wide scarious margin;
outer bracts lanceolate, often without scarious margin or with scarious
border on one or both sides; all bracts acuminate. Ligulate florets 7—
12, with obovate or linear ligule 6—8(12) mm long, about 2 times as
long as tube and with 4—6 veins. Tubular florets numerous, up to
30(40), with 6.5-8.0 mm-long corolla; limb weakly enlarged, narrowly
campanulate, almost tubular, roughly 2 times as long as tube; anthers
with narrow, lanceolate, acuminate apical appendages and short basal
appendages filaments in upper part enlarged into antheropodia; style
branches short, with conical, somewhat broadened appendage, with a
tuft of small hairs outside at its base and with very short hairs below.
Achenes brownish-yellow, 5.0-5.6 mm long and 1.5-1.7 mm wide;
pappus somewhat longer than achenes. Flowering second half of June
to first half of August; fruiting from September.

Alpine (up to 3,600 m) and subalpine zones on meadow and stony
slopes, in juniper forests, and in Tien Shan in spruce forests.—Soviet
Central Asia: Pamiro-Alai (Trans-Alai, Alai ranges, eastern and western
Pamirs, Darvaz, Karategin, Peter the First ranges and western part of
Hissar Range—in basin of Sardai-Mion River), Tien Shan, western
(Fergana, Chatkal, Tashkent ranges, Angren Plateau), central and
eastern. General distribution: Dzhungaria-Kashgaria (Kashgaria and
Dzhungaria—Borokhoro Range). Described from south of Pamirs from
Mats River. Type in Leningrad.

30. L. altaica DC. Prodr. VI (1837) 315; Bong. and Meyer, Suppl.
fl. Alt. (1841) 149; Ldb. Fl. Ross. II, 621, (excl. synon. Linn. Gmel.
and area praeter Altai); O. and B. Fedtsch. Perech. Rast. Turk. III, 211

844

802

p. p. (excl. pl. tiansch. pamiroal.); Kryl. Fl. Zap. Sib. XI, 2865;
Pojarkova in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XII, 313.—
Senecillis glauca auct. non Gaertn. (1791); Ldb. Fl. Alt. IV (1833)
113 (excl. synon.).—Cineraria altaica C.A.M. ex DC. loc. cit. nom. in
synon.—Senecio altaicus Sch. Bip. in Flora, XXVIII (1845) 50; Franch.
in Bull. Soc. bot. Fr. 39 (1892) 292 (excl. pl. e Kansu).—L. glauca
var. altaica Kryl. Fl. Alt. III (1904) 664.—Exs.: P. Smirn. FI. alt. exs.
No. 94.

Perennial. Rhizome short, with numerous slender, filiform fibrous
roots, in upper part densely covered with fibrous remnants of dead
leaves. Stem usually solitary, glaucous or reddish, glabrous or
inflorescence including peduncles covered with rather small, papilliform
hairs, 10-65(80) cm high. Leaves glaucous, always glabrous on both
sides and entire; basal leaves and leaves of sterile rosettes with their
lamina elliptical or less often oblong or ovate, 4-18 cm long, 1.5-9.0
cm wide, decurrent on petiole, their petioles as long as lamina or up
to two thirds as long, in leaves of sterile rosettes thickened toward
base, not winged, in basal leaves usually winged throughout,
amplexicaul; cauline leaves 3-4, lower leaves identical with basal
leaves, others strongly reduced; all leaves sessile, semiamplexicaul.
Inflorescence with 5-25 capitula, usually aggregated in dense spicate
raceme, simple or lower peduncles bearing 2-3 capitula, very rarely as
elongated lax raceme. Capitula drooping after flowering, 2.5 (3.0) mm
in dia (including ligulate florets), with 3-10 mm-long peduncles, arising
from axils of narrowly linear bracts usually shorter than peduncles.
Involucre campanulate-bowl-shaped or campanulate, 5—9 mm in dia in
upper part, often with 1-2 small bracts at base; involucral bracts 6-10
mm long, 7-9, dimorphic, usually distinctly in 2 whorls; inner bracts
ovate, with wide scarious margin, outer bracts lanceolate, often lacking
border; all bracts acuminate, finely floccose at apex, on back always
glabrous. Ligulate florets 4-5, obovate, with 8-10 mm-long and 4—6
mm-wide ligule, with 4—7(8) veins. Tubular florets (8)10-—20; limb
narrowly campanulate; somewhat longer than tube; stamens with basal
appendages of anthers very short, filaments flat, thickened above in
antheropodia; style branches short, with conical appendage, with a tuft
of short hairs outside at its base and much shorter hairs below sometimes
up to very base of branches. Achenes 4.5-5.5 mm long and 1.5-2.0
mm wide; pappus white, slightly longer than achene. Flowering end of
May to August; fruiting from August. (Plate XXXIX, Fig. 1).

Alpine zone to upper part of forest zone, mostly in meadows and
along river valleys and streams.—Western Siberia: Altai; Soviet Central
Asia: Dzhungaria-Tarbagatai (Tarbagatai, Saur, Dzhungarian Alatau
mountains). General distribution: Mongolia, northwestern part

803

Plate XL.
1 — L. mongolica (Turcz.) DC.; 2— L. schmidtii (Maxim.) Makino.

847

804

(Mongolian Altai, region of Lake Ubs). Described from Kurchum River
in Altai. Holotype in Paris; isotype in Leningrad.

Subsection 2. Senecillis (Gaertn.) Pojark. op. cit. 894.—Senecillis
Gaertn. De fruct. II (1791) 453, tab. 173 p. p.—Capitula in simple,
usually dense, spicate racemes, less often with lower peduncles bearing
2-3 capitula. Involucre broadly campanulate or bowl-shaped; involucral
bracts 7-12, pappus hairs very short, 0.5-2.0 mm long, stiff, angular-
barbed. Stems hollow.

Type of subsection: L. glauca (L.) O. Hoffm.

Note. The two rather similar species belonging to this subsection
are characterized by a very short pappus, and, on this basis, L. glauca
was split off into the separate genus Senecillis Gaertn., whose
independent status is even now recognized by a number of authors.
However, I am not inclined to attach. much taxonomic significance to
this character, because it is observed even in certain species that are
not at all related to each other (L. jacquemontiana (Decne) Kitam., L.
angusta (Nakai) Kitam). As regards our species of subsection Senecillis,
they are undoubtedly closely related to the species of series Alpigenae,
which is manifest not only in their habit but also in the structure of all
floral parts except the pappus (cf. Plate XX XIX), whereas their links
with L. angusta and particularly with L. jacquemontiana are rather
distant.

31. L. glauca (L.) O. Hoffm. Pflanzenfam. IV (1897) 288; Kryl.
Fl. Zap. Sib. XI, 2864.—Cineraria glauca L. Sp. pl. ed. 2 (1763)
1242.—Senecillis glauca Gaertn. De fruct. et sem. II (1791) 413; DC.
Prodr. VI, 313 (excl. area Podolia).—Senecio senecillis Maxim. in
Mélang. biol. XI (1883) 243; Franch. in Bull. Soc. bot. Fr. 39, 291 p.
p. (quoad pl. alt. and excl. var. carpatica).—L. glauca var. typica
Kryl. Fl. Alt. III (1904) 664.—Ic.: Gaertn. op. cit. tab. 173, fig. 4.

Perennial. Plant glaucous, glabrous; rhizome short, with numerous
slender, filiform, fibrous adventitious roots and in upper part densely
covered with fibrous remnants of dead petioles. Stem erect, 50-150
cm high, (3)4—-10 mm thick at base, ribbed, often reddish-violet as also
petioles, below inflorescence sometimes including peduncles covered
with papilliform hairs. Leaves succulent, always with entire cartilaginous
margin; basal leaf and sometimes lower cauline leaf petiolate, petiole
as long or up to 1.5 times as long as lamina, their laminas oblong or
ovate, obtuse or acute, with truncate-sinuate or cordate, less often
almost hastate base, decurrent on petiole, 7-24 mm long and 4-13 mm
wide, much smaller in lowermost leaf than in others; cauline leaves
oblong-ovate or broadly ovate, lower 1-2 leaves with petiole usually

848

805

broadly winged others sessile, semiamplexicaul, upright, upper leaves
strongly reduced, often lanceolate. Capitula (8)10—20, mostly in dense
ovoid or less often elongated racemose inflorescence; in the latter
case, usually with few, rather remote capitula, elongated after flowering,
peduncles 7-10(12) mm long, up to 5 cm in lower capitula, lower
peduncles sometimes bearing 2-3 capitula; bracteal leaves lanceolate-
subulate, shorter than peduncles. Involucre bowl-shaped, 8—10(12) mm
in dia in upper part, after fruiting stellate, with 1-3 small bracts at
base; involucral bracts 10-—12(15), 5—8(11) mm long, often dimorphic.
Ligulate florets 7-12, with bright yellow, elliptical-oblong ligule, 8-
15(20) mm long and 4-7 mm wide, with 4-6 veins; tube a fourth to
a third as long as ligule. Tubular florets 18-29, with 7-8 mm-long
corolla; limb campanulate, 5 mm long and lobed into oblong-deltoid,
1 mm long teeth, but abruptly narrowed below into short narrow tube;
anthers with narrowly acute apical appendages and short, subacute
basal appendages, antheropodia rather wide; style branches acuminate
and outside covered with short hairs only in upper part. Achenes
yellowish-brown, 4—8 mm long, terete or somewhat cuneate, narrowed
toward base, angular, with 5-6 ribs; pappus caducous, of very short,
0.5—2.0 mm-long, thickish hairs, with dense barbs. Flowering second
half of June to July; fruiting from August. (Plate XXXIX, Fig. 2).

In floodplain and dry valley meadows, open birch and larch
forests.—Western Siberia: Ob River Area (southeastern part). Irtysh
(east), Altai, Angara-Sayans (western part). Described from Western
Siberia. Type in London.

32. L. carpatica (Schott, Nym., and Kotschy) Pojark. comb.
nova.—Senecillis carpatica Schott, Nym., and Kotschy, Analect. bot.
(1854); Schur. Enum. pl. Transsilv. 342 (excl. syn.).—Senecillis glauca
auct. non Gaertn. (1791) nec Ldb. (1833): Bess. Enum. pl. Volhyn.
(1822) 76; DC. Prodr. VI, 313 p. p. (quoad pl. Podol. and excl. synon);
Knapp. Pflanz. Galiz. 132 (excl. synon. Linn.): Schur. Sert. fl. Transsilv.
41; Enum. pl. Transsilv. 342; Katina, in Vizn. Rosl. URSR, 552.—
Senecillis glauca (podolica) Schott. Nym. Kotschy, op. cit. 5.—
Senecillis glauca var. transsilvanica Schur. op. cit. (1866) 342, nom.
in synon.—Senecillis heterophylla Schur. ibid., nom. in synon.—Senecio
senecillis auct. non Maxim. (1883); Franch. in Bull. Soc. bot. Fr. 39
(1892) 291 p. p. (excl. pl. sibir.); Schmalh. Fl. Sred. i Yuzhn. Ross.
II, 84.—Senecio senecillis var. carpatica Janka ex Franch. op. cit. 291.—
Lingularia glauca ssp. carpatica Heyek. f1. III (1923) 145.—Senecillis f.
carpatica Javorka, Magyar fl. Pflanzendecke Oesterr.-Ung. I (1945) 420.—
Ic.: Reichb. Ic. Fl. Germ. XVI (1853) tab. 958; Hayek, loc. cit. fig. 238;
Javorka, Iconogr. fl. Hung. (1934) tab. 533, fig. 3821.

849

806

Perennial. Distinguished from L. glauca (L.) O. Hoffm. by toothed
margins of all or only lower leaves as also by corolla of disk florets
with narrower limb and shorter tube and also by large number of veins
(6-10) on ligules of peripheral florets. Flowering July; fruiting
September.

Forest edges, herb slopes.—European part: Middle Dnieper
(Podolia, Pantalikh steppe near Trembovl). General distribution: Central
Europe (Transylvanian Mountains). Described from Transylvanian
mountains, Rodna District. Type not known.

Note. I had an opportunity to become familiar with L. carpatica
on the basis of two specimens (flowering and fruiting) from Besser’s
herbarium. Both specimens differ sharply, by their sinuate-toothed
leaves and narrow limb of the tubular florets, from the numerous speci-
mens of the western Siberian species L. glauca (Gaertn.) O. Hoffm.
examined by me.

In describing this species for the first time on the basis of material
from the vicinity of Rodna in Transylvania under the name Senecillis
carpatica Schott, Nym. et Kotschy, the authors contrasted it with the
Podolie golden-ray, which they mistakenly identified with Senecillis
glauca Gaertn. As the basic differences of the Transylvanian plant
from the Podolie plant, the following were mentioned: the absence of
a cartilaginous margin of the leaves (which has not been confirmed by
more recent studies), usually smaller and often simple racemose
inflorescences, a narrower ligule of the peripheral florets with fewer
(not 8-10 but 6-7) longitudinal veins, and also narrow, almost linear-
fusiform (and not cuneate) achenes. Judging from the data of Schur
(1866), who cited two species, Senecillis carpatica and “S. glauca”
(which is, in fact, the same as S. carpatica), with detailed descriptions
for Transylvania, including for Rodna), there is no basis to attach
special significance to the above-mentioned differences. This
presumption is indirectly supported by the examination of extensive
herbarium material of the vicariant species L. glauca (Gaertn.) O.
Hoffm., which revealed similar parallel variation in this species of the
above-mentioned organs, as was indicated for the plants from
Transylvania.

Subsection 3. Stereochaetae Pojark. op. cit. 894.—Involucral
bracts free or fused into 3—4-toothed cup; pappus hairs brown, stiff,
shorter than tubular florets; capitula small, borne in dense oblong
panicles. Stems solid; rhizome hairy.

Type of subsection: L. mongolica (Turcz.) DC.

Species of the eastern border of Eastern Asia.

850

807

Series 1. Mongolicae Pojark. ibid.—Involucral bracts free; pap-
pus 3-7 mm long.

Besides our species, this series includes L. fauriei (Fr.) Koidz.
(Korea) and L. taquetii (Lévl. and Vaniot) Nakai (Quelpart[Chuju]
Island).

33. L. mongolica (Turcz.) DC. Prodr. VI (1837) 315; Korshinski
in Tr. SPb. Bot. Sada, XII, 357; Kom. Fl. Manchzn. III, 819; Hand.-
Mazz. in Engl. Bot. Jahrb. LXIX, 1, 121; Kitag. Lineam. fl. Manshur.
458.—Cineraria mongolica Turcz. in Bull. Soc. Nat. Mosc. V (1832)
199.—Senecio mongolicus Sch. Bip. in Flora, XXVIII (1845) 50;
Maxim. in Bull. Soc. Nat. Mosc. LIV, 1, 28; in Mélang. biol. XI, 242;
Franch. in Bull. Soc. bot. Fr. 39, 292.—Senecio putjatae Winkl. in Tr.
SPb. Bot. Sada, XIV (1895) 125.—L. putjatae Hand.-Mazz. op. cit.
120.

Perennial. Plant entirely glabrous, smooth, glaucous, 50-100 cm
(and probably more) high; rhizome short, covered with numerous, thick,
cord-like, adventitious roots but very densely covered in upper part
with fibrous remnants of leaves and together with stem base pubescent
with weakly flexuous hairs. Stem solitary, 3.5—4.5 mm thick at base,
solid, basally mostly purple. Leaves fleshy, when dry usually rather
thick, with very distinct network of veins beneath; basal and sometimes
lowermost cauline leaf, as also leaves of sterile rosette, long-petiolate,
petioles as long as lamina or slightly longer, reddish at base and with
narrow sheath, lamina oblong-ovate, less often ovate or oblong, obtuse
or subacute, with roundish-truncate or cuneate base, decurrent on
petiole; cauline leaves 2—3, lower leaves identical to basal, other leaves
oblong, sessile, amplexicaul or lower leaves with short, broadly winged
petiole, uppermost leaves strongly reduced. Inflorescence 2—8 cm long,
as dense spicate, ovoid or cylindrical elongated raceme, sometimes
with 1-2 remote capitula; peduncles thick, 2-3 mm long, lower up to
1 cm long, borne in axil of narrow, lanceolate-subulate bracteal leaves
as long as capitula, lower peduncles sometimes bearing 1-2 similar
bracts. Involucre cylindrical, 9-12 mm long, lacking bracts at base;
involucral bracts 5(6), dimorphic, in 2 distinct whorls; inner bracts
oblong-obovate, obtuse or subacute, with wide scarious margin; outer
bracts almost a half as long, acuminate, with very narrow scarious
margin on one or both sides; all bracts hairy in upper part along margin
and at apex with thickish hairs. Ligulate florets (1)2—4, with golden-
yellow, oblong, 13-20 mm-long ligule, with 8—10(12) veins; tube a
half as long as ligule. Tubular florets 6-8, with narrowly campanulate
limb lobed almost up to middle into narrowly deltoid teeth; tube
approximately as long as limb; pappus as long as corolla; anthers with

851

808

subacute basal appendages, filaments of stamens thickened in upper
part; style branches long and slender, with very short conical appendage,
rather densely hairy from base of this appendage to very point of
bifurcation of style with long flattened hairs. Achenes almost fusiform,
grayish-brown, 4.0-5.5 mm long and 1.5 mm wide, with 5 weakly
developed ribs; pappus of thickish, reddish-brown hairs, with frequent
barbs such that hairs appear almost plumose. Flowering June to first
half of July; fruiting from July (Plate XL, Fig. 1).

Meadows.—Far East: Zeya-Bureya (one locality known near the
village of Konstantinovka-on-Amur). General distribution: China,
northeastern part in Greater Khingan, Inner Mongolia—in vicinity of
Kalgan and Hubai Province west of Peking. Described from southern
part of Inner Mongolia, from vicinity of Boroldzhi. Type and isotype
in Leningrad.

Series 2. Schmidtianae Pojark.—Cyathocephalum Nakai in Bot.
Mag. Tokyo, XXIX (1915) 11.—Involucral bracts fused into goblet
with 3-4 lobes; pappus 2.5—4.5 mm long.

Besides L. schmidtii (Maxim.) Nakai, the Japanese species L.
angusta (Nakai) Kitam. must also be included in this series.

Note. I agree with Kitamura (Kitamura, Compos. Japon. Ill, 185)
that fusion of the involucral bracts cannot be a criterion for separating
the above-mentioned two eastern Asian species into an independent
genus. In the subtribe Senecioninae, partial fusion of the involucral
bracts is observed also in certain species of the genus Cremanthodium.

34. L. schmidtii (Maxim.) Makino in Bot. Mag. Tokyo, XVII
(1903) 191 (quoad typ. excl. specim.); Kom. Fl. Manchzh. III (1907)
697; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1053; Kitag.
Lineam. fl. Mansh. 458; Kitam. Compos. Japon. III, 186.—Senecillis
schmidtii Maxim. in Bull. Acad. Sc. Pétersb. XVI (1871) 222 (in
Mélang. biol. VIII, 16) p. p. (quoad. specim. Schmidt.).—Senecio
schmidtii Franch. and Sav. Enum. pl. jap. I (1875) 246 (quoad typ.,
excl. specim.); Forb. and Hemsl. in Journ. Linn. soc. XXIII, 457;
Franch. in Soc. bot. Fr. 39, 291 p. p. (quoad. pl. Mandsh.).—
Cyathocephalum schmidtii Nakai in Bot. Mag. Tokyo, XXIX (1915)
deli;

Perennial. Plant glabrous, smooth, glaucous, up to 1-2 m high;
rhizome short, thick, densely covered with thick, cord-like, adventitious
roots, similarly densely covered in upper part with soft, fibrous remnants
of dead leaves, pubescent with simple flexuous hairs. Stem sulcate,
2.5—7.0 mm thick at base, solid. Leaves fleshy, dry, rather thick, with
prominent veins beneath; basal leaves long-petiolate, lamina 10-30 cm

852

809

long and 5-22 cm wide, broadly ovate to oblong, obtuse or short-
acute, with truncate or cuneate base, decurrent as wing mostly only on
upper part of petiole, more or less irregularly sinuate-toothed, less
often only undulate; cauline leaves oblong and oblong-ovate, lower
sometimes with short, broadly winged petiole, others sessile,
amplexicaul, adpressed to stem and upright, uppermost leaves strongly
reduced, small. Raceme at anthesis condensed, (3)6—13(18)[c¢m] long,
sometimes, including remote lower capitula, up to 23 cm long, simple,
less often with lower peduncles bearing 2-3 capitula; peduncles 10-15
mm long, lower 12-15 mm long, in remote lower capitula up to 4(7)
cm long, elongating after flowering and raceme becoming lax; lower
bracteal leaves small, with peduncles many times as long and usually
fused with them over considerable part, others not developed. Involucre
green, campanulate, fused, much broader at fruiting, bowl-shaped, with
3-5 teeth or lobes, obtuse or subacute, often with 2 very small short-
pubescent teeth at apex. Ligulate florets 3-6, with bright yellow ligule,
13-22 mm long and 2.2-6.0 mm wide, with 6-8 veins, tube a fourth
as long as ligule. Tubular florets 12-22, with 7-10 mm-long corolla;
limb campanulate, 5-7 mm long, lobed up to a fifth into narrowly
deltoid teeth and abruptly narrowed into short, narrow, 2.5-3.0 mm-
long tube. Pappus 3.0-4.5 mm long, somewhat longer than corolla
tube; anthers with narrow, acute, apical appendages and very short
basal appendages, filaments with narrow antheropodia; style branches
somewhat long, narrow, attenuate, outside up to point of bifurcation
covered with flat hairs. Achenes almost terete, slightly narrowed at tip,
ribbed, grayish-brown, 7-8 mm long; pappus dirty reddish-brown, a
half as long as achene, consisting of stiff hairs with dense short barbs.
Flowering second half of June to July; fruiting September (Plate XL,
Fig. 2).

Meadow slopes, in scrubs, open, oak forests, wet and open places.—
Far East: Ussuri (southern district). General distribution: China,
northeast (southern part), Korea. Described from Posjet in Ussuri.
Lectotype in Leningrad.

Section 5. Glossophyllum Pojark. sect. nova in Addenda, XXV,
894.—Capitula 1-3, with more or less distinctly 2 whorled involucre
of inner and outer bracts quite different from each other; bracteal
leaves small, green, lanceolate; style branches with conical appendage,
and tuft of long hairs outside at its base, but short hairs below; stigma
surface as 2 wide, raised lateral stripes divided by a groove;
antheropodia thickened, distinctly thicker than filament. Basal leaves
oblong, narrowed into petiole, entire or with a few weakly developed
teeth.

853

810

Type of section: L. pavlovii (Lipsch.) Cretz.

Note. The species L. pavlovii (Lipsch.) Cretz., included in this
section, differs rather sharply from other species of the subgenus
Ligularia by its characters (the structure of the stylodes and stamens,
shape of the leaves, presence of nonleafy membranous sheaths at the
base of the stem), and in the flora of Soviet Central Asia it is definitely
an ancient, narrowly endemic relict. Of all the species of the genus
Ligularia known to us, L. virgaurea (Maxim.) Mattf. can, to some
extent, be considered close to L. pavlovii.

35. L. pavlovii (Lipsch.) Cretz. in Journ. Jap. Bot. XVII (1941)
409.—Senecio pavlovii Lipsch. in Fedde, Repert. sp. nov. XXIX (1931)
162.—Ic.: Lipsch. ibid., tab. CX VIII.

Perennial. Rhizome short, densely covered with slender, filiform
adventitious roots and fibrous remnants of dead leaves. Stems one or
many, 18-35 cm high, and 2—3 mm thick, with few leaf rosettes, erect,
sulcate, covered, as also leaves beneath, with arachnoid white tomentum,
denser in upper part, with few nonleafy membranous swollen
amplexicaul, sheaths at base, sheaths initially brownish, later almost
white, transparent, along margin and at apex white-arachnoid-floccose.
Leaves thin; basal leaves in sterile rosettes with oblong-elliptical lamina,
5-10 cm long and 1.6-28.0 mm wide, acute or obtuse, decurrent on
petiole, petiole in first{lower] leaves two-thirds as long as lamina, in
secondary [upper] leaves half or less as long, midrib prominent, strongly
raised beneath, lateral veins very slender, 4—5 pairs; cauline leaves 2—
4, elliptical or oblong-elliptical, with short, amplexicaul, sheath-like
petiole, sometimes with 2 auricles, upper leaves sessile; all leaves
entire or sometimes with few remote teeth. Capitula solitary or often
2-3, borne corymbosely at branch apices; peduncles as also base of
capitula arachnoid-tomentose, 1-6 cm long, borne in axils of small,
sessile, lanceolate, acuminate leaves. Involucre 1.5—2.0 cm in dia,
campanulate-bowl-shaped, with 1-2 small lanceolate or linear bracts
at base; involucral bracts 8-12, pale green, on back weakly arachnoid-
hairy, distinctly 2 whorled, dimorphic: inner bracts oblong-ovate, short-
acuminate, with wide scarious border; outer bracts narrowly lanceolate,
often finely long-acuminate, without or with narrow scarious border on
one or both sides. Ligulate florets about 8, small, 9-14 mm long, with
elliptical, 7 mm long and 2.0-3.5 mm wide ligule, with 5 parallel
veins. Tubular florets up to 30, with 7-8 mm-long corolla; limb narrowly
campanulate, as long as or slightly longer than tube; apical appendage
of anthers short, ovoid, basal appendage short, filament enlarged in
upper part into antheropodia; style branches short, erect, with short,
conical, apical appendage, with a tuft of somewhat long hairs outside

854

811

at its base, but much shorter and thicker hairs below, often up to point
of bifurcation. Achenes lighter, stramineous, narrow, 6.5—7.0 mm long;
pappus snow-white, as long as achene. Flowering second half of April;
fruiting from end of May. (Plate XXXVIII, Fig. 2).

Stony and rubbly slopes.—Soviet Central Asia: Tien Shan (Syr-
Daryian, Kara-Tau Range). Endemic. Described from Ush-Uzen River
in Kara-Tau Range. Holotype in Herbarium of Moscow State Univer-
sity; isotype in Leningrad.

Subgenus 2. Dolichorrhiza Pojark. subg. n. op. cit. 895.—Rhi-
zome slender, woody, creeping. Capitula large, 1-6; involucre indis-
tinctly in 2 whorls, but involucral bracts more or less distinctly dimor-
phic, broader (inner bracts), with scarious margin and narrower bracts
enclosing them, lacking scarious margin and[or] with very narrow
scarious border on one or both sides. Antheropodia wide and thick;
style branches with very short, roundish, semiconical appendage, flat
on inner side, hemispherical in cross section; stigma surface as 2 raised
stripes divided by a furrow.

Type of subgenus: L. renifolia (C.A.M.) DC.

Three very similar Caucasian species belong to this subgenus.

36. L. renifolia (C.A.M.) DC. Prodr. VI (1837) 316; Fisch. and
Mey. Ind. sem. hort. Petrop. VII, 52; Ldb. Fl. Ross. II, 622.—Cineraria
renifolia C.A.M. Verz. Pflanz. Caucas. Casp. Meer. (1831) 82.—
Cineraria geifolia C.A.M. in sched. ex DC. op. cit. 316 (in synon.)
non L. (1763).—L. geifolia DC. ibid. Fisch. and Mey. loc. cit.—Senecio
renifolius Sch. Bip. in Flora, XXVIII (1845) 50, non Port. and Coult.;
Boiss. Fl. or. III, 401; Grossh. Fl. Kavk. IV, 147; Manden. in FI.
Gruzii, VII, 404.—Ic.: Manden. ibid., fig. 402.

Perennial. Rhizome oblique, creeping, up to 10 cm long, with few
branches at base and often arachnoid-tomentose at nodes, usually 3-
5 cm thick, narrowed toward apex, bearing one scale leaf and a tuft of
slender adventitious roots at each node, often developing 1, less often
2-3 flowering stems and few leaf rosettes, and then rhizome thicker
(up to 1 cm). Stem 12-55 cm high, slender, 1.5—2.0 mm thick in lower
part, glabrous, only at base arachnoid-tomentose, usually with | basal
leaf and 2 cauline leaves. Leaves reniform, often strongly expanded,
apically roundish, sometimes with a notch or groove, thin-papery when
dry, glabrous excluding margins, sparsely covered with thick, multi-
cellular hairs, sinuate-toothed with rather large, deltoid, abruptly
narrowed, callous, acute teeth separated by broad notches; basal leaf
1.5-7.0 cm long and 2.5-12.0 mm wide, petiole thin, 2-4 times as
long as lamina; lower cauline leaf arising from lower part of stem,

855

812

often larger than basal leaf and with as long petiole; upper cauline leaf
usually much smaller and short-petiolate; all petioles expanded into
short and narrow sheath, on inner side arachnoid-tomentose.
Inflorescence of 1 capitulum, rarely 2—4(5) capitula with remote,
peduncles 3.5-13.0 cm long, with thickish hairs’ at apex, in axils of
narrow, small, 0.8—3.5 cm-long, lanceolate or (lower) rhombic leaves
and with remote, subulate, small leaves, of which 2-3 upper leaves
often near base of involucre; capitula 3.5-5.5 cm in dia; involucre
bowl-shaped, 10-13 mm long; involucral bracts (7)9-14, often of
different shapes: lanceolate or lanceolate-linear, with distinct
membranous border, short- or long-acuminate, and much narrower,
linear, long-acuminate, lacking membranous border or with very narrow
border on one, less often on both margins; all bracts floccose-pubescent
at apices but elsewhere glabrous. Ligulate florets 8-13, with bright
yellow, 15-20 mm long, linear-elliptical to oblong-elliptical ligule,
with 4-6 veins, tube 4-5 mm long. Disk florets numerous, with 6-7
mm-long corolla; limb almost tubular, gradually narrowed into short,
about 2 mm-long tube; anthers with very short basal appendage; filament
with weakly enlarged antheropodia; style branches long, twisted, with
thick semiconical appendage, with a dense tuft of long (as long as
appendage) flat hairs at base of this appendage on inside of branch,
style branches in ligulate florets shorter and often lacking tuft of hairs.
Achenes 5 mm long, lighter, brownish-yellow, with few ribs; pappus
white as long as achene or slightly longer. Flowering second half of June
to first half of August; fruiting second half of August to September.

Alpine and subalpine zones in meadows and upper part of forest
zone, in pine and birch forests and shrub thickets in rather wet and
shady places, mostly in crevices of rocks and stones.—Caucasus:
Ciscaucasia, Western Transcaucasia, Dagestan, Glavnyi [Main]
Caucasus Range. Endemic. Described from western part of Glavnyi
[Main] Caucasus Range. Lectotype in Leningrad.

37. L. caucasica (MB.) G. Don, in Sweet, Hort. brit. ed. 3 (1839)
382.—Cineraria caucasica MB. FI. taur.-cauc. II (1808) 313, III, I,
573.—Senecio caucasicus DC. Prodr. VI (1837) 348; Ldb. Fl. Ross.
II, 622; Boiss. Fl. or. III, 401; Grossh. Fl. Kavk. IV, 147; Manden. in
Fl. Gruzii, VII, 404.—Arnica auriculata Willd. in sched. ex Ldb. loc.
cit. (pro synon.).—Arnica scorpioides Adams. in sched. ex Ldb. loc.
cit. pro synon.—Ic.: Manden. loc. cit. fig. 403.

Perennial. Rhizome as in previous species, but always lacking
arachnoid hairs and often thicker and then with thicker adventitious
roots. Stem erect, strong, sulcate, 23-85 cm high, 2-5 mm thick in
lower part, leafy, from middle or above irregularly and sparsely white-

856

813

arachnoid-hairy and usually densely pubescent below capitulum. Leaves
relatively thicker than in previous species, glabrous or covered with
thickish hairs along veins beneath and margin, sinuate-toothed; teeth
usually small, deltoid, with callous acute tip; basal leaf usually 1 (less
often 2), with strongly broadened, lamina 2.0—3.5 cm long and 5-9 cm
wide, reniform, with deeply notched base, roundish apex and large
roundish lobes at base, petiole long, thin (3)4—8 times as long as lamina;
cauline leaves (2)3—6, lower of them usually larger than basal leaf,
also reniform but less expanded and with shorter petiole; upper leaves
reduced upward, mostly hastate and short-petiolate, uppermost leaves
almost sessile, sometimes lanceolate; petioles of lower leaves usually
glabrous and sparsely pubescent, with narrow sheath, upper petioles
usually more densely pubescent, with mixture of arachnoid hairs, and
usually with 2 auricles, often large (up to 3 cm long), less often petioles
of all cauline leaves auriculate (var. auriculata Somm. and Lev. in Tr.
SPb. Bot. Sada, XVI (1900) 241). Common inflorescence of 1, less
often 2-6 capitula, corymbose; peduncles 3.5—11.0(13.0) cm long, with
a small lanceolate leaf at base and 1—3(4) remote, very small, often
subulate leaves above it. Capitula including ligulate florets (3.5)4—7
cm in dia. Involucre bowl-shaped (18-22 involucral bracts) with 3-5
narrow subulate bracts at base, often only slightly shorter than involucral
bracts; latter not sharply differing from each other in shape, narrowly
lanceolate or lanceolate, long—acuminate, with membranous margin on
one or both sides, along margin ciliate, apically floccose, glabrous
outside or sparsely arachnoid-hairy. Ligulate florets 15-17, with linear
or narrowly elliptical, saffron-yellow ligule 1.5—3.0 cm long, with 7—
10 veins; tube about 5 mm long. Disk florets numerous, with 9-11
mm-long corolla; limb campanulate-tubular and as long as tube; anthers,
shape of appendages of style branches, and hairs on the latter same as
in L. renifolia,; achenes 4—6 mm long, light colored, brownish-yellow,
ribbed-sulcate; pappus white, as long as achene or scarcely longer.
Flowering July to August; fruiting from second half of August.

Alpine and subalpine meadows also subalpine birch groves, par-
ticularly in crevices and shade of rocks and along streams. —Caucasus:
Ciscaucasia, Glavnyi [Main] Caucasus Range, Eastern Transcaucasia,
Western Transcaucasia (north and Kuba region), Southern Transcaucasia
(near Lake Sevan). Endemic. Described from Georgia. Type in
Leningrad.

38. L. correvoniana (Alb.) Pojark. comb. nova.—Senecio
correvonianus Alb. in Bull. Herb. Boiss. III (1895) 91 and in Tr. Tifl.
Bot. Sada, I (1895) 140; Grossh. Fl. Kavk. IV, 147; Manden. in FI.
Gruzii, VII, 403.—Ic.: Manden. ibid. fig. 401.

857

814

Perennial. Rhizome oblique, branched, with slender adventitious
roots, basally covered with large brown scales, developing many
flowering and vegetative shoots, glabrous like base of stems. Stems
slender, 20-45 cm high, finely sulcate, with 3—5 remote, small, almost
scaly, subulate or linear leaves and rosette of basal leaves, of which 1—
2 leaves sometimes in lower part of stem. Leaves thick, finely
coriaceous, glabrous, glaucous above, reniform, expanded, some
roundish, with broad or often narrow and deep notch and then lateral
lobes overlapping with their inner margins, sinuate-toothed, with usually
weakly developed teeth, often as one callous cusp, separated by shallow
notches, petiole 2.5—4 times as long as lamina, at base weakly expanded
into inconspicuous sheath, glabrous or in beginning with sparse
floccose-arachnoid tomentum. Capitula solitary, 3.5-5.0 cm in dia;
stem below capitulum grayish-tomentose for some distance. Involucre
10-12 mm long, goblet-shaped; involucral bracts 10-16, distinctly
dimorphic; broader, oblong-elliptical or elliptical-lanceolate with wide
membranous margin and not entirely regular, subsequent bracts
narrower, narrowly lanceolate or linear-lanceolate, with or without
membranous border on one or both sides; all bracts glabrous on back,
tomentose at apex, with 3—4 longitudinal veins and 1-3 small, linear,
apical bracteal leaves at base, sometimes bracts distinctly pushed
downward on peduncle. Ligulate florets 10-20, with oblong-elliptical
or linear ligule, 11-12 mm long and 2.0-4.5 mm wide, with 4—5 veins;
tube 3.5-5.0 mm long; style branches long, usually lacking hairs at
base of appendage; sometimes florets with staminodes also found.
Tubular florets 6.5—7.0 mm long; limb 4 mm long, tubular-campanulate,
with narrowly deltoid teeth and 2.5-3.0 mm-long tube; stamens with
very short basal appendages of anthers; style branches with a long tuft
of hairs outside at base of appendage. Achenes with white pappus,
according to Albov—as long as achene or up to twice as long. Flowering
June to July. (Plate XXXVII, Fig. 2).

Calcareous wet rocks and talus from the upper part of forest zone
to alpine zone.—Caucasus: Western Ciscaucasia, Western Trans-
caucasia. Endemic. Described from material from Abkhazia (author of
treatment not known to me) and from Western Transcaucasia
(Cherkesia). Type in Tbilisi or in Geneva.

Tribe 9. CALENDULEAE Cass. in Dict. Sc. Nat. XX (1821)
366, XXX (1824) 322, LX (1830) 573; Baill. Hist. des Plantes, VIII
(1882) 194; O. Hoffm. in Pflanzenfam. IV, 5 (1890-1894) 303.—Trib.
Calendulaceae Cass. in Bull. Soc. Philom. (1815) 173 and in Dict. Sc.
Nat. VI, Suppl. (1817) 35; Benth. in Benth. and Hook. fil. Gen. PI. II
(1873) 209.—Trib. Cynareae Less. subtrib. Calendulaceae DC. Prodr.

858

815

VI (1837)! 449, 451, p. p—Fam. Calendulaceae Bessey in Ann. Miss.
Gard. 2 (1915) 163.—Capitula heterogamous, with ray of pseudo-
ligulate florets; receptacle glabrous, less often weakly setose; anthers
mostly sagittate; style branches of bisexual florets flat, truncate; corolla
yellow or orange, less often ray florets white; pappus lacking or poorly
developed; achenes curved, heteromorphic, leaves as a rule, alternate.

Herbs, shrubs, and semishrubs, growing wild only in Old World,
mostly in tropics and subtropics; most members of Calenduleae are
concentrated in South Africa.

GENUS 1565. Calendula L.'”

ii Spuplhs ¢1753):-928:

Capitula many-flowered, terminal; involucre of 1-2 ead of
elongate involucral bracts. Outer (ligulate florets pistillate, with linear
stigma; inner florets tubular, bisexual, but sterile, with capitate stigma.
Achenes in 2-3 whorls, curved (to annular), heteromorphic: outer
achenes differing from middle and inner achenes by the form and
relief of their surface. Annual or perennial herbs, floccose-glandular,
with yellow or orange flowers; 20-30 species in the Mediterranean
Region, Western Europe and Middle East.

Some species (C. officinalis—“pot-marigold”’) are among the most
popular ornamental plants.

1. Capitula large (up to 4—5 cm in dia); cultivated plant ...........

as Bienes Ss Rhee uh. based wniets *C. officinalis L.
+ Capitula smaller; wild plants or weeds ............cccscceesseeeeseeeeeeeees 2.
Gi: . fiDuteriachenes daleate f weakly: qunyed . 4..4:.1...:s0scccedeoeses sete Lequengens 3.
+ Outer achenes strongly curved, more or less annular .................. 4.
3» Outer achenes not'winged,)/more/or less terete .......1....:..ssssessssvenanyge»

Ad. chet bed BE das BOR. Lies LB ccateatibenecncins 3. C. arvensis L.
+ Outer achenes winged with setose-toothed, lateral wings, lanceolate
etek ee, puch capa dak. Raaecialenmes. duc itacds 4. C. karakalensis Vess.
4. Outer achenes spinose-setose, usually winged (rarely lacking wings),
3 times as large as inner; capitula with fruits lax ............ce
esos cane Septem. ton eet toes mh is seek his Oa so as 1. C. persica C.A.M.
+ All achenes usually lacking wings; outer achenes tuberculate or
short-subulate, but not setose-spinose, not more than 1.5—2 times as

11937 in the original—General Editor.

'Treatment by I.T. Vassilczenko.
*From the Latin word calendae—first day of the month; indicative of prolonged
flowering.

859

816

long as inner, sometimes almost equal. Capitula with fruits more
SHSE UAT. LL cleat ei as OR oes 2. C. gracilis DC.

1. C. persica C.A.M. Verzeichn. Pfl. Cauc. (1831) 72; DC. Prodr.
VI, 453; Ldb. Fl. Ross. II, 2, 650; Boiss. Fl. or. III, 418; O.A. and
B.A. Fedtsch. Perech. Rast. Turk. [V, 221; Grossh. Fl. Kavk. IV, 155.—
C. arvensis MB. FI. taur.-cauc. III (1819) 596.—C. officinalis auct.
non L.: Hohen. in Bull. Soc. Nat. Mosc. VI (1833) 256.

Annual. Plants finely and glandular-floccose; stems few, (5)8-
20(30) cm long, ascending or procumbent, arising from branched root
collar. Basal leaves petiolate, oblong or oblong-obovate; cauline leaves
lanceolate or oblong-lanceolate, acute, usually sparsely toothed, with
small teeth or almost entire, sessile. Capitula 8-10 mm in dia, solitary,
at fruiting usually globose, at apices of stems.

Involucre in 1-2 whorls; involucral bracts uniformly long, oblong-
lanceolate, woolly-glandular, acuminate and often colored (dark purple).
Receptacle glabrous, weakly convex; ligules narrow, filiform-linear,
golden-yellow, 1.5—2 times as long as involucre. All achenes annular-
curved, densely pubescent; outer achenes brownish or brownish-
stramineous, considerably (thrice) as long as inner, 7-8 mm wide,
entirely spinulose on back, winged on inner side and along edges (lateral
wings often pectinate, with lanceolate unequal teeth or involute, and
then achenes carinate); inner achenes grayish, lacking wings, finely
punctate-tuberculate, on back often with obtuse small tubercles.
Flowering April to May.

Dry stony slopes in foothills, rubbly, sandy-stony and sandy plains
(foothills), sometimes as weed in fields.—Caucasus: Eastern and
Southern Transcaucasia; Soviet Central Asia: mountainous Turkmenia.
General distribution: Iran-Afghanistan, Armenia and Kurdistan, Iraq.
Described from the vicinity of Baku. Type in Leningrad.

2. C. gracilis DC. Prodr. VI (1837) 453; Ldb. Fl. Ross. II, 2, 650;
Grossh. Fl. Kavk. IV, 155.—C. persica C.A.M. var. gracilis Boiss. Fl.
or. III, 418.

Annual. Plant floccose-glandular. Stem usually branched from base,
ascending, 5—-15(30) cm long. Basal leaves oblong, with remote teeth,
sometimes indistinctly sinuate, acute, petiolate; cauline leaves shorter,
oblong-lanceolate, sessile. Capitula terminal, small, about (3)4-5 mm
in dia (at anthesis). Involucre in 1-2 whorls; involucral bracts oblong-
lanceolate, acuminate; ligules yellow, 1.5—2 times as long as involucre,
filiform-narrowly linear. All achenes curved, annular or almost annular,
lacking wings and beak; outer achenes only slightly larger than inner
ones (not more than 1.5—2 times as long), on back rugose-tuberculate

860

817

or spinulose, not setose-spinose; middle achenes broadly carinate, more
or less smooth, inner rugose tuberculate; all achenes more or less
homochromous, dark gray, crowded in capitulum forming more or less
dense “heads.” Flowering April to May.

Weed of fields, roadsides, sometimes on dry (clayey and stony)
slopes.—Caucasus: Eastern and Southern Transcaucasia; Soviet Central
Asia: Western Turkmenia (Krasnovodsk, Kara-Kala). General
distribution: Iran, Balkans-Asia Minor, Mediterranean Region.
Described from Karabakh. Type in Paris.

Note. This species (seemingly in secondary habitats) produces a
number of forms transitional to C. persica (hybrids?). In natural habitats,
C. persica is confined to sands and light (sandy, rubbly) soils but C.
gracilis to more compact, clayey-stony soils. This information needs
to be verified with more material.

Under better moisture conditions, the following features are
observed in plants of C. gracilis: sparser pubescence, thickened stems,
wider and longer ligules. Such plants are known, for example, from the
Firyuza Valley as also from the Alty-Togdan stream near Kara-Kala.
I refer them to a separate variety (C. gracilis var. firjusensis Vass.),
which, after collection of more complete material, may possibly be
considered as a species.

3. C. arvensis L. Sp. pl. (1763) 1303; DC. Prodr. VI, 452; Ldb.
Fl. Ross. II, 2, 650; Boiss. Fl. or. III, 418; Grossh. Fl. Kavk. IV, 155;
Klokov in Vizn. Ros. URSR, 558.—C. brachyglossa Rupr. in Bull.
Acad. Sc. Pétersb. XIV (1856) 230.—C. echinata DC. Prodr. VI (1837)
453.—C. byzantina DC. ibid.—Ic.: Sibthorp. Fl. Graeca, X (1840)
tabl. 920.

Annual. Plant floccose-glandular. Stem simple or branched, 10-30
cm high. Basal leaves oblong-obovate or oblong-spatulate, indistinctly
undulate-sinuate, weakly toothed; cauline leaves sessile, oblong, more
or less amplexicaul. Capitula 7-9 mm in dia, solitary, terminal.
Involucre in 1-2 whorls of acuminate bracts; ligulate florets twice as
long as involucre, sulfur-yellow. Outer achenes weakly curved, apically
attenuate (with truncate beak), long-spinose on back; middle achenes
carinate, bulged; inner achenes small, annular, curved, rugose. Flowering
May to June.

Fields, roadsides.—European part: Middle Dnieper, Bessarabia,
Black Sea Region, Crimea; Caucasus: Very rarely as an introduced
plant. General distribution: Balkans-Asia Minor, Mediterranean Region,
Central Europe. Type in London.

86

—_

818

4. C. karakalensis Vess. sp. nova in Addenda, XXV, 895.

Annual. Plants finely and profusely glandular-pubescent. Stem
branched from base. All leaves entire; lower leaves oblong, acute,
petiolate; upper ones oblong, also acute, sessile. Capitula small;
involucre in single whorl of oblong, acuminate, involucral bracts; ligules
yellow, only slightly (not more than 1.5 times) longer than involucre.
Outer achenes falcate, large (15-20 mm long), broadly lanceolate, with
longitudinal groups of spines and bristles on back in furrow formed by
2 lateral setose-toothed wings, winged on inner side and more or less
irregularly toothed; middle achenes carinate (with membranous wings
involute), with teeth and spines on back, broadly ovoid or ovoid-
roundish, a half as large as outer ones; inner achenes annular, lacking
wings, spinose-toothed on back, usually with membranous unilateral
process at apex. Flowering April to May.

Clayey slopes.—Soviet Central Asia: Mountainous Turkmenia
(western Kopetdag). Endemic. Described from vicinity of Kara-Kala.
Type in Leningrad.

*C. officinalis L. Sp. pl. (1753) 921; DC. Prodr. VI, 452; Ldb. FI.
Ross. II, 2, 650; Boiss. Fl. or. III, 416 (in nota); Grossh. Fl. Kavk. IV,
155; Klokov in Vizn. Rosl. URSR, 558; Pidotti, Opred. Semyan.
Dakorat. Rast. 81.—Ic.: Pidotti, ibid. Plate XXIV, 3, 4, 5 (pl.*).

Annual. Plant 20-50(75) cm high; stem often branched from base,
green, more or less floccose-glandular. Basal leaves oblong-obovate,
with remote teeth or entire, large, petiolate; cauline leaves oblong-
lanceolate or oblong-obovate, sessile, more or less amplexicaul,
indistinctly undulate and with small, occasional teeth. Capitula large
(up to 4—5 cm in dia); involucre in 1—2 whorls of oblong-lanceolate,
acuminate, involucral bracts. Florets yellow or orange, sometimes dull,
twice as long as involucre; ligules broad, up to 4.5 mm wide. All
achenes curved, yellowish or brownish to gray, trimorphic: outer
achenes semiannular, large (up to 15 mm long), on outside usually
with longitudinal rows of spines, with long, hollow, inwardly bent
beak, seemingly truncate at apex; middle achenes carinate, smaller and
broader (up to 10-12 mm long, 8-10 mm wide), with a longitudinal
rib and spines on back (sometimes without), with wing-like involute
lateral processes and a similar middle process; inner achenes grayish,
small, annular, spinose or tuberculate. Transients between these forms
of achenes observed. Flowering June to September.

It is cultivated everywhere as one of the most popular ornamental
plants; sometimes it is found growing as a weed in gardens, kitchen
gardens, etc. Described from Europe. Type in London.

819

Economic Importance. For some time, it has been cultivated for
a dye that is used in the food industry. It is also used as a medicinal
plant.

GENUS 1565a. Dipterocoma Fisch. and Mey.’ *

Fisch. and Mey. Ind. Sem. Hort. Petrop. 1 (1835) 26.

Capitula few-flowered, heterogamous. All flowers tubular; outer
flowers pistillate, weakly bilabiate; inner flowers staminate with 5 teeth.
Achenes terete, curved spinose on back, with 2 curved cornicles at
apex, sometimes some achenes without such cornicles and with short,
radial spines. Annuals with narrow alternate leaves.

One species, found in countries of the Middle East.

1. D. pusilla Fisch. and Mey. Ind. Sem. Hort. Petrop. I (1835) 26;
Boiss. Fl. or. III, 420; Jaub. and Spach, III, pl. or III (1847-1850) 34;
Grossh. Fl. Kavk. IV, 156; Nikitin in Fl. Turkm. 7, 140.—Jaubertia
koelpinioides Spach in Jaub. and Spach, op. cit. 131.—Koelpinia sessilis
Boiss. Diagn. ser. I, II (1845) 34.—Iec.: Jaub. and Spach, loc. cit. tab.
289.

Annual. Leaves sessile, somewhat thick, with single vein, obtuse,
initially weakly arachnoid-hairy like stem but later glabrous. Capitula
with 10-15 florets, 2-3 mm wide; involucre imbricate; achenes stellately
arranged, not shedding; 2 cornicles at apex, outer cornicle as long as
achene. Flowering March to April.

Clayey deserts, loess foothills—Caucasus: Eastern and Southern
Transcaucasia; Soviet Central Asia: mountainous Turkmenia, Kara-
Kum, Kyzyl-Kum; Amu-Darya, Pamiro-Alai Region. General
distribution: Eastern Mediterranean Region, Asia Minor, Iran. Described
from Transcaucasia. Type in Leningrad.

Tribe 10. ARCTOTIDEAE Cass. in Dict. Sc. Nat. II, Suppl.
(1816) 118; XX (1821) 364, XXIX (1823) 447, LX (1830) 573; Benth.
in Benth. and Hook. fil. Gen. Pl. II, 110; O. Hoffm. in Pflanzenfam.
IV, 5, 307.—Astérées on Corymbiferes sect. 8. Arctotides Cass. in
Bull. Soc. Philom. (1812) 191.—Trib. Cynareae Less. subtrib. VI.
Arctotideae Less. Synops. Comp. (1832) 15; DC. Prodr. VI, 449, 484.—
Fam. Arctotidaceae Bessey in Ann. Miss. Gard. 2 (1915) 163.—Capitula

'From the Greek words dis—twice, pteron—wing, and come—hair, so named
because of the corniculate protuberances of the achenes.
*Brought here from the errata on p. 756 of vol. XXVII.

820

heterogamous or homogamous. Receptacle glabrous, squamose, setose
or alveolate. Anthers at base undivided or sagittate, but lacking caudate
appendage. Style branches of bisexual florets very short, style of sterile
florets not lobed. Leaves mostly alternate, often basal only.

One of the small tribes of the family Compositae. whose species
are found only in Old World and, that too, almost exclusively in South
Africa.

Subtribe Gndelinae (“Gundelieae”’) Benth. in Benth. and Hook.
fil. Gen. Pl. II (1873) 211; O. Hoffm. in Pflanzenfam. IV, 5, 311.—
Cinarocephalae Less. div. II. Gundeliaceae DC. in Ann. Mus. Paris.
XVI (1810) 153.—Trib. Gundelieae Boiss. FI. or. III (1875) 421.—
Individual capitula aggregated in globose-capitate compound
inflorescence; involucral bracts spiny. Herbaceous plants.

GENUS 1566. Gundelia L.'?

L. Sp. pl. (1753) 814.

Capitula with 5-7 florets, of which one (central) flower fertile and
others (peripheral) sterile. All capitula clustered in globose-capitate
glomerules resembling one large head; involucral bracts in certain
capitula fused into testaceous membrane, with 5-7 spinules at apex;
corolla infundibuliform, exserted from openings of testaceous membrane
formed by fused involucral bracts. Achenes large, short-obpyramidal,
with small cupule instead of pappus. Perennial herbs with spiny leaves.
One species.

1. G. tournefortii L. Sp. pl. (1753) 814; DC. Prodr. V, 88; Ldb.
Fl. Ross. Il, 2, 464; Boiss. Fl. or. III, 421; Post, Fl. Syr. 2, 74; Grossh.
Fl. Kavk. IV, 156; Nevski in Sorn. Rast. SSSR, IV, 265.—G. glabra
Mill. Dict. ed. 8 (1768) 2.—G. purpurascens Bornm. in Beih. Bot.
Centralbl. IX (1939) 197.—Exs.: Bornm. (1892-1893) 1407.

Perennial. Plant large, glabrous or more or less arachnoid-hairy;
root thick, containing latex; stem 30-60 cm high, rough and thick,
simple or weakly branched above. Leaves coriaceous, stiff, large, oblong
or oblong-lanceolate, pinnately divided or lobate, with more or less
spiny-toothed lobes apically attenuate into spine, midrib thick and wide.
Capitula clustered; general inflorescence (glomerule of capitula),
globose-ovoid, surrounded by spiny bracteal leaves, 2-5 cm wide; bracts

'Treatment by I.T. Vassilczenko.
*Named after A. Gundelscheimer (1668-1718), one of the associates of Tournefort.

821

in individual capitula terminating into long coarse spine, exceeding
capitula. Corolla purple; fruit (achenes) enclosed in a hull (sheath)
formed by fused involucral bracts; nut-like “pseudo-fruit” gray or
grayish-brown, about 1.2—1.5(1.7) cm long and 0.8-1.2 cm wide,
apically with prickly spines. Flowering May to June; fruiting July.

Dry stony slopes in foothills and as weed in non-irrigated crop
field.—Caucasus: Eastern and Southern Transcaucasia; Soviet Central
Asia: mountains Turkmenia. General distribution: Eastern
Mediterranean Region, Asia Minor, Iran. Described from Syria. Type
in London.

Note. Plants with arachnoid-tomentose bracts and a finely sca-
brous sheath belong to a special variety (var. asperrima Trautv.).
Boissier (loc. cit.) distinguished one more variety, var. tenuisecta Boiss.,
with pinnate (to dissected) leaves with narrow lobes (segments).

Economic Importance: Achenes edible and contain oil; the young
shoots and leaves are also edible. The latex has irritant properties
(causes vomiting).

863

865

ADDENDA XXV
DIAGNOSES PLANTARUM NOVARUM IN TOMO XxXVI
FLORAE URSS COMMEMORATARUM

Diagnoses of the New Species in Volume XXVI

November, 1961

ANTHEMIS L.

1. Sect. Rumata Fed. sect. nova.

Achenia transverse angulata obconica vel obpyramidata non
compressa. Paleae lanceolatae plus-minus carinatae saepe subulatae.
Receptaculum convexum vel hemisphaericum. Involucri phylla nigro-
marginata vel omnino pallida saepe sphacelata. Herbae perennes vel
plantae suffruticulosae, rhizomate indurato vel radice lignosa; caules
simplices basi foliosi capitulum saepe unum sat magnum gerentes; folia
inciso-lobata vel pinnata ad basin caulis congesta et rosulam diffusam
formantia.

Sectionis typus—A. Soportana Alb.

Sectio nostra ob acheniorum formam cum sectione typica
Anthemidis generis convenit, sed ab ea omnibus characteribus,
praecipuae involucri phyllis sphacelatis, caudicibus hypogaeis
surculisque sterilibus e collo radici rhizomatisque emittentibus etc.
etc., eximie differt.

Sectionis nomen per conversionem anagrammaticam nominis
Marutae constitutum.

2. A. anahytae Woron. (descripsit Fed.).

Perennis, subglabra vel pilis sparsis obsita, rhizomate indurato
fusco caudicibus brevibus et surculis sterilibus foliorum rosulis
terminatis instructo; caules sat humiles, ca 10—15(20) cm alti,
adscendentes vel erecti, praecipue ad basin foliosi; folia caulina
inferiora et surculorum sterilium ambitu obovata vel spathulata, plus-
minusve petiolata, caetera sessilia, pinnata, segmentis oblongo-
lanceolatis vel linearibus integris vel in lacinulas partitis apice indivisis
vel tridentatis, rachide latiuscula edentula; folia caulina superiora
diminuta lobulata vel indivisa; capitula sat magna, ca 4—5 cm in diam.;
involucri plus-minus pilosi phyllis externis trianlaribus, internis oblongis
vel oblongo-obovatis, omnibus fuscis et ad margines late scariosis
membranaceis et atro-fuscis vel nigris; ad apicem rotundatis, ad externa
ad internis gradatim sphacelatis; ligulis albis oblongo-ellipticis, apice

866

823

obsolete tridentatis basin versus valde angustatis; achenia videtur A.
ibericae consimilia.

Habitat: in regione alpina montium Caucasi minoris Anataliae
orientalis.

T y pus: Azerbaidzhania, in monte Gjamisch jugi Murov-dagh,
ca. 3000 m s. m. 15 VII 1909. Leg. A.V. Schelkovnikov. Conservatur
in Herb. Inst. Bot. Acad. Sci. URSS in Leningrad (Holotypus et
Isotypus).

Affinitas: Ex affinitate Anthemidis ibericae M.B. et praesertim
ob formam ligulae florum radiantium insignis.

O bs.: Nomen speciei a cl. G. Woronow in honorem collaboratoris
suis nomine armeniaco femineo Anahyt dictis et in nostro tempore
incognitis datum.

3. A. tranzcheliana Fed. sp. nova.

Perennis, tota pilis minutis lanuginosis obsita, radice lignosa
indurata, caudice elongato apice ramoso residuis petiolorum emortuorum
praedito et caules rosulasque foliorum emittente; caules valde elongati,
30-45 cm alti, simplices, monocephali, ad basin et in parte inferiore
mediaque tantum foliosi apicem versus nudi, ad basin arcuati erecti vel
(in sicco) vix flexuosi, obsolete longitudinaliter striati, infra capitulum
non incrassati; folia caulina inferiora et surculorum sterilium petiolata
ambitu late ovata in lacinulas lineares inter se distantes indivisas vel
apice bi-tripartitas divaricatim bipinnatisecta utringue pilosa, rachide
foliorum edentula angusta; capitula mediocria, ca 3-4 cm in diam.,
involucri plus-minus pilosi phyllis externis triangularibus pallide fuscis,
internis-oblongo lanceolatis stramineis ad dorsum nervo mediano atro
fusco donatis apice scarioso-membranaceis et interdum nigro-marginatis;
ligulis albis late-ovatis integerrimis videtur reflexis; receptaculi
hemisphaerici paleis densis stramineis membranaceis acuminatis;
achenia (immatura) fusca laeviuscula videtur calva vel apice minutissime
marginata. Fl. V—VI.

Habitat: in fissuris rupium umbrosarum solo vulcanico, in
regione media montana Tauriae australis juxta Pontum Euxinum.

Typus: Tauria, distr. Sudak-Theodossia, prope stationem
scientificam marinam. Ad rupes umbrosas infra summum montis
Karadagh. 9 VI 1926. Leg. V. Tranzschel. Conservatur in Herb. Inst.
Bot. Acad. Sci. URSS in Leningrad (Holotypus). Specimina Topo-ac
paratypica: ibidem 29 V 1952. Leg. Al. Theodorov, An. Fedorov et M.
Kirpicznikov.

Affinitas: Affinis Anthemide sterili Stev., sed eximie differt
statura robustiore, caulibus valde elongatis, capitulis majoribus, foliis
divaricatim laciniatis, lacinulis inter se distantibus (nec approximatis)
longioribus, ligulis latioribus.

867

824

Obs.: Nomen specificum dedi in memoriam beati botanici
clarissimi professoris doctoris V. Tranzschel, qui plantam hoc loco
descriptam legit.

Sect. Cota (J. Gay ex Guss.) Rupr.

4. A. maris-nigri Fed. sp. nova.

Perennis, parce pilosiuscula vel pilosa, viridis vel vix incanescens;
caules e radice indurata a basi arcuata erecti stricte-ramosi, ca. 40-
60(70) cm alti, rigidi longitudinaliter striatuli, usque ad mediam partem
et ultra foliosi superne nudi; folia radicalia nulla vel cito decidua,
caulina—ambitu obovata pinnata, in segmenta latiuscula et obtusiuscula
oblongo linearia margine acuti-denticulata fissa, dentibus inaequelibus
sursum orientibus minutissime mucronulatis; capitula sat parva, ca, 2
cm in diam., ligulis luteis abbreviatus obtusis apice obsolete tridentatis
disco subduplo brevioribus; involucri villoso-lanati phylla externa tri-
angularia acutissima, interna oblongolanceolata obtusiuscula, omnia
ad costam mediam nigrescentia apice fusco barbellata; receptaculi
hemisphaerici paleae fuscae oblongae in mucronem rigidum
nigrescentem subito contractae; achenia (immatura) straminea transverse
subtetragona apice obsolete denticulata facie laeviuscula. VI-IX.

Habitat: in regione silvatica montium Caucasicarum
occidentalium.

T y pus: Caucasus occidentalis (olim Circassia), in abruptis inter
fluvios Schache et Czuchuk. 10 IX 1927. Leg. G. Woronow et W.
Steup. Specimina paratypica: Abchasia, in pratulis juxta confluvium
Bzybj et Gega. 23 VII 1930. Leg. P. Panjutin; Prope urbem Tuapse.
23 VI 1915. Leg. D. Litwinow. Conservantur in Herb. Inst. Bot. Acad.
Sci. URSS in Leningrad.

Affinitas: Inter plures species generis Anthemidis e grege
Tinctoriae m. ob ligulas abbreviatas eminens; differt ab Anthemide
tinctoria, A. monantha et al., praeter notas indicatas, foliis abbreviatis
virescentibus, caulibus longissimis stricte-ramosis, involucri phyllis
apice fusco fimbriatis et habitatione silvatica montana.

5. A. markhotensis Fed. sp. nova.

Perennis, tota ob pilos densos sericeo lanata; caulis in media
parte ramosus foliosusque, ca 40-50(60) cm altus, basi et sub capitalum
efoliatus, striatellus, rubescens; folia sat densa majuscula ca 3-4 cm
longa, ambitu obovata pinnata, segmentis latiusculis oblongo obovatis
laciniatis, lacinulis inciso mucronulato denticulatis; capitula majuscula,
ca 2-2.5 cm in diam.; ligulis luteis abbreviatis apice rotundatis
integerrimis vel interdum obsolete denticulatis, oblongo ovatis disco

868

825

manifeste brevioribus; involucri lanati phylla externa triangularia acuti-
acuminata, interna oblonga obtusiuscula, omnia pallida; receptaculi
hemisphaerici paleae fuscae, externae in mucronem sensium angustatae,
internae—e basi retusa et bilobulata subito contractae; achenia straminea
laevia costulata apice inaequaliter et obsolete denticulata. VI-IX.

Habitat: in abruptis sterilibus vel herbosis solo calcareo, in
jugo Markchot et ad littora Maris Nigri.

Ty pus: Caucasus occidentalis, in distr. Gelendzhik, in jugo
Markhot. n° 58, 1936 IX. Leg. E. Crniakowska, Specimina paratypica:
Distr. Gelendzhik, ad promontorium Tolstyi-myss, in abruptis calcareis
littoralibus. n° 25, 1936 IX. Leg. E. Czerniakowska; ad summum jugi
Markhot supra sinum Maris Nigri, in declivibus herbosis. 6 VI 1911.
Leg. I. Palibin et N. Vorobjev..

Affinitas: Inter omnes species seriei Tinctoriae m. ob
pubescentiam densam sericeo-lanatam foliorum forman et magnitudinem
eorum insignis.

6. A. debilis Fed. sp. nova.

Perennis, ob pilos breves sat densos canescens; caulibus e radice
indurata fusca adscendentibus partim subprostratis debilibus parum
ramosis vel simplicibus sat tenuibus, ca 15—20(40) cm altis, usque ad
partem mediam foliosis superne nudis; folia tantum caulina valde
approximata abbreviata parva, ca 7-10(15) mm longa, ambitu late ovata
pinnata, segmentis linearibus acutis et acute dentato-lanciniatis
approximatis, lacinulis acutis mucronulatisque; capitula mediocria vel
parva, ca 2.5-3 cm in diam., ligulis aureo-luteis, interdum reflexis
oblongo lanceolatis apice obsolete denticulatis; involucri lanati phylla
omnia pallida, externa triangularia ad costam mediam virescentia,
interna oblongo lanceolata obtusiuscula apice densius tomentella;
achenia ignota. V—VI.

Habitat: in declivibus montosis siccis apertis et inter arbores.

T y pus: Asia minor, distr. Artvin, in angust. Godrochevi. 11 VI
1909. Leg. P. Nesterov. Specimina paratypica: In ejusden districtu,
Sanonet—tapa, 25 VI 1909; prope pag. Toigom, 13 VI 1909; prope
pag. Kvartzchany, 21 VI 1909; prope custodiam Gurzhany, 30 V 1910
(Omnia specim. a cl. P. Nesterov lecta).

Affinitas: E grege Anthemidis tinctoriae s. 1., differt foliis
valde abbreviatis parvis et approximatis, capitulis minoribus, ligulis
abbreviatis, caulibus debilibus parum subprostratis. Probabiliter cum
Anthemide brachyglossa C. Koch convenit, sed ob defectum
speciminum authenticum Kochianum mea manu in statu speciei novae
hoc loco descripta est. Plusminus accedit ad Anthemidem cretaceam
Zefir. sed ab ea ob caules simplices distinguenda.

869

826

7. A. linezevskyi Fed. sp. nova.

Perennis, virescens, radice dura, tota pilis adpressiusculis parce
tecta; caules elongati stricti, ad basin subarcuati dein erecti,
longitudinaliter striati, monocephali, ca 60 cm alti, simplices, a basi
usque ad dimidiam longitudinis foliosi, superne aphylli, maturitate rigidi
virguliformes; folia radicalia cito decidua; caulina—ambitu oblonga
sursum declinantia, pinnata, in segmenta anguste-linearia laciniato-
denticulata divisa, dentibus laciniisque sursum orientibus inaequalibus
acutissimis; rhachide foliorum angusta plus minus laciniata, capitula
mediocria, ca 3 cm in diam.; ligulis albis disco vix brevioribus oblongo
ellipticis utrinque angustatis apice retusis vel obsolete bi-tridentatis;
involucri villosuli phylla omnia pallida, externa—triangularia lanceolata,
interna—oblongo lanceolata obtusiuscula apice fusca vix fimbriata;
receptaculi hemisphaerici paleae straminae membranaceae acutissimae
acuminatae, achenia violacea apice latiuscule marginata. V—VI.

Habitat: in declivibus sat aridis et in dumetis, in montibus
Kopetdaghensibus Turcomaniae.

T ypus: Montes Kopet-dagh occidentales, distr. Kara Kala, in
declivibus australibus montis Chasardagh, in angust. Joldere, alt. 1350
m s. m. 23 VI 1931, n° 322. Leg. I. A. Linczevsky. Specimina
paratypica: In ejusdem districtu. ad declivitatem montis Sjunt prope
summum, alt. 1550-1650 m s. m. 11 VI 1931. n° 362 ej. lectum.; in
ejusdem distr., sine indicatione exacta; in monte Sjunt, ad decliv.
borealem, inter frutices, alt. 1100 m s. m. 12 VI 1930. Leg. E. G.
Bobrovl; in declivibus occidentalibus montis Chasardagh, in umbra
arborum alt. 1350 m. 19 VI 1930, n° 355. Leg. E.G. Bobrov; Kopet-
dagh, in angust. Joldere 21 V 1917. n° 149. Leg. E. Korovin;
Turcomania, distr. Kara-kala. In montibus Kassa-dagh orient., in agris
derelictis. 24 VII 1935. Basil. Nikitin.

Affinitas: Plus minusve affinis Anthemide dumetori Sosn.,
sed ob folia magis elongata virescentia (nec canescentia), segmenta
angustiora valde elongata, inter se distantia, involucri phylla interna
apice fusco-fimbriata et ob habitum alienum primo aspectu insignis.

Obs.: Nomen specificum dedi in honorem amiciss I. A.
Linczevsky, investigatoris cl. florae Asiae Mediae, Sinae austro-
occidentalis et tropicae etc., etc., qui plantam hoc loco descriptam in
variis locis Turcomaniae legit.

8. A. talyschensis Fed. sp. nova.

Perennis, radice dura pluricepiti fusca ad collum residuis petiolorum
emarcidorum instructa; tota planta ob pilos densos vel densissimos villosos
tomentum formantes incana; caules e basi incurvata adscendentes vel erecti
plus minus ramosi, ramis capituliferis monocephalis vel partim sterilibus,

870

827

usque ad dimidium longitudinis foliosi dein nudi, ca 20-30 cm alti,
internodiis valde albo lanuginosis abbreviatis; folia brevia ambitu late ovata
omnia sessilia in segmenta oblongolinearia denticulata approximata fissa;
capitula medioria, ca 3-4 cm in diam.; ligulis albis oblongis utrinque
angustatis apice obsolete tridenticulatis discum aequantibus vel eo vix
longioribus; involucri villosi phylla externa triangularia, interna oblongo
lanceolata, omnia pallida, apice margineque scarioso membranacea;
receptaculi hemisphaerici paleae fuscae membranaceae oblongae in
mucronem acutum brunneum abrupte angustatae; achenia obconica
prominenter costulata fusca apice coronulis obsolete denticulatis terminata;
fl. V-VII; fr. VIII.

Habitat: in declivibus lapidosis montium Talyschensium in
regione media ac arida.

T y pus: Prov. Baku, distr. Lenkoran, inter pagos Tuli et Dzhoni.
12 V 1912. Leg. A. A. Grossheim. Specimina paratypica: montes
Talyschenses, prope pag. Czairu, alt. 4800 m s. m. 4 VIII 1897. Leg.
Th. Alexeenko; Talysch, in steppis inter pagos Amuram et Dugja, alt.
4000 m s. m. 12 VI 1894. Leg. A. Lomakin; Transcaucasia,
Azerbaidzhan, prov. Baku, distr. Lenkoran, inter pagos Lerik et Tellavaj.
11 VII 1930. Leg. L. Prilipko; Ad trajectum inter pagos Lerik et
Busaczar, in declivitate australi ac austro-orientali lapidosa, alt. ca
1700 m s. m. 3 VII 1931, n° 587. Leg. N. Schipezinsky; Distr. Lenkoran,
inter montes Naduz-galassi et Tevile-band. 13 VII 1929. Leg. L.
Prilipko.

Affinitas: Ex affinitate Anthemidis dumetori Sosn. et ab ea
optime differt foliis valde abbreviatis densius lanuginoso-villosis,
internodiis caulibusque abbreviatis, involucri phyllis villosis et
ramificatione aliena. Ob pubescentiae densitatem, foliorum formam,
habitum quasi alpinum, involucrum characterem et aream geographicam
separatam inter omnes species seriei Rigescentis m. insignis et
incomparabilis.

9. A. abagensis Fed. sp. nova.

Perennis, glabriuscula vel pilis scabridis albidis parciore obsita,
sed saturate-viridis, radice fusca lignosa; caulis erectum monocephalus
usque ad mediam partem foliosus superne nudus striatellus, ca 40-50
cm altus, basi arcuatim in curvatus; folia radicalia cito decidua, caetera
caulina sessilia ambitu lato obovata pinnata, in segmenta latiuscula
circumscriptione late ellipsoidea pectinatim incis et obtusi-dentata
divisa, dentibus apice callosis et minutissime subulatis; capitula
medioria ca 4—4.5 cm in diam.; involucri subglabri vel pilosiusculi
phyllis externis obongo-trigoniformibus marginibus anguste atro fuscis,
internis-oblongis ad dorsum flavido-viridibus, ad margines apicemque

828

late-atrofuscis vel nigrescentibus scarioso membranaceis fimbriatisque;
receptaculi haemisphaerici paleis membranaceis pallidis acuminatis;
ligulis albis in sicco vix lutescentibus oblongis disco vix brevioribus
integerrimis; achenia ad huc ignota. VII.

Habitat: in pratis subalpinis montium Caucasicorum tantum
occidentalium.

T y pus: Caucasus occidentalis, Reservatum Caucasicum, in monte
Abago. 24 VII 1956. Leg. Erschov. Conservatur in Herb. Inst. Bot.
Acad. Sc. URSS (Holotypus). Isotypus—in Herb. Reserv. Caucasici.

Affinitas: Indubie ex affinitate Anthemidis zyghiae Woron.,
sed primo intuitu foliis glabriusculis saturate viridibus (nec cinereo-
incanis) vel ob pilos sparsos scabridis (nec velutino-lanuginosis) eximie
differt.

10. A. schischkiniana Fed. sp. nova.

Perennis, pilis albidis parce obsita, viridis; radice lignosa; caulos
plures vel solitarii, monocephali, a basi et ad mediam partem foliosi,
superne nudi, ca 50-60 cm alt., e basi arcuata erecti, striatelli; folia
lasalia nulla vel probabiliter cito decidua; caulina-ambitu late ovata,
sessilia, in segmenta lacinulasque bi vel tripinnatisecta, lacinulis
oblongo linearibus acutis minutissime mucronulatis; capitula medioria,
ca 4 cm in diam., involucri subglabri vel parce pilosi phylla externa
oblongotrigona, interna—oblongo lanceolata fusco scariosa
membranacea; receptaculi hemisphaerici paleae membranaceae
acuminatae discum vix superantes; ligulae albae oblongo-latelanceolatae
sursum deorsumque angustatae apice obsolete bi-tridentatae vel retusae
in sicco vix lutescentes; achenia nulla. VIJ—VIII.

Habitat: in rupibus regionis montanae mediae partim silvosae
Transcaucasiae austro-occidentalis.

Typus: Transcaucasia, Georgia, prov. Kutais, distr. Ozurgety
(Guria), prope Bachmaro, ca 2000 m, in rupestribus. 17 VII 1925.
Leg. A. Grossheim (A. Grossheim et B. Schischkin, Plantae orient.
exs., n° 245, Anno 1928), (sub Anthemide platyglossa). Specimina
paratypica: Jugum Adzharo-Imereticum, in monte Schavnabad regione
alpina. 6 VIII 1914. Leg. E. I. Kikodze; in districtu Achaltziche, prope
Abbas-tuman, in silvis coniferis ac mixtis, prope Zekaram. 12 VIII
1914. Leg. E. I. Kikodze. Conservatur in Herb. Inst. Bot. Acad. Sci.
URSS in Leningrad.

Affinitas: Ob folia saepius tripinnatisecta et ob lacinulas
minutissimas inter omnes species seriei Rigescentis m. insignis.

Nomen speciei descriptae dedimus in honorem B.C. Schischkin,
redactoris «Florae URSS>» illustrissimi ac praeclari.

871

829
MATRICARIA L.

11. M. tzvelevii Pobed., sp. nov.

Annua. Radix brevis, autumno dura; caulis tenuis, a basi
multiramosus, 8-15 cm altus, primo vere pilis setosis strictis brevibus,
post anthesin solum sub calathidiis pilosus; folia oblonga, 15-20 mm
lg., 3-5 mm It. imparipinnata foliolis simplicibus crassiusculis teretibus
mucronulatis, hispidis vel glabris, suprema-saepe indivisa linearia;
pedunculi monocephali, tenues, 2-5.5 cm lg. Calathidia 7-10 mm in
diam., receptaculum anguste-conicum, cavum, torulosum; involucrum
subuniseriatum, phyila involucri late elliptica, margine albo-
membranaceo lacera; flores radii 1.5-3 mm lg., 1.5 mm It., ligulae
breves, latae, albae, apice tridentatae, sub anthesi revoluti, flores disci
tubulosi flavi limbo 5-dentato, dentibus subtriangularibus, tubo in medio
strangulato; antherae apice appendicibus brevibus lanceolatis, stylus
flosculi radii exsertus, stylus flosculi disci occultatus; achenia 1.5 mm
lg., 0.5 mm It., teretiuscula, compressa, apice obliqua, costis guinque
tenuibus, pappus nullus. VIII.

Habitat: in declivibus schistosis.

Spec.typus: Tauria, dist. Sudak, in declivibus montis «<Bolvan»,
prope pagum Ujutnoe, 11 VIII 1960, leg. N. Tzvelev.

A. Matricaria recutita L. caulibus humilioribus, 8-15 cm alt. (nec
10-30 cm), prostratis, foliis saepe hispidis, calathidiis minoribus 7—10
mm in diam. (nec 10-15 mm) differt.

PYRETHRUM Zinn

12. Sect. Leptanthemum Tzvel. sect. nova.—Plantae herbaceae,
15-50 cm alt., pilis accumbentibus bifurcatis et simplicibus plus
minusve tectae; caules parce foliati, foliis bipinnatisectis. Calathidia
vulgo numerosa, in inflorescentiam laxam corymbiforme-paniculatam
congesta; involucra late semiglobosa, 6-11 mm in diam., phyllis internis
exteris sesqui, rarius duplo longioribus; receptaculum subplanum;
ligulae late ovales, S-7 mm lg. Achenia 1.5—2 mm lg., nervis (4)5(7)
longitudinalibus percursa, pappo circa 0.2 mm lg., irregulariter dentato
vel lobato.

Typus sectionis (species unica): Pyrethrum leptophyllum Stev.

13. Sect. Balsamitopsis Tzvel. sect. nova.—Plantae herbaceae,
30-70 cm alt., pilis bifurcatis adpressis valde compressis dense tectae;
caules parce foliati; laminae foliorum radicalium integrae. obovatae
vel late ellipticae, caulinorum—lyrato-pinnatifidae. Calathidia 1-6,

872

830

saepe inflorescentiam corymbiformi-racemosam formantia; involucra
semiglobosa, 8-15 mm in diam., receptaculum parum convexum,
subplanum; ligulae oblongo-lineares 9-17 mm lg. Achenia 2.5-2.8 mm
lg., nervis 5-8 longitudinalibus percursa, pappo 0.3-0.4 mm lg.,
irregulariter obtuse dentato.

Typus sectionis (species unica): Pyrethrum Kelleri (Kryl. et Plotn.)
Krasch.

14. Sect. Pyrethrellum Tzvel. sect. nova.—Plantae herbaceae, 5—
80 cm alt., pilis brevibus bifurcatis et simplicibus vel solum simplicibus
plus minusve tectae, saepe subglabrae; laminae foliorum plus minusve
pinnatisectae. Calathidia vulgo solitaria, rarius 2-—10(40) et
inflorescentiam laxam irregulariter corymbiformem formantia; involucra
semiglobosa, 8—20 mm. in diam.; receptaculum vulgo valde convexum
semiglobosum, rarius parum convexum; ligulae oblongae vel lineares,
7-25 mm lg. Achenia 1.8-3.2 mm lg., nervis 5-8 longitudinalibus
percursa, pappo 0.1-0.5 mm lg., rarissime (P. Komarovii Sosn.) 0.5-
1.3 mm lg., irregulariter dentato vel lobato.

Typus sectionis: Pyrethrum coccineum (Willd.) Worosch.

15. Sect. Xylopyrethrum Tzvel. sect. nova.—Plantae prope basin
plus minusve lignescentes, 5-20 cm alt., pilis adpressis vel
accumbentibus bifurcatis vel bifurcatis et simplicibus plus minusve
tectae; caules parce foliati; laminae foliorum plus minusve pinnati-vel
palmati-sectae, lobulis terminalibus angustissimis. Calathidia solitaria,
involucra semiglobosa, 5.5-12 mm in diam.; receptaculum valde
convexum conico-semiglobosum; ligulae ovales vel oblongae, 7-12
mm lg. Achenia 2-3 mm lg., nervis 5—7 longitudinalibus percursa,
pappo 0.2-1 mm lg., irregulariter dentato vel lobato.

Typus sectionis: Pyrethrum Kotschyi Boiss.

16. Sect. Richteriopsis Tzvel. sect. nova.—Plantae herbaceae, 15-—
35 cm alt., pilis accumbentibus bifurcatis (rarius partim simplicibus)
dense tectae; caules parce foliati; laminae foliorum bipinnatisectae,
lobulis terminalibus vulgo angustissimis. Calathidia solitaria, rarius 2—
8, involucra semiglobosa, 9-20 mm in diam.; receptaculum parum
convexum, subplanum; ligulae oblongae vel oblongo-lineares, 7-20
mm lg. Achenia 1.8-3.2 mm lg., nervis 8—12 longitudinalibus percursa,
pappo 0.1—0.3 mm lg., irregulariter dentato vel lobato.

Typus sectionis: Pyrethrum sericeum (Adam) M.B.

17. Sect. Brachyglossa Tzvel. sect. nova.—Plantae prope basin
plus minusve lignescentes, 10-50 cm alt., pilis sat longis bifurcatis et

873

831

simplicibus vel solum simplicibus plus minusve tectae; caules parce
foliati; laminae foliorum pinnatisectae. Calathidia solitaria, rarius 2-5;
involucra anguste semiglobosa, 6-13 mm in diam., receptaculum parum
convexum, subplanum; ligulae ovales, rarius oblongae, 6-12 mm lg.
Achenia 2.3—3.2 mm lg., nervis 6-8 longitudinalibus percursa, pappo
0.2-0.6 mm lg., irregulariter lobato vel dentato.

Typus sectionis: Pyrethrum hissaricum Krasch.

18. Sect. Trichanthemopsis Tzvel. sect. nova.—Plantae prope
basin plus minusve lignescentes, 6-25 cm alt., pilis patulis simplicibus
sparse tectae, saepe glabrae; caules parce foliati; laminae foliorum
pinnatisectae (vulgo bipinnatisectae), lobulis terminalibus angustissimis.
Calathidia solitaria; involucra late semiglobosa, 10-16 mm in diam.;
receptaculum valde convexum, semiglobosum; ligulae oblongoovales,
6-15 mm lg. Achenia 2.5-3.5 mm lg., nervis 5-10 longitudinalibus
percursa, pappo 1.2—2.5 mm lg., usque ad basin lobis 5—10 dissecto.

Typus sectionis (species unica): Pyrethrum tianschanicum Krasch.

19. P. corymbiforme Tzvel. sp. nova.—P. corymbosum var. «fol.
segmentis profundius divisis, pappo coroniforme breviore» Ldb. FI.
Ross. II, 551.

Planta 30-100 cm alt., pilis sparsis simplicibus tecta, saepe
subglabra; caules erecti, sat parce foliati, simplices (inflorescentia
exclusa); folia radicalia et caulina inferiora ad 20-25 cm lg. et 6-8 cm
It., longe petiolata, laminis ambitu oblongis vel late linearibus,
piunatisectis, segmentis pinnatifidis vel pinnatilobatis, margine plus
minusve dentatis, dentibus longe acutatis. Calathidia (1)2—6(10), in
inflorescentiam laxam corymbiforme—racemosam congesta, pedunculis
longis, ad 15-18 cm lg.; involucra 8-12 mm in diam. et 3.5—-4.5 cm
alt., glabra, phyllis margine angustissime fuscescente membranaceis,
ligulae albae, 9-15 mm lg. Achenia 2—2.5 mm lg., nervis 5-8
longitudinalibus percursa, pappo 0.3-0.5 mm lg., irregulariter lobato
et dentato.

Habitat: in pratis et fruticetis montium Alatau songoricus.

T y pus: In rupestribus Alatau ad fl. Sarchan. Enum. 1841, n°
441, Karelin et Kirillov. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad)
conservatur.

A P. corymboso (L.) Willd. pubescentia (pilis solum simplicibus),
involucris glabris, inflorescentia corymbiforme-racemosa et pappo
breviore (non 0.5—0.8 mm lg. differt.

874

832

20. P. mikeschinii Tzvel. sp. nova.

Planta 20-30 cm alt., rhizomate incrassato, caudiculos plus minusve
lignescentes caulibus floriferis et rosulis foliorum terminatos emittente;
caules erecti vel basi ascendentes, parce foliati, solum prope basin
ramosi; folia canescente-viridia, pilis accumbentibus bifurcatis (partim
simplicibus) copiose tecta, radicalia ad 10 cm lg. et 1.3 cm It., longe
petiolata (sed petiolis laminis brevioribus), basi valde dilatata, laminis
ambitu late linearibus, pinnatisectis, segmentis oblongis, plus minusve
pinnatisectis vel pinnatilobatis, lobulis terminalibus lineari-lanceolatis
ad 1.2 mm It., apice breviter mucronatis, caulina pauca et minora,
sessilia vel subsessilia. Calathidia solitaria, longe pedunculata; involu-
cra 6-8 mm in diam. et 5-7 mm alt., pilosa phyllis coriaceo-herbaceis,
exteris lanceolato-ovatis, internis oblongo-lanceolatis margine
fuscescente membranaceis exteris sesqui vel duplo longioribus; recep-
taculum subplanum; flores evoluti deficientes. Achenia matura ignota;
pappus circa 0.2 mm lg., fere ad basin irregulariter lobatus.

Habitat:. in declivitatibus lapidosis et stepposis jugi
Turkestanici.

T y pus: Declivitas borealis jugi Turkestanici, in fontibus fl. Isfara
supra pag. Voruch, alt. circa 2900 m s. m., 22 VI 1938, n° 58, G.
Mikeschin. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur.

A P. hissarico Krasch. et P. Galae M. Pop. forma et pubescentia
foliorum bene differt.

21. P. songaricum Tzvel. sp. nova.—P. pyrethroides ssp.
songaricum Krasch. in herb.

Planta 15-35 cm alt., griseo-viridia, laxiuscule tomentosa; caules
erecti parce foliati, simplices vel pauciramosi (ramis 1-3); folia radicalia
ad 8-10 cm lg. et 3 cm It., longe petiolata, laminis ambitu late linearibus
vel lineari-oblongis, tripinnatisectis, lobulis terminalibus lineari-
lanceolatis et lanceolato-oblongis, ad 1 mm It., caulina vulgo minora,
sessilia, laminis bipinnatisectis. Calathidia solitaria, rarius 2-4, longe
pedunculata; involucra 10-18 mm in diam. et 5—8 mm alt., plus minusve
tomentosa, phyllis herbosis, margine sat late atrofusce membranaceis;
corollae florum ligulatorum albae, ligulis 8-14 mm lg. Achenia 2.5—3
mm lg., nervis 6-8 longitudinalibus percursa, pappo 1-1.5 mm lg.,
vulgo ad medium (et minus) irregulariter dentato vel lobato.

Habitat: in decliviis lapidosis et saxosis supra 2000 m. s.

Typus: Alatau songoricus, systema fl. Koksu prope montem
Kuvtas, 11 VIII 1948, V. Goloskokov. In Herb. Inst. Bot. Acad. Sc.
URSS (Leningrad) conservatur.

A P. pyrethroides (Kar. et Kir.) B. Fedtsch. ex Krasch. caulibus
altioribus saepe ramosis et pappo acheniis irregulariter dentato vel
lobato (non ad basin dissecto) differt.

875

833

22. P. neglectum Tzvel. sp. nova.

Planta 12-30 cm alt., copiose sed laxiuscule albide tomentosa;
caules erecti, parce foliati, simplices vel pauciramosi (ramis 1-2); folia
radicalia ad 8-10 cm lg. et 2.5 cm It., longe petiolata (petiolis saepe
laminis subaequalibus), laminis ambitu oblongo-ovatis vel’ oblongis,
tripinnatisectis, lobulis terminalibus lanceolato-linearibus vel lanceolato-
ovatis, ad 0.6 mm It., caulina vulgo minora et minus dissecta, superiora
sessilia. Calathidia solitaria, rarius 2-3, longe pedunculata; involucra
7-12 mm in diam. et 4—5 mm alt., copiose tomentosa, phyllis herbosis
margine comparate anguste fusco vel atrofusco membranaceis; corollae
florum ligulatorum albae vel roseo-albae, ligulis 6-10 mm lg. Achenia
1.8-2.3 mm lg., nervis 6-8 longitudinalibus percursa, pappo 0.6—0.8
mm lg., usque ad basin lobis 5-8 dissecto.

Habitat: in decliviis lapidosis et saxosis montium Tjan-Schan.

T y pus: Jugum Czatkalense, montes Kuldambes prope lac. Sary-
Czilek. 17 VII 1908, V. Rovinski. In Herb. Inst. Bot. Acad. Sc. URSS
(Leningrad) conservatur.

A P. pyrethroides (Kar. et Kir.) B. Fedtsch. ex Krasch. pubescentia
albide tomentosa et involucri phyllis margine vulgo fusco et angustius
membranaceis differt.

23. P. urgutense M. Popov in Trud. Uzbeksk. Gos. Univers. Nov.
Ser. n° 27, Biol. XIV (1941) 80, Russk. diagn et in litt.

Planta perennis, 18-40 cm alt.; caules numerosi, erecti vel
ascendentes, parce foliosi, ramosi, basi tomento cingentes; folia pilosa,
ad 10 cm lg., laminis ambitu lineari-oblongis, bi-vel tri-pinnatisectis,
lobulus terminalibus ad 1 mm lg. Calathidia (1)2-6(8), in
inflorescentiam laxam corymbiforme—racemosam congesta, homogama;
involucra 7-12 mm in diam.; receptaculum convexum, sparse pilosum
vel glabrum; flores omnes tubulosi, corollis flavis, 2.2-2.8 mm lg.,
superne copiose pilosis. Achenia 2—2.4 mm lg., 5—6-costata, pappo
0.3-0.6 mm lg., irregulariter acute dentato.

Habitat: in decliviis saxosis et lapidosis jugi Zeravschanici.

T y pus: Jugum Zeravschanicum, infra trajectum Kirk-arcza prope
pag. Farab, 18 VII 1934, K. Zakirov. In Herb. Univers. Asiae mediae
(Taschkent) conservatur.

A specie proxima Tanaceto Krascheninnikovii Nevski praesertim
corollis florum superne copiose pilosis differt.

876

834
24. SPATHIPAPPUS Tzvel gen. nov.

Calathidia vulgo solitaria (sed in planta saepe numerosa)
heterogama; involucra late semiglobosa, 10-18 mm in diametro et 4—
6 mm alt., phyllis herbaceis, imbricatis, 3—4-serialibus, exteris
lanceolatis, internis late linearibus sesqui vel duplo longioribus;
receptaculum leviter convexum, glabrum; flores marginales ligulati,
uniseriales, sat numerosi, gynoeciati, sed semper steriles; corollae albae
cum germine epapposo vel subepapposo concrescentes, tubo valde
compresso, 1.2—1.6 mm lg., ligulis oblongo-linearibus 7-10 mm lg.;
flores disci numerosi hermaphroditi, tubulosi, flavi, tubo corollae
angusto et sat longo, superne sensim dilatato; antherarum appendices
apicales lanceolato-ovatae, obtusiusculae; rami stylorum lineares,
truncati. Achenia florum disci conformia, glabra, subteretia, 3-3.5 mm
lg. et circa 0.5-0.6 mm It., costis longitudinalibus 5-6 leviter
prominentibus percursa, pappo integro secundo auriculiformi 1.5-2.4
mm lg. coronata. Plantae perennes; caules numerosi, prope basin
lignescentes, calathidiorum pedunculis longis terminati; folia alterna;
laminis bipinnatifidis.

Typus generis: Spathipappus Griffithii (Clarke) Tzvel.

A genere Pyrethrum Zinn pappo secundo auriculiformi et floribus
ligulatis sterilibus differt.

WALDHEIMIA Kar. et Kir.

25. W. transalaica Tzvel. sp. nova.

Planta perennis, ad 10 cm alt., laxe griseo-tomentosa; caules
numerosi, simplices, ascendentes vel erecti, sed prope basin vulgo
procumbentes; folia ad 2.5—3 cm lg. et 1(1.5) cm It., longe petiolata,
laminis ambitu subovatis, bipinnatifidis rarius pinnatifidis, segmentis e
quovis latere 2—5, lobulis terminalibus lanceolatis vel ovatis, ad 1 mm
It., apice mucronatis. Calathidia solitaria, heterogama; involucra 1.2—
2 cm in diam. et 4-6 mm alt.; flores ligulati 6-20, albi, ligulis 7-12
mm lg. et 2-4 mm It. Achenia 3-4 mm lg., usque ad basin dense
pilosa, pappo 5-7 mm lg. (tab. XI, fig. 1).

Habitat: in locis glareosis et lapidosis jugi Transalaici supra
3000 m s. m.

Typus: Kirghisia, jugum Transalaicum, locus glareosus in
fontibus fl. Kapstur, 11 VII 1933, n° 867, I. Tysczenko. In Herb. Inst.
Bot. Acad. Sc. URSS (Leningrad) conservatur.

A specie proxima W. tomentosa (Decne.) Rgl. forma foliorum et
acheniisusque ad basin dense pilosis differt.

835
CANCRINIA Kar. et Kir. emend. Tzvel.

26. Sect. Polychrysum Tzvel. sect. nov.—Calathidia numerosissima
(plus 40), parva, in inflorescentia corymbiformi ad 25 cm in diametro
dense congesta; involucra semiglobosa, 3—4 mm in diametro;
receptaculum convexum, sed margine subplanum, glabrum. Achenia
1.3-1.8 mm lg. sat dense, sed breviter glandulose pilosa, pappo 0.4—
0.7 mm lg. fere ad basin dissecto coronata. Plantae perennes,
monocarpicae, 30-150 cm alt.

Typus sectionis (et species unica): Cancrinia tadshikorum (Kudr.)
Tzvel.

27. Sect. Tanacetopsis Tzvel. sect. nova.—Calathidia vulgo
numerosa (ad 40), in inflorescentia paniculate-corymbiformi vel
corymbiformi laxe, rarius dense congesta; involucra late semiglobosa,
6-14 mm in diametro; receptaculum convexum, glabrum, rarius sparse
pilosum. Achenia 1-2.4 mm lg., glabra, pappo 0.1—-1.5 mm lg. plus
dissecto vel dentato coronata. Plantae perennes, polycarpicae, (8)20-
70 cm alt.

Typus sectionis: Cancrinia mucronata (Rgl. et Schmalh.) Tzvel.

28. Sect. Matricarioides Tzvel. sect. nova.—Calathidia solitaria
(sed in planta saepe numerosa); involucra late semiglobosa, 7-12 mm
in diametro; receptaculum valde convexum, subconicum, glabrum.
Achenia 1.8—2.2 mm lg., glabra, pappo 0.8-1 mm lg., fere ad medium
irregulariter et obtuse lobato coronata. Plantae biennes vel perennes
(?), monocarpicae, 5—20 cm alt.

Typus sectionis (et species unica): Cancrinia discoidea (Ldb.)
Poljak.

29. CANCRINIELLA Tzvel. gen. nov.

Calathidia solitaria (sed in planta saepe numerosa), homogama,
involucra semiglobosa, 6-10 mm in diametro et 3.5—-6 mm alt., phyllis
herbaceis, bi-vel uniserialibus, subaequilongis, lanceolatis vel oblongo-
lanceolatis; receptaculum valde convexum, parce et breviter pilosum;
flores disci tubulosi, flavi, tubo corollae parum et sensim dilatato,;
antherarum appendices apicales lanceolato-ovatae, obtusae; rami
stylorum lineares, truncati. Achenia conformia, glabra, teretia, 2.4—2.8
mm lg. et 0.6—0.7 mm It., costis longitudinalibus 10 valde prominentibus
percursa, pappo 0.4-0.6 mm lg., ad (vel fere ad) basin lobis 6-10
magrine irregulariter et obtuse dentatis dissecto coronata. Plantae

877

836

perennes, herbaceae (sed prope basin sublignescentes), humiles, copiose
et longe pilosae; inrovationes sat numerosae valde abbreviatae et dense
congestae, rosulis foliorum et calathidiis in pedunculis longis aphyllis
vel subaphyllis terminatae.

Typus generis (species unica): Cancriniella Krascheninnikovii
(Rubtz.) Tzvel.

A genere proximo Cancrinia Kar. et Kir. habitu singulari, involucri
phyllis subaequilongis et acheniorum costis 10 valde prominentibus
bene differt.

TRICHANTHEMIS Rgl. et Schmalh. emend. Tzvel.

30. Sect. Subbulbosa Tzvel. sect. nova.—Involucra 12—20 mm in
diametro, phyllis internis exteris sesqui vel duplo longioribus;
receptaculum valde convexum, dense pilosum; flores ligulati numerosi,
flavi; corollae florum omnium extus pilosae. Achenia 2.8—3.6 mm lg.,
pappo 3-4.2 mm lg. ad basin dissecto coronata. Plantae herbaceae,
caulibus basi subbulbose incrassatis.

Typus sectionis (species unica): Trichanthemis radiata Krasch. et
Vved.

31. Sect. Pyrethroides Tzvel. sect. nova.—Involucra 9-12 mm in
diametro, phyllis internis exteris duplo vel triplo longioribus;
receptaculum convexum, glabrum vel subglabrum; flores ligulati
numerosi albi vel deficientes; corollae florum semper glabrae. Achenia
2-3 mm lg., pappo 1.5-2 mm lg. ad basin dissecto coronata. Plantae
caulibus prope basin valde lignescentibus.

Typus sectionis: Trichanthemis Litwinowii (Krasch.) Tzvel.

32. XYLANTHEMUM Tzvel. gen. nov.

Calathidia in apicibus ramorum solitaria, longe pedunculata,
homogama; involucra semiglobosa vel breviter cylindracea, 4-8 mm in
diam. et 4.5-8 mm alt., phyllis coriaceo-herbaceis, imbricatis, 4—5-
serialibus, exteris ovatis vel late lanceolatis, apice obtusiusculis, margine
angustissime hyalinis, internis late linearibus, vulgo triplo-quadruplo
longioribus, margine late hyalinis; receptaculum planum vel subplanum,
glabrum; flores disci tubulosi, hermaphroditi, flavi, corollis, 2.6—3.2
mm lg., tubo corollae longo et angusto; antherarum appendices apicales
lanceolato-ovatae, obtusiusculae; rami stylorum lineares, truncati.
Achenia conformia, glabra, subteretia, 1.5-3.5 mm lg. et circa 0.6—0.7

878

837

mm It., costis longitudinalibus 5-6 percursa, pappo secundo,
auriculiformi 1.2-2.2 mm lg., integro vel usque ad basin dissecto
coronata. Plantae perennes; caules prope basin lignescentes,
ramosissimi, superne simplices, calathidiorum pedunculis longis
terminatis; folia alterna, basalia, cito emortua, caulina sat numerosa,
parva laminis pinnatisectis.

Typus generis: Xylanthemum Fischerae (Aitch. et Hemsl.) Tzvel.
A generibus proximis Tanacetum L., Cancrinia Kar. et Kir. et
Trichanthemis Rg}. et Schmalh. differt: a primo calathidiis homogamis,
receptaculo plano, acheniorum pappo auriculiformi et habitu; a
secundo—pappo auriculiformi, receptaculo plano, involucris
angustioribus phyllis internis exteris multo longioribus et caulibus basi
valde lignescentibus, a tertio-acheniis glabris, pappo auriculiformi et
habitu.

33. PPEUDOHANDELIA Tzvel. gen. nov.

Calathidia 20-100, breviter pedunculata (pedunculis ad 1.5 cm
lg.), in corymbum umbelliformem dense congesta, homogama; involucra
semiglobosa, 6-9 mm in diam. et 3-5 mm alt., phyllis herbaceis,
subaequilongis, exteris et mebiis oblongis margine late hyalinis, internis
angustioribus margine angustissime hyalinis bracteiformibus;
receptaculum valde convexum, obtuse, conicum, glabrum; flores omnes
tubulosi, hermaphroditi; corollae flavae, 1.8-2.5 mm lg., tubo longo et
angustissimo, breviter glanduloso-piloso, in parte superiore subito
campanulato-dilatato, quinquedentatae, dentibus lanceolato-
triangularibus patentibus tubo 7-—8-plo breviorbus; antherarum
appendices apicales ovato-lanceolatae, obtusiusculae; rami stylorum
lineares, truncati. Achenia conformia, subteretia, arcuate curvata, sat
copiose et breviter glanduloso-pilosa, 1.8—2.4 mm lg. et circa 0.3 mm
It., mervis longitudinalibus 4-5 percursa, epapposa. Plantae
monocarpicae biennes vel perennes, laxe arachnoideo-tomentosae;
caules erecti, simplices (inflorescentia exclusa) 20-100 cm alt.; folia
alterna, laminis bipinnatisectis.

Typus generis (species unica): Pseudohandelia umbellifera (Boiss.)
Tzvel.

A generibus proximis Tanacetum L. et Cancrinia Kar. et Kir.
differt: a primo-calathidiis homogamis, acheniis epapposis glanduloso-
pilosis et forma corollarum florum, a secundo-acheniis epapposis
glanduloso-pilosis, receptaculo plano, phyllis involucri subaequilongis
et habitu.

879

838
TANACETUM L.

34. Sect. Asterotricha Tzvel. sect. nova.—Plantae herbaceae, pilis
accumbentibus stellatis dense tectae, 5-30 cm alt.; caules parce foliati.
Calathidia solitaria, heterogama; involucra 3.5-10 mm in diam., phyllis
internis externis duplo vel triplo longioribus; receptaculum subplanum;
flores marginales ligulati flavi: flores disci tubulosi flavi. Achenia 2—
3.2 mm lg., nervis 5—6 longitudinalibus percursa, pappo 0.2—0.3 mm
lg., irregulariter obtuse lobato.

Typus sectionis: Tanacetum Walteri (Winkl.) Tzvel.

35. T. ulutavicum Tzvel. sp. nova.

Planta 12—25 cm alta, pilis bifurcatis (et partim simplicibus) sparse
tecta; caules erecti vel basi ascendentes, parce foliati, simplices vel
supra medium plus minusve ramosi; folia sparse pilosa, radicalia ad 8
cm lg. et 1.3 cm It., longe pedunculata, laminis ambitu late linearibus
pinnatisectis, segmentis pinnatilobatis vel pinnatifidis, lobulis
terminalibus ad 1.5 mm It., breviter acutatis. Calathidia solitaria vel 3-
5, longe pedunculata (pedunculis ad 8-10 cm lg.); involucra 6-11 mm
in diam. et 4.5-6 mm alt., subglabra vel sparse pilosa, phyllis
coriaceoherbaceis, exteris lanceolato-ovatis, internis lanceolato-
oblongae sesqui vel duplo longioribus, omnibus margine sat late
membranaceis; flores marginales subligulati vel tubulosi, disci tubulosi.
Achenia 1.8—2.5 mm lg. et circa 0.6 mm It., nervis 6—9 longitudinalibus
percursa, pappo circa 0.3 mm lg. margine irregulariter dentato.

Habitat: in decliviis lapidosis et saxosis montium Ulutau.

Ty pus: Prov. Karaganda, distr. Karsakpaj, in declivitate montis
Kazantau non procul a pag. Ulutavskoe, 12 VII 1929, n° 298, I.
Schipezinski. In Herb. Inst. Bot. Akad. Sc. URSS (Leningrad)
conservatur.

A specei proxima T. achilleifolia (M.B.) Sch. Bip. pubescentia
sparsiore, calathidiis majoribus, foliis minus dissectis et floribus
marginalibus tubulosis vel subligulatis differt.

T. canescens ssp. erivanense Tzvel. ssp. nova.—A typo segmentis
foliorum minus numerosis et minus dissectis apice acutiusculis differt.

Typus subspeciei: In viciniis urb. Erevan prope pag. Dzhirvicz. V
1922, A. Grossheim.

T. myriophyllum ssp. borshomicum Tzvel. ssp. nova.—A typo
pedunculis in parte superiore et involucris multo laxius tomentosis
differt.

Typus subspeciei: Distr. Borshomi, in declivitate saxosa inter st.
Sakogavi et pag. Sakogavi, 14 VI 1918, V. Koslovski.

880

839
DENDRANTHEMA (DC.) Des Moul. emend. Tzvel.

36. D. sichotense Tzvel. sp. nova.—Chrysanthemum sibiricum var.
alpinum Nakai in Tokyo Bot. Mag. XXXI (1917) 109.—Ch. Zawadskii
var. alpinum (Nakai) Kitam. in Acta Phyt. et Geob. VII (1938) 210.

Planta perennis, 10-25 cm alta; caules erecti, foliosi, simplices
vel pauciramosi (ramis 1-2); folia laxe tomentosa griseo-viridia,
radicalia et caulina infreriora numerosa, ad 4 cm lg. et 2 cm lt., longe
petiolata; laminae ambitu a reniformibus ad subrotundas, bipinnatifidae,
segmentis e quovis latere 2-3, lobulis terminalibus anguste linearibus
(ad 1 mm It.), apice longe et sensim acutatis; folia caulina media et
superiora minora, breviter petiolata vel sessilia, minus dissecta.
Calathidia vulgo solitaria, rarius 2-3, sat magna; involucra 12-18 mm
in diametro et 4.5-6 mm alt., extus lanuginoso-tomentosa, phyllis
margine late hyalinis et nigrofuscis; flores ligulati rosei, ligulis 12-24
mm lg.; flores tubulosi flavi. Achenia 1.5—2.5 mm lg., pappo nullo
(tab. XVI, fig. 1).

Hab itat: in declivitatibus lapidosis et rupibus montium Sichote-
Alinj Oriens Extremus) supra 1200 m s. m.

T y pus: Oriens Extremus. Montes Sichote-Alinj (pars australis)
in monte Snezhnaja, alt. circa 1500 m s. m. 15 VII 1930, n° 767, I.
Schischkin. An Herb. Inst. Bot. Acad. Sc. URSS (Leningrad)
conservatur.

A specie proxima D. Maximowiczii (Kom.) Tzvel. pubescentia,
forma foliorum radicalium et phyllis involucri margine late nigrofuscis
bene differt

37. Sect. Arctanthemum Tzvel. sect. nova.—Antherarum
appendices apicales oblongae, apice late rotundatae; receptaculum
glabrum. Achenia sine cellulis muciferis. Plantae glabrae vel subglabrae,
rhizomate incrassato, laminis folio rum crassiusculis plus minusve
lobatis vel dissectis.

Typus sectionis: Dendranthema arcticum (L.) Tzvel.

38. Sect. Haplophylla Tzvel. sect. nova.—Antherarum appendices
apica les oblongae, apice late rotundatae; receptaculum parce pilosum.
Achenia sine cellulis muciferis. Plantae humillimae, caulibus
subaphyllis, foliis radicalibus numerosis integerrimis anguste linearibus.

Typus sectionis (species unica): Dendranthema integrifolium
(Richards.) Tzvel.

_

840
BRACHANTHEMUM DC.

39. Sect. Dendranthemopsis Tzvel. sect. nova.—Brachanthemum
ser. Pyrethroidea Krasch. Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR
XI (1949) 196, diagn. russk.—Calathidiorum involucra semiglobosa
phyllis internis exteris. duplo-triplo longioribus; receptaculum sub-
planum, breviter et parce pilosum; corollae florum ligulatorum albae,
ligulis 6-8 mm lg.; antherarum appendices apicales ovato-lanceolatae.

Typus sectionis (et species unica): Brachanthemum Baranovii
(Krasch. et Poljak.) Krasch.

AJANIA Poljak.

40. A. trilobata Poljak. sp. nova in litt.

Haec species A. Scharnhorstii (Rgl. et Schmalh.) Tzvel. proxima,
sed caulibus validioribus (5—15 cm altis), calathidiis magis numerosis
(vulgo 3-10 in corymbo) et minoribus (vulgo 5—7 mm in diametro)
differt.

Habitat: in decliviis glareosis et lapidosis supra 2500 m s. m.
montium Tjanj-Schanj.

T y pus: Tjanj-Schanj centralis, in systemate fluminis Sary-Dzhas,
in decliviis infra Nansen-montem, alt. 2950 m s. m., 29 VII 1955, M.
Grudzinski. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad)
conservatur.

41. TURANIPHYTUM Poljak. gen. nov.

Calathidia heterogama in numero 6-18 in glomerulos subglobosos
dense congesta, rarius partim solitaria; involucra oligophylla, phyllis
subaequilongis, internis carinatis scaphoideis; receptaculum convexum
semisphaericum, glabrum, flosculi disci staminati, pistillo rudimentali,
corolla tubulosa, superne sat sensim dilatata, quinquedentata, apice
longe et rigide pilosa, plus minusve roseola; antherae appendices
superiores oblongae apice attenuatae; pistillum rudimentale apice
integrum conicum; flosculi radii pistillati, fertiles, in phyllis involucri
scaphoideis dispositi, corolla parva, late tubulosa, apice inaequaliter
crenata; pistillum apice bifidum, ramis linearibus divergentibus; achenia
ovata, brunnea, epapposa. Plantae perennes sat longe et molliter pilosae;
folia pinnatisecta; glomeruli et calathidia solitaria in inflorescentia
spiciformi interrupta congesti.

841

Typus generis: Turaniphytum eranthemum (Bge.) Poljak. comb.
nova-Artemisia eranthema Bge.

Affinitas: Hoc genus a genere Artemisia L. flosculis radii in
involucri phyllis scaphoideis dispositis et calathidiis vulgo in glomerulos
dense congestos differt.

42. T. kopetdaghense Poljak. sp. nova.

Planta perennis, incano-viridis. pilis sat longis mollibus dense tecta;
radix crassa, sublignosa; surculi abbreviati, caespites humiles compactos
formantes; caules plus minusve numerosi, 15-25 cm alt., erecti vel
basi ascendentes, semper simplices, parce foliosi; folia radicalia longe
petiolata, 3.5—6 cm lg., 1.2—1.5 cm It., ambitu oblonga, bipinnatisecta,
lobis primariis 2—3-jugis, remotis; lobulis 3-5 mm lg. et 0.4-0.6 mm
It., linearibus, acuminatis; folia caulina minora, sessilia vel subsessilia,
pinnatisecta; folia suprema parva, linearia. Inflorescentia spicata,
interrupta; calathidia sessilia, ovata vel subglobosa, 2—2.5 mm It., in
inflorescentiae parte superiore plus minusve conferta,; involucrum
oligophyllum; flosculi disci in numero 16-18, radii in numero 8-9 in
involucri phyllis scaphoideis dispositi. Achenia 1.2-1.4 mm lg. et circa
0.6 mm It.

Typus: speciei. Turcomania, distr. Aschchabad, montes
Kopetdaghenses, 2 km versus orient. a pag. Sybir, 2600-2400 m s. m.,
24 VIII 1947, V. Nikitina. In Herb. Inst. bot. Ac. Sc. URSS (Leningrad)
conservatur.

Affinitas: A. T. eranthemo (Bge.) Poljak. calathidiis multo
majoribus et in glomerulos subglobosos non congestis, floribus 20-30
(non 2-10) constantibus differt.

SENECIO L.

43. S. ferganensis Schischk. sp. nov. (Sect. Jacobaea DC. Prodr.
VI (1837) 348).

Biennis; caulis erectus, superne ramosus, ramis oblique sursum
directis unacum foliis araneoso-flosculosus, 60-75 cm altus; folia
bipinnatim secta, radicalia et caulina infima cito marcescentia caulina
media numerosa anguste ovata, sessilia basi dilatata et auriculis vix
evolutis munita, 6-15 cm longa et 1.5-2 cm lata. Inflorescentia
corymbosa, ramis oblique sursum directis; calathidia numerosa in
numero 15-30, involucrum late campanulatum 5 mm longum et 10 mm
latum, foliola involucri externa 3—5, linearia, interioribus adpressa vel
leviter patentia, duplo triplove interioribus breviora, interiora anguste
ovata, acutiuscula vix araneosa vel glaberrima; ligulae luteae involucro
duplo longiores, achenia glabra ad 2 mm longa. Fl. VI-VII.

882

842

Typus: Asia Media. Jugum Alaicum, in faucibus fluminis
Czigirczik, in clivo montis supra ripam. Fl. 11 VII 1930 n° 295. S.
Juzepcezuk; in Herb. Inst. bot. Ac. Sc. URSS conservatur.

Affinitas: Nostra species nova appropiquat ad S.
borysthenicum Andrz sed foliis bipinnatim (nec tripinnatim) sectis et
acheniis disci glaberrimis (ne pilosis) sa discrepat.

44. S. massagetovii Schischk. sp. nov. (Sect. Incani DC. Prodr. V
(1837) 355).

Radix 0.5 cm crassa; caulis solitarius tomento araneoso leviter
detergibii tectus, costatus, superne ramosus, 50-75 cm altus; folia
radicalia et caulina infima petiolata, petioli dilatati laminae aequales
vel ea longiores, lamina ovato oblonga 6-20 cm long. et 3-10 cm lata,
supra fuscoviridia, subglabra, subtu breviter et dense araneosa, lyrato-
pinnata, lobis 2-3, distantibus, deorsum deminutis, inaequaliter dentatis;
lobus supremus ceteris major; folia caulina media radicalibus similia, sed
sessilia et auriculis semiamplexicaulibus instructa. Calathidia numerosa (10—
40), inflorescentiam laxam corymbosao-paniculatam formantia
pedunculi calathidiis aequales vel eis multo longiores; involucrum
flocoso-araneosum, 6-10 mm longum et 10 mm Iat., foliola involucri
externa anguste linearia, in numero 1-5, internis duplo breviora
(interdum nulla), foliola interna oblongo-linearia, dorso carinata, ligulae
luteae in numero 13-15, 10 mm longae, 2-3 mm latae, achenia
cylindrica glabra, vix costata, 4-4.5 mm longa. Fl. VI-VII.

T y pus: Grusinskaja SSR, In faucibus fluminis Borzhomka prope
stationem viae ferreae Sakoczavi, in saxosis 25 VI 1934 P. Massagetov;
in Herb. Inst. bot. Ac. Sc. URSS conservatur.

Affinitas: Nostra. species nova valde affinis S$. Lorentii Hochst.
et §. Lipskyi Lom., sed ab ambabus involucris 10 mm latis (nec 12-
15 mm), calathidiis numerosis usque ad 40 (nec 10-25) distinguitir.

45. S. iljinii Schischk. sp. nova. (Sect. Pseudooliganthi Sof. in
Izv. Ak. Nauk Azerb. CCP (1957) 88).

Ut videtur, perennis, radix deficiens; caulis erectus, intus solidus,
pilis albis, brevibus, srispatis tectus, 40-50 cm altus, dense foliosus;
folia ovata, sessilia vel petiolis brevibus instructa, acuta, marginibus
grandidentata, dentibus vulgo obtusis, supra fusco-viridia, infra
pallidiora, subglabra vel vix araneosa, 8-12 cm longa et 3-5 cm lata.
Inflorescentia paniculata, polycephala pedunculi densiuscule albo-
tomentosi; involucrum 5 mm longum ac latum, foliola involucri externa
in numero 3-4, linearia, internis vix longiora et eis appressa, foliola
interna ovoidea, apice acutiuscula, in numero 8-10, dorso breviter
crispule pubescentia, marginibus ciliata; ligulae 8, luteae, involucro

883

843

triplo longiores; achenia juvenalia glaberrima, 3-4 mm longa. FI. VIII.
Ty pus: Kazachskaja SSR. Districtus Dzharkend apud flum.
Tyschkan. Fl. 8 VIII 1910—A. Michelson; in herb. Inst. bot. Ac. Sc.
SSSR conservatur.
Affinitas: Nostra species affinis est $. nemorensi L. sed foliis
marginibus grosse dentatis, dentibus inaequalibus, obtusis (nec dentibus
aequalibus acutis), pedunculis dense albo tomentosis (nec subglabris).

46. Sect. 7. Extremiorientales Schischk. sect. nov.—Plantae
perennes, 1-2 m alt., foliis digitatis vel integerrimis, basi saepe auriculis
magnis instructis; calathidia numerosa parva, involucrum ad 5 mm
longum ac latum; achenia glabra 3-S mm longa.

Duae species extremiorientales: §. cannabifolius Less. et S.
Litvinovii Schischk. Typus sectionis: S. cannabifolius Less.

47. Sect. 8. Carniolici Schischk. sect. nov.—Folia radicalia diu
persistentia, pinnatim incisa, lobis late linearibus, foliola interna
involucri apice macula nigra instructa. Ligulae luteae 2-4, achenia
glabra, pappus sordide fuscus.

Sect. monotypica: S. carniolicus Willd.

48. Sect. 9. Montani Schischk. sect. nov.—Folia radicalia petiolata,
petioli laminae aequales vel ea duplo triplove longiores ovoidea,
dentibus majusculis inaequalibus vel folia pinnatim secta; foliola
involucri externa linearia in numero 2-8, ligulae flavae in numero 10—
12; achenia glabra.

Duae species: S. tianschanicus Rg]. et Schmalh., S. Renardii Winkl.
montibus Asiae Mediae propriae. Typus sectionis: S. tianschanicus
Rgl. et Schmalh.

49. Sect. 10. Scaposi (Hoffm.) Schischk. ex Krylov Flora Zap.
Sib. XI (1949) 2890.—Scaposi Hoffm. in Engl. Pflanzenf. 4.5 (1889)
297 part.

Plantae perennes, humiles, semper monocephalae, folia pinnata
vel lyrata; tota planta glaberrima.

Duae species: S. residifolius Less. et S. whanithaaie Herb. Typus
sectionis: S$. resedifolius Less.

50. Sect. 11. Karelinoidei O. et B. Fedtsch.
IV (1911) 220 nom. nud.—Plantae perennes, folia subintegerrima ovata
vel ovato-lanceolata; foliola involucri interna in numero 8, externa |-
2, vel interdum deficientia, ligulae luteae in numero 3-5.

884

844
Sectio monotypica: S. karelinoides Winkl. Asiae Mediae propria.

51. S. amurensis Schischk. sp. nov. (Sect. Tephroseris (Rchb.)
DC. Prodr. VI (1837) 359).

Perennis; radix fibrillosa; caules solitarii vel 2—3, erecti, intus solidi,
arachnoideo-albo-pubescentes (10)20—60 cm alti; folia radicalia ovata,
anguste ovata vel interdum lanceolata, obtusa vel acutiuscula, basi in
petiolum latum angustata, supra et subtus araneoso-pubescentia,
marginibus dentibus minimis nigris distantibus instructa; lamina 1.5-
8 cm longa et 1-5 cm lata; caulina non numerosa, ad apicem caulis
gradatim deminuentia, infima caulina radicalibus similia, breviter
petiolata, cetera sessilia basi amplexicaulia, lanceolata, acuta 4-8 cm
longa et 1—1.5 cm lata, suprema lanceolato-linearia minora. Calathidia
in numero 3-10 inflorescentiam umbelliformem formantia, pedunculi
araneoso-pubescentes, vulgo calathidiis multo longiores, basi umbellae
foliis linearibus parvis munitis, foliola involucri unifaria, involucrum
late campanulatum, 6-7 mm longum et 10 mm latum; ligulae pallide
flavae, oblongae, in numero 12, 10 mm longae et 2—2.5 mm latae;
ovarium et fructus pubescentes. Fl. V—VI.

T y pus: Sibiria orientalis, in pratis siccis silvaticis prope oppidum
Blagovesczensk V—VI 1903 fl. F. Karo.

Affinitas: Nostra species nova valde affinis S. Karelinii sed
differt ab eo foliis dentibus vulgo nigris minimis distantibus instructis,
nec dentibus deficientibus.

52. S. lenensis Schischk. sp. nova (Sect. Tephroseris (Rchb.) DC.
Prodr. VI (1837) 359).

Rhizoma adscendens, 8-10 mm crassum, radicibus numerosis
tenuibus tectum; caulis glaber vel floccoso pubescens, in dimid.o
inferiore aequifoliosas, in dimidio superiore fere efoliatus, intus cavus,
tenuiter costatus, 25-50 cm altus; folia radicalia petiolata, petiolis
lamina brevioribus vel vix longioribus, lamina ovata vel lanceolato-
ovata, 4-6 cm longa et 1.5—3 cm lata, apice obtusa vel acuta, marginibus
dentibus parvis nigris distantibus instructa, glabra vel floccoso-
pubescens; folia caulina inferiora radicalibus angustiora, rite lanceolata,
acuta vel interdum acuminata. sessilia rarissime + longe petiolata, 4—
12 cm longa et 0.5-2 cm lata superne fere linearia, parvula.
Inflorescentia umbellata, calathidiis 3-7 plus minusve longe
pedunculatis; involucrum unifarium, 5-8 mm longum et 10-15 mm
lata, foliola leviter pilis brevibus tecta, lanceolato-linearia, apice tenuiter
acuminata et hic nigrescentia; ligulae luteae, interdum durantiacae 12
mm longae et 3 mm latae, in numero ad 12; achenia 2—3.5 mm longa,
pilis brevibus mollibus pubescentia. Fl. VII.

885

845

Ty pus: In valle fluminis Lenae prope Bulun. fl. 3 VII 1914. V.
Kaschkarov.

Affinitas: Nostra species nova S. lenensis Schischk. affinis
est S. Sukaczevii Schischk. sed foliolis involucri apice maculis
nigriusculis munitis (nec deficientibus) et seminibus hirsutis (nec
glabris) differt.

53. S. aquilonaris Schischk. sp. nov. (Sect. Tephroseris (Rchb.)
DC. Prodr. VI (1837)359).

Perennis, rhizoma sat longum, 3-8 mm crassum, radicibus longis,
tenuibus, non multis obsitum; caules erecti vel vix flexuosi, simplices,
pauci vel solitarii, vulgo violaceo-colorati, costati, leviter araneosi vel
subglabri, 20-30 cm alti; folia radicalia petiolata, petiolis dilatatis,
lamina brevioribus vel longioribus, lamina ovata vel interdum oblonga,
apice obtusa vel acutiuscula, marginibus dentata, dentibus late
triangularibus, brevibus, acutis, non multis, rarius integerirma, supra et
subtus glabra, 2-7 cm longa et 1—2.5 cm lata; folia caulina in numero
2-4, sessilia, amplexicaulia, lanceolata marginibus integerrima, glabra,
3-6 cm longa et 0.8—1.2 cm lata, suprema minora. Calathidia in apice
caulium in numero 2-5 raro solitaria, pedunculi breves dense araneoso-
pubescentes; involucrum 10-12 mm longum et 12-15 mm latum,
unifarium, foliola involucri lanceolato-linearia, fusco-purpurea, tomento
reticulato griseolo tecta; ligulae in numero 10-12, luteae, involucro
1%-plo longiores, 3—4 mm latae, non raro ligulae deficientes; achenia
cylindrica, glaberrima, 3-5 mm longa, pappus achenio duplo triplove
longior. Fl. VI-VII.

T y pus: Ad litus maris orientali-sibirici. In valle amnis Medveshja,
prope ostium. Fl. 8 VIII 1950. G. Nepli.

Affinitas: Nostra species affinis S$. Sukaczevii Schischk. sed
involucro tomento griseolo reticulatim tecto (nec glabro vel vix
araneoso) statim dignoscitur.

54. S. igoschinae Schischk. sp. nov. (Sect. Tephroseris (Rchb.
DC. Prodr. VI (1837) 359.

Perennis, rhizoma horisontale, longum, caulis glaber, 30-40 cm
alt.; folia radicalia in numero 2-3 et caulina inferiora lamina duplo
triplove longiora, lamina anguste ovata, 2—5 cm long. et 0.7-1.5 cm
lata, apice rotundata, basi cuneatim angustata, glabra, marginibus
breviter dentata vel integerrima, folia caulina media lanceolata,
acutiuscula basi sensim in petiolum angustata, laminae aequale, folia
suprema linearia, integra. Calathidia in numero 5 inflorescentiam
umbelliformem formantia; involucrum 5 mm longum et 10 mm latum,
unifarium, foliola involucri linearia subglabra vel vix pubescentia;

886

846

ligulae luteae, involucro duplo longiores, in numero 15; achenia
cylindrica, ad 4 mm longa, glabra vel interdum puberula. Fl. VI—VII.

Typus: Montes uralenses australes, prope pag. Machmutovo
non procul a flumine Belaja in pratis montanis. Fl. 23 VI 1957. K.
Igoschina.

Affinitas: Valde affinis S$. Reverdattoi, sed foliolis involucri
viridibus (nec nigro-violaceis), ligulis vitellinis (nec aurantiacis)
discrepat.

55. S. tichomirovii Schischk. sp. nov. (Sect. Tephroseris (Rchb.)
DC. Prodr. VI (1837) 359).

Perennis; rhizoma abbreviatum, ascendens; caules solitarii, erecti,
crassiusculi, 4-6 mm in diametro, 10-15 cm alti, a basi usque ad
dimidium glabri, superne, pilis hepaticis araneoso-tomentosi, folia
radicalia 4—6, longe petiolata, petiolis lamina duplo triplove longioribus,
lamina anguste ovata 3—4 cm longa et 0.5—1.5 cm lata, glabra, pallidi-
viridia, apice acutiuscula, marginibus dentata, dentibus crassiusculis
callosis distantibus; folia caulina radicalibus similia, sed sessilia et
amplexicaulia, glabra, folia suprema minora et ut caulis dense
tomentosa. Calathidia in apice caulis solitaria; involucrum 1.5 cm
longum et 2.5 cm latum, foliola involucri uniserialia, ut caulis, dense
villoso-tomentosa, pili fusco-flavius-culo-hepatici, ligulae fusco-flavae
lineares in numero 20-25, involucro duplo longiores. Fl. VII—VIII.

T y pus: Peninsula Czukotka, in tundra fl. 20 VIII 1958. Leg. B.
Tichomirov.

Affinitas: Affinis est S. atripurpureo (Ldb.) B. Fedtsch. sed
ligulis sordide purpureis (nec luteis vel aurantiacis) involucro 1.5 cm
longo et 2.5 cm lato, caule superne pilis hepaticis araneoso-tomentoso
nec pilis albis vel atri-purpureis statim dignoscitur.

LIGULARIA Cass

56. Subgenus Ligularia—Syn. cfr. supra 797.—

Rhizoma abbreviatum incrassatum radices numerosas chordoideas
emittens. Calathidia parva, mediocria vel magna, involucro quinque—
polyphyllo, phyllis saepissime inter se liberis, sed raro (subsect.
Stereochaete series Schmidtianae) in cyathium apice paucidentatum
connatis. Florum disci styli ramuli apice angustati aut appendice brevi,
conica vel subplana, apice rotundata vel acutata acuti; stigmata (proprio
sensu) ramulorum totam faciem superiorem occupantia aut strias
marginales continuas, latiusculas, prominulas, usque ad appendicis basin
extensas formantia; pili ad paginam inferiorem infra medium vel saepe

887

847

ad basin, rarius sub appendice tantum siti. Antheropodium filamento
paulo vel distincte latius. Pappi setae achenio saepissime subaequilongae
tenues, scabrae vel brevissime barbellatae, aut raro (Sect. Senecillis)
breviusculae vel brevissimae, achenio multo breviores, manifeste
barbellatae.

Typus subgeneris—L. sibirica (L.) Cass.

57. L. trichocephala Pojark. sp. nova.—L. Hodgsonii var.
sachalinensis Nakai in Tokyo. Bot. Mag. XXIV (1910) 264 (non L.
sachalinensis Nakai (1944.—L. calthaefolia auct. non Maxim. (1870):
Sugaw. Ill. Fl. Saghal. IV (1940) 1857.

Caulis superne, rarius ad basin pilis crassiusculis fuscis et praeterea
albis arachnoideis vestitus. Folia plerumque rotundato-vel triangulari—
cordata, rarius subhastata vel subreniformia, apice saepe mucronulata.
Inflorescentia subcorymbosa vel racemosa, pedunculis inferioribus ad
11-13 mm longis, ut folia pedunculos fulcrantia albo-arachnoideis et
pilis fuscis crassiusculis tectis. Calathidia sat magna, involucro 10-15
mm longo, 10-18 mm lato. Reliqua L. Hodgsonii Hook.

Habitat: in pratis paludosis et in betuletis humidis ins. Sachalin
et Moneron.

T y pus: Sachalin australis in vicinitate opp. Jushno-Sachalinsk,
in angustiis Tankovoje jugi Sudzuski, in declivibus montium, 600-700
ms. m., 28 VII 1948, fl., M.G. Popov. Typus et isotypus in Herb. Inst.
Bot. Acad. Sc. URSS in Leningrad conservantur.

Affinitas: A proxima L. Hodgsonii Hook. bene differt caulibus
pedunculis, foliis floralibus nec non involucri phyllis pilis fuscis
crassiusculis vestitis.

58. Sect. Ligularia.—Syn. cfr. supra 801.

Calathidia in inflorescentiam simplicem racemosam aggregata
(pedunculi infimi nonnunquam calathidia 2—3 ferunt). Folia pedunculos
fulcrantia sat magna, saepe colorata, squamiformia, inferiora saltem
naviculiformia petiolum et non raro involucrum amplectentia. Pappi
setae brunnescentes vel rufescentes. Floribus disci vel eius parte
tubuloso subaequilongae. Antheropodium filamento paulo latius; styli
rami apice angustati vel appendice acutata vel rotundata aucti; stigmata
ramorum totam paginam superiorem occupantia. Calathidia mediocria,
involucris angustis cylindraceis campanulatisve.

Typus sectionis—L. sibirica (L.) Cass.

Spesis sectionis Ligulariae Asiae austro-orientalis, orientalis,
borealisque nes non Caucasi et Europae incolae sunt.

59. Series Speciosae Pojark. ser. nova.—Pappi setae breviusculae
florum disci parte tubuloso aequilongae vel vix longiores.

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848

Typus seriei—L. Fischeri (Ldb,) Turcz.

Species Sibiriae orientalis, Orientis extremi (Korea, Mandshuria
et Japonia inclusae) nec non China temperatae incolae.

Series Racemosae Kitam. Comp. jap. III (1942) 190, p. min. p.
emes Pappi flori corollae aequilongae vel vix breviores. Plantae pro
norma altae, caulibus robustis, foliis magnis, calathidiis numerosis.

Typus seriei—L. sibirica (L.) Cass.

Area geographica seriei: Asiae regiones temperatae,
Caucasus, montes Europae centralis occidentalisque.

61. L. abakanica Pojark. sp. nova.—Cineraria sibirica auct. non
L. (1763); Ldb. FI. alt. IV (1833) 102, p. p. et excl. syn.—L. sibirica
var. hirsuta Serg. in Kryl. Fl. Zap. Sib. XI (1949) 2861.

Rhizoma abreviatum radicibus crassiusculis funaliformibus, superne
residuis fibrosis petiolorum annorum priorum dense vestitum. Caules
1-2, erecti 20-70 cm alti, 3.5-8 mm lati, robusti, a basi ob pilos
fuscescentes perplexos villosi vel tomentosi. Folia in sicco rigidiuscula,
subtus dense vel densiuscule fusco pubescentia, margine plus minusve
regulariter acute et plerumque crebre dentata, radicalia 1-3, saepius 2,
late cordata vel triangulari-cordata, saepe longa quam lata, rarius
cordato-reniformia, 6.5—12 cm longa, 7-40 cm lata, apice breviter
acutata vel obtusa, basi plerumque anguste et profunde, rarius lata
incisa, petiolo crassiusculo, quam lamina sesqui—duplo longiore, sed
nonnunquam ei aequilongo vel paulo breviore, pubescentia fusca,
patenti, villosa vel tomentosa vesito;-folia caulina 1-3, saepius 2,
inferiora radicalibus conformia, sed interdum brevius petiolata, petiolo
basi anguste vaginato; folium supremum lamina parva triangulari,
petiolo brevi in vaginam amplam inflatam dilatato, folia sub
inflorescentiam plerumque 1-3 in numero, ovata sessilia amplexicaulia,
foliis pedunculos fulcrantibus conformia sed majora. Calathidia 7-12
in numero, in inflorescentiam racemosam, 6.5—20 cm longam, ellipticam
vel oblongam, compactam aggregata; pedunculi crassiusculi, superiores
2-8 mm longi, inferiores 10-12 mm raro 3-5(6) cm longi et interdum
calathidia 2—3-ferunt, omnes pilis longis fuscis multicellularibus et
insuper albis arachnoideis dense vestiti; folia pedunculos fulcrantia
omnia lata, ovata, sessilia, extus praecipue ad margines alboarachnoidea,
plus minusve anthocyano colorata, apice non raro acute dentata,
inferiora 2—4 cm longa, naviculiformia, superiora plana involucro
sublongiora; calathidia cum ligulis 3.5-4.5 cm diam., involucrum basi
foliis 2-3 fulcrantibus ei subaequilongis, linearibus vel lineari-
lanceolatis praeditum, late campanulatum, basi pilosum, indistincte
biseriale, phyllis (7)9—11, glabris vel parce araneosis 9.5—11 mm longis,
dimorphis interioribus 3—4 mm latis, sat breviter acutatis, late scarioso-

849

marginatis, exterioribus duplo angustioribus, lanceolato-linearibus,
margine scarioso nullo vel angusto. Flores marginales 7-12, ligulis
oblongis vel obovato-oblongis 15-20 mm longis 4—6 mm latis, nervis
(5)7-10, tubo 5—6.5 mm longo; flores disci 26-42, corolla 7.5-9 mm
longa, tubo breviusculo 2.75—3.5 mm longo, parte dilatata campanulata
vel rarius anguste campanulata, 4.5-5 mm longa, dentibus oblongo-
triangularibus; pappus albido-fuscus, corolla subbrevior; antherae basi
sagittatae, auriculis breviusculis subulatis appendice apicali ovato-
lanceolata; antheropodium haud latum; styli ramuli planiusculi, apice
spathulatim dilatati, rotundati, extus ad basin pilosi. Achenia angusta,
cylindracea, brunnescentia, 5 mm longa, pappo acheniis subaequilongo.

Habitat: in pratis humidis necnon ad ripas paludosas fluviorum
lacuumque Sibiriae: in regionibus montanis et praemontanis jugorum
Altai, Kuznetzki et Abakanski.

T y pus: Sibiria centralis (distr. Minussinsk) in valle fl. Uibat, in
pratis humidis, 1(14) VIII 1909, fl. V. I. Smirnov. Typus et isotypus
in Herb. Inst. Bot. Acad. Sc. URSS in Leningrad conservantur.

Affinitas: L. subsagittatae Pojark. praecipue similis, a qua
notis sequentibus differt: foliis radicalibus latioribus et ut caulis densius
pubescentibus, foliis pedunculos fulcrantibus omnibus ovatis (nec
lanceolatis vel linearibus), florum radii ligulis (5)7—10-nerviis (nec 4—
5-nerviis), florum disci parte angustata quam dilatata 1.5-1.75-plo
breviore (non subaequilonga).

62. L. sichotensis Pojark. sp. nova.

Rhizoma speciei praecedentis; caulis 18-80 cm longus, 2—3.5 cm
crassus, plus minusve purpureo coloratus, ut pedunculi glaber vel pilis
patentibus perplexis fuscis vestitus. Folia in sicco rigida subcoriacea,
supra laete viridia, subtus albido-glaucescentia, utrinque glabra vel
subtus ad nervos pilis crassiusculis haud dense obsita, radicalia petiolis
luminis sesqui-duplo brevioribus vel eis subaequilongis, saepe tenuibus,
laminis 5—11 cm longis et latis, cordatis vel orbiculato-cordatis, lobis
basalibus breviusculis obtusis, apice rotundatis vel obtusis, margine
non raro integerrimis vel emarginato-dentatis, dentibus latis vel
nonnunquam obsoletis; folia caulina nulla vel solitaria, infra medium
caulis disposita, lamina parvula 1.5—2 (rarius ad 4.5) cm longa petiolo
brevi fere omnino in vaginam transformato, folia apicalia 1-4, foliis
pedunculos fulcrantibus conformia. Inflorescentia simplex racemosa e
calathidiis (1-3)5—10; pedunculi involucro vulgo breviores, 3-10 mm
longi, rarius infimi 2-—3.5 cm longi, ut rachis racemi plus minusve
albo-arachnoidei et praeterea pilis patentibus breviusculis obsiti; folia
pedunculos fulcrantia omnia angusta, sublinearia vel infima anguste-
lanceolate, viridia vel purpurascentia, subaraneosa, pedunculis longiora,

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850

infima tantum interdum eis breviora. Inflorescentia racemosa; calathidia
cum ligulis 3.5—-4 cm diam., involucro basi phyllis fulcrantibus binis
eo subaequilongis instructo, campanulato vel campanulato-cylindraceo;
involucri phylla 7-8, acuminata, (8)9-10 mm longa, biformia, interna
ovata late membranaceo-marginata, externa lanceolata margine anguste
marginata, dorso omnia glabra, vel interdum pilis crassiusculis parce
obsita vel subaraneosa. Flores marginales (4)5-7, ligulis oblongis vel
oblongo-ellipticis, 10-17 mm longis, tubo (5)7 mm longo, flores disci
ad 20 in numero, corolla 8-9 mm longa, parte superiore longiuscule
campanulata parte tubulosae subaequilonga; styli ramuli spathulato
dilatati, apice rotundati, pappus corolla brevior. Achenia immatura 5—
6 mm longa.

Habitat: in declivibus siccis lapidosis et herbosis, ad margines
silvarum paludumque a maris litore usque ad regionem alpinam Orientis
extremi. Planta declivi orientali montium Sichote-Alinj endemica.

T y pus: Oriens extremus, regio Primorski, montes Sichote—Alinj:
vallis fl. Botchi, in parte superiore fl. Jasy, in declivibus apertis, ad
1000 m s. m., 29 VII 1924, fl. I. Schischkin, n° 380. Holotypus in
Herb. Inst. Bot. Acad. Sc. URSS in Leningrad conservatur isotypi—in
Leningrad et in Valdivostok conservantur.

Affinitas: A L. sibirica (L.). Cass. foliis minoribus, saepe
integerrimis, inflorescentia depauperata, e calathidiis (1—3)5—10 (nec
30-50) composita, calathidiis paucifloris, florum disci corollae parte
dilatata late campanulata tubo aequilongo (nec tubuloso-campanulata
tubo sesqui longiore).

63. L. subsagittata Pojark. sp. nova.—Cineraria sibirica auct. fl.
cauc. non L. (1763).—L. sibirica auct. fl. cauc. non Cass. (1823).

Rhizoma abbreviatum radices crassiusculas numerosas emittens
caulis basi residuis fibrosis tenuibus foliorum vetustorum cinctus,
erectus, robustus, sulcato-costatus, glabratus, sub inflorescentia tantum
(ut pedunculi) pilis patentibus crassiusculis brunnescentibus et praeterea
albis araneosis sparse aut dense vestitus, 25-90 cm altus, 4-7 mm
crassus. Folia supra saturati viridia, glabra, subtus pallidiora
glaucescentia, tota pagina pilis breviobus brunnescentibus densiuscule
vel sparse obsita, textura saepe subcoriacea, radicalia 2-3, cordata vel
cordato-sagittata, rarius triangulari-sagittata, 5.5—18(24) cm longa, 4.5—
15 cm lata, sesqui (duplo) longiora quam lata, apice breviter acutata
vel obtusa, basi profunde incisa, lobis basalibus majusculis, acutiusculis
vel obtusis, deorsum spectantibus vel divergentibus, margine
inaequaliter grandidentata, petiolo 1.5—2.5-plo lamina longiore, basi in
vaginam haud longam breviusculam dilatato; folia caulina saepe 3,
infimum radicalibus conforme, sed petiolo breviore, longius vaginato;

890

851

folium medium lamina minore petiolo ei subaequilongo, longe vaginato,
folium superius parvulum, petiolo in vaginam amplam inflatam
transformato; folia superma (1)2-6 sessilia, amplexicaulia, violaceo
colorata, foliis pedunculos fulcrantibus similia sed majora. Calathidia
7-35, sub anthesi nutantia, inflorescentiam racemosam 4.5—30(34) cm
longam formantia; pedunculi inferiores ad 3.5 cm longi, superiores
0.3-1 cm longi; folia pedunculos fulcrantia plus minusve fusco-violacea,
glabra vel subaraneosa, lanceolata, longe tenuiterque acutata, inferiora
22-35 mm longa, naviculiformia, superiora minora, involucro
sublongiora, nonnunquam linearia. Involucrum basi calyculi phyllis
duobus linearibus eo subaequilongis instructum campanulato
cylindraceum obsolete biseriale, phyllis 8-10, tenuibus chartaceis,
pallidis, interdum plus minusve anthocyano coloratis, 10—-12(14) mm
longis, glabris aut parce pilis crassiusculis patentibus vel appressis
arachnoideis tectis, dimorphis, internis lanceolatis, marginibus late
membranaceis, externis angustioribus linearibus, non vel angusie
membranaceis, omnibus longe acuminatis. Flores radii 7-12, ligulis
aureo-luteis, 8-15 mm longis, 2.5-3.5 mm latis, nervis 4-5, tubo 4—
5.5(7) mm longo; flores disci 17-32, corolla 8-11 mm longa, parte
dilatata anguste campanulata, 3.5-5 mm longa, dentibus oblongo-
triangularibus 1-1.5 mm longis, parte angustata 4-4.5 mm longa;
stamina et styli ramuli speciei praecedentis; achenia anguste cylindracea,
5-6 mm longa, pappo subaequilongo albido-rufescenti.

Habitat: in pratis alpinis humidis fere toti Caucasi.

Ty pus: Caucasus Magnus: prope stationem Kazbek, in prato
paludoso vallis fl. Terek in declivo occidentali montium, 10 VIII, 1916,
P.N. Krylow et E.I. Steinberg. In Herb. Inst. Bot. Acad. Sc. URSS
conservatur.

Affinitas: cfr. supra p. 888.

64. Series Longipedes Pojark. ser. nova.—Pappo setae florum disci
corollae subaequilongae. Plantae pro norma humiles, caulibus, tenuibus,
foliorum radicalium petiolis saepe longissimis, laminis parvis, 1.5—
8(10) cm longis, saepissime sagittatis, lobis basalibus elongatis.
Calathidia enumerosa.

Typus seriei: L. longipes Pojark.

Species tres in Sibiria, Europa arctica et subarctica nec non in
montibus Carpaticis distributae.

65. L. longipes Pojark. sp. n.

Rhizoma abbreviatum, incrassatum, radicibus numerosis
funaliformibus, superne residuis fibrosis foliorum vetustorum dense
obsitum; caules 30-65 cm alti, 1.5—2.5 mm diam., tenues, remote foliati,

891

852

tenuiter striati, glabri, subinflorescentia tantum pilosi, basi vel tota
altitudine purpureo colorati. Folia supra laete viridia, subtus pallidiora,
glaucescentia utrinque glabra, textura in sicco tenui sed rigidiuscula,
margine sat regulariter emarginato-denticulata, dentibus latis
triangularibus, in mucronem callosum subito contractis; folia radicalia
2-3(4), petiolis tenuissimis 0.75—2 mm crassis, glabris, longis, lamina
2.5—5-plo longioribus, non raro purpureis, laminis 2.3-8(10) cm longis,
saepe elongatis, sagittatis vel triangulari-sagittatis, vel triangularibus,
rarius reniformi-sagittatis vel reniformi-cordatis, apice plerumque
acuminatis, rarius obtusis, basi profunde incisis, lobis basalibus saepe
longis, non raro dimidio totae laminae aequilongis, rarius ad 1.5—2.5-
plo ea brevioribus, apice acuminatis vel rotundatis; folia caulina 2
(rarius 3) valde remota, inferum radicalibus conforme, sed nonnunquam
brevius petiolatum et latius vaginatum, superus lamina parvula, petiola
in vaginam amplam inflatam dilatato; folia apicalia nulla aut 1—2(3),
subsquamiformia, plus minusve purpurascentia, ovata sessilia, foliis
pedunculos fulcrantibus inferioribus similia sed majora. Calathidia
interdum 1-2 («L. sibirica var. alpestris Turcz.») sed vulgo 3-15
(rarissime ad 20) in numero, in racemum laxiusculum, 2—3.5 cm longum
aggregata, cum ligulis 2—3.5 cm diam., breviter pedunculata, primo
suberecta dein nutantia, cauli parallela, pedunculis superioribus 2 mm
longis, inferioribus 8-30 mm (rarissime ad 4—7 cm) longis, ut rachis
pilis crassiusculis albidis demum brunnescentibus et interdum albis
araneosis sparse tectis; folia pedunculos fulcrantia ovata, cymbiformia,
purpurea. Involucrum basi foliis fulcrantibus binis filiformibus vel
lanceolato-linearibus praeditum, cylindraceum vel anguste
campanulatum, indistincte biseriale, phyllis 7-8 glabris apice tantum
pubescentibus 8-11 mm longis, plus minusve purpureo coloratis,
biformibus, internis lanceolato-ovatis, margine late membranaceis,
breviter acutatis, externis multo angustioribus, linearibus vel lanceolato-
linearibus, tenuiter longeque acuminatis, haud membranaceis. Flores
marginales, 6-7, ligulis oblongis, 11-15 mm longis, 3-4 mm latis,
tubo 4.5-6 mm longo; flores disci 15-20, corolla 7-9.5 mm longa,
parte apicali dilatata, breviuscule campanulata, parte basali tubulosae
subaequilonga; antherae basi breviter sagittatae, appendice apicali
longiuscula; antheropodium elongatum. Styli rami tenues, appendice
leviter dilatata, extus ad basin pilis brevibus conicis obsiti. Achenia
pallida, S-6 mm longa, pappo rufescenti, achenio paulo longiore.

Habitat: in pratis paludosis salsuginosisque, in vallibus
fluminum, in paludibus herbosis in regionibus silvatica, stepposa
alpinaque. Sibiriae occidentalis orientalisque, Mongoliae borealis, nec
non Orientis extremi.

T y pus: Transbaicalia occidentalis in districtu lacuum Eravinskije
prope pag. Constantinovka, in salebris silvae, 3 VII 1912, fl., M.

892

853

Korotki, Z. Lebedeva et M. Okuschko, n° 263. Typus et isotypus in
Herb. Inst. Bot. Acad. Sc. URSS in Leningrad conservantur.

Affinitas: AL. arctica Pojark. foliis angustioribus, sagittatis
(non cordatis vel triangulari-cordatis), lobis basalibus longioribus,
petiolo longiore, lamina 2.5—5-plo (nec 1.5—2 rarius 3-plo) longiore
nec non florum radii ligulis latioribus differt.

66. L. arctica Pojark.—L. sibirica var. minor Nym. Consp. fl.
Europ. (1879) 350.

Rhizoma speciei praecedentis; caulis 15-70 cm altus, tenuis 1.5-
3 mm crassus, fere efoliatus, saepissime violaceo-purpureus, raro in
parte media viridis, sub inflorescentia solum pilis crassiusculis
brunnescentibus obsitus. Folia in sicco tenuia, sed nonnunquam sat
rigidiuscula, supra saturate, subtus laete viri-dia; folia radicalia saepe
2, (1.7)2-7(11) cm longa, 2.2-7(10) mm lata, cordata, triangulari-
cordata vel cordato-sagittata, apice obtusa vel breviter acutata, lobis
basalibus divergentibus obtusis vel breviuscule acuminatis, margine
irregulariter dentata, dentibus latis triangularibus in mucronum brevem
contractis, nonnunquam indistincte remote dentata petiolis tenuibus,
lamina 1.5—2(3) plo brevioribus; folia caulina 2, inferus radicalibus
conforme, sed petiolo breviore et longius vaginato; superus longe
remotum, lamina parvula, petiolo in vaginam amplam amplexicaulem
transformato; folia apicalia saepe nulla, rarius unicum squamiforme,
sessile, lanceolatum, foliis floralibus conforme, sed majus. Inflorescentia
simplex racemosa laxa e calathidiis 2-7(12) composita; pedunculi pilis
crassiusculis fuscescentibus obsiti, inferiores 12—35(50) mm longi,
superiores 6-10 m longi; folia pedunculos fulcrantia fusco-purpurea,
lanceolata, infima interdum ovata, longe acuminata, integerrima,
pedunculum cum involucro amplectentia, superiora plana, non raro
linearia. Calathidia cum ligulis 2-3 cm diam., post anthesin nutantia;
involucrum basi foliis fulcrantibus binis linearibus instructum,
cylindraceum vel cylindraceo-campanulatum, plyllis 6—-8(9), glabris,
totis vel partim, purpureis, 6—-10(12) mm longis, biformibus, internis
lanceolatis vel subovatis late membranaceo marginatis, externis
angustioribus margine haud membranaceis. Flores radii 5—6(7), ligulis
linearibus vel oblongis, 8-10 mm longis, 1.25—4 mm latis, 3—-6-nerviis,
tubo 3-4 mm longo; flores disci 17-26, corolla 7-8 mm longa, parte
superiore campanulata vel late campanulata, 3—3.5 mm longa, in tubum
tenuem subaequilongum subito contracta; filamenta et styli ramuli
speciei praecedentis. Achenia anguste cylindracea, 5—6.5 mm longa,
pappo albido-fuscescenti eis sublongiore.

Habitat: in pratis paludosis, in paludibus herbosis, ad ripas
humidas lapidosasque nec non in tundris fruticosis et silvarum pratulis

893

854

in Rossiae europaeae regionibus arctica et subarctica, in jugo Ural et
in parte boreali-occidentali Sibiriae occidentalis.

T y pus: Rossia septentrionalis: peninsula Kanin, in declivi boreali
jugi Pae-hoi, in saliceto vallis lapidosae, 26 VII 1928, fl. V.N. Andrejev,
n° 525.

Affinitas: Cfr. supra

67. Sectio Stenostegia Pojark. sect. nova.—Folia pedunculos
fulcrantia angusta, parva, linearia vel subulata, inferiora pedunculos
multo breviora. Antheropodium filamento paulo latius. Stigmatis striae
latae, prominulae, usque ad appendicem extensae. Pappi pili albi,
molles. Folia viridia (nec glauca) textura molli (mon carnosa, nec
subcoriacea).

Typus sectionis—L. narynensis (Winkl.) O. et B. Fedtsch.

Species Asiae praecipue mediae centralisque propriae, paucae
Orientis extremi incolae.

68. Subsectio Oligocephalae Pojark. subsect. nova.—Calathidia
magna aut mediocria, solitaria vel 2-10 in numero, in inflorescentiam
racemosam aggregata. Involucrum cupuliforme. Rhizoma tomento dense
vestitum. Folia radicalia ovatovel orbiculato-cordata, rarius
subreniformia.

Typus subsectionis: L. narynensis (Winkl.) O. et B. Fedtsch.

Species Asiae mediae incolae.

69. Subsectio Racemiferae Pojark. subsect. nova.—Calathidia
mediocria (4.45 mm diam.), numerosa, in racemum longum angustum
aggregata. Involucrum campanulatum. Folia radicalia reniformi-
triangularia vel triangulari ovata Rhizoma nudum.

Typus subsectionis—L. jaluensis Kom.

Area geographica subsectionis—Korea et regio Ussuriensis.

70. Subsectio Paniculatae Pojark. subsect. nova.—Calathidia
parva, plerumque paniculatim aggregata. Rhizoma tomento denso
vestitum.

Typus subsectionis—L. thyrsoidea (Ldb.) DC.

Species Asiae mediae centralisque nec non Sibiriae austro-
occidentalis incolae.

71. Series Thyrsoideae Pojark. ser. nova.—Involucrum apertum
campanulatum vel cupiliforme. Paniculae rami laterales etiam inferiores
rachide breviores, calathidia racemosim aggregata ferentia. Folia
oblongo-ovata, basi cordata vel sagittata.

894

855

Typus seriei—L. thyrsoidea (Ldb.) DC.
Series monotypica.

72. Series Songaricae Pojark. ser. nova—Involucrum angustum
cylindraceum vel obconicum. Calathidia in paniculam pyramidatam
vel corymbosam (inflorescentiis partialibus racemosis) aut in racemum
aggregata. Folia oblongo-triangularia, acuta, basi cordata.

Typus seriei—L. songarica (Fisch.) Ling.

Species duae Asiae mediae incolae.

73. Series Thomsonianae Pojark. ser. nova.—Involucrum anguste
cylindraceum vel obconicum, aureo-fuscum. Inflorescentia generalis
paniculato-corymbosa, inflorescentiis partialibus corymbosis ad apicem
ramorum lateralium dispositis. Folia orbiculari-ovata usque ad
reniformi-triangularia, texturamolli supra atriviridia.

Typus seriei—L. Thomsonii (Clarke) Pojark.

Species duae Asiae mediae, Afghanistaniae nec non Himalayae
occidentalis incolae.

74. Series Altissimae Pojark. ser. nova.—Involucrum tubuloso-
campanulatum. Inflorescentia generalis paniculato-pyramidata,
inflorescentiis ramorum lateralium racemosis subpaniculatisve
apicalibus. Folia laete viridia, crassiuscula.

Typus seriei—L. altissima Pojark.

Series monotypica.

75. Sectio Senecillis (Gaertn.) Pojark. sectio nova.—Senecillis
Gaertn. De fruct. et sem. II (1791) 453.—Senecio groupe Ligularia B.
Euligularia 1. Racemosi Franch. in Bull. soc. bot. Fr. 39 (1892) 189,
p. p.—Cyathocephalum Nakai in Tokyo Bot. Mag. XXIX (1915) II.—
Ligularia sect. Cyathocephalum Kitam. |. c. 185.—descript. emend.

Calathidia plerumque parva, rarius mediocria in racemum simplicem
vel in paniculam elongatam angustam aggregata. Involucrum tubulosum
aut fere cupuliforme, phyllis liberis aut in cyathium apice 3-4 dentatum
connatis. Pappi setae albae vel fuscescentes, molles vel rigiusculae
longae vel breviusculae brevissimaeque. Antheropodium filamento
distincte latius. Styli rami appendice conica aucti. Folia laeta glauca,
crassiuscula, succosa vel subcoriacea.

Typus sectionis—L. glauca (L.) O. Hoffm.

Species huius sectionis in Asia orientali, media, centralique, Irania,
Sibiria et Europa distributa sunt.

895

856

76. Subsect. Sericochaete Pojark. subsect. nova.—Pappi setae
albae, molles, longae. Involucri phylla inter se libera. Caules fistulosi.
Rhizoma glabrum.

Typus subsectionis—L. macrophylla (Ldb.) DC.

Species Asiae mediae, Chinae centralis nec non Iraniae incolae.

77. Subsect. Stereochaetae Pojark. subsect. nova.—Involucri phylla
libera vel in cyathium 3-—4-dentatum connata. Pappi setae fuscae
rigidiusculae, floribus disci breviores. Calathidia in paniculam oblongam
densam aggregata. Caules solidi. Rhizoma pilis flexuosis vestitum.

Typus seriei—L. mongolica (Turcz.) DC.

Species Orientis extremi incolae.

78. Series Mongolicae Pojark. ser nova.—Involucri phylla libera.
Pappi setae 3-7 mm longae.

Typus seriei—L. mongolica (Turcz.) DC.

Species tres in regione Amurensi, in China boreali-orientali nec
non in Korea (insula Quelpart inclusa) distributa.

79. Subsectio Senecillis (Gaerth.) Pojark. subsect. nova.—Senecillis
Gaertn. De fruct. et sem. II (1791) 453, tab. 173, p. p. typica.—
Calathidia in inflorescentiam racemosam densam. aggregata (pedunculis
inferioribus nonnunquam calathidia 2—3 ferentibus). Involucrum
apertum, late campanulatum vel cupuliforme, phyllis 7-12. Pappi setae
brevissimae, 0.5—2 mm longae, rigidiusculae, aculeato-barbellatae.
Caules fistulosi.

Typus subsectionis—L. glauca (L.) O. Hoffm.

Species duae Sibiriae austro-occidentalis nec non Podoliae et
Transsilvaniae propriae.

80. Sectio Glossophyllum Pojark. sect. nova.

Calathidia mediocria, 1-3 in numero. Involucrum campanulato-
cupuliforme, manifeste biseriale, phyllis distincte biformibus. Folia
pedunculos fulcrantia parva, viridia, lanceolata. Antheropodium
incrassatum, filamento conspicue latius. Styli rami appendice conica
instructi, facie superiore stigmatis striis marginalibus latiusculis,
elevatis, pagina inferiore sub appendicis basi barbellata. Folia radicalia
elongata in petiolum angustata, integerrima vel indistincte remote
dentata.

Typus sectionis—L. Pavlovii (Lipsch.) Cretz.

Sectio monotypica montium Kara-tau Asiae mediae incola.

857

81. Subgenus Dolichorrhiza Pojark. subgen. nov.

Rhizoma tenue, lignescens, repens, radicibus tenuibus. Calathidia
magna solitaria vel 2-6 in numero. Involucrum apertum cupuliforme,
indistincte biseriale, phyllis (7)9-22, plus minusve manifeste
biformibus, internis latioribus anguste membranaceo marginatis
exterioribus (interiorum margines tegentibus) angustioribus margine
haud vel vix membranaceis. Antheropodium latum incrassatum. Styli
rami appendice semiconico praediti, facie superiore stigmatis striis
marginalibus prominulis, facie inferiore sub appendicis basi barbellata.
Folia reniformia.

Typus subgeneris—L. renifolia (C.A.M.) DC.

Species tres Caucaso endemicae.

CALENDULA L.

82. C. karakalensis Vass. sp. nova.

Planta annus dense glandulosula caule a basi ramoso, foliis
integerremis acutis oblongis, foliis inferioribus petiolatis foliis summis
sessilibus. Calathidia parva; involucri phylla uniserialia oblonga acuta.
Flosculi radiales lutei involucra 1-1%-plo longiores. Achaenia externa
falcata 15-20 mm lg. late lanceolata, dorso obsita seriebus
longitudinalibus aculeolorum setaciiformum in vallecula duobus
lateralibus margine setaceo-denticulatis alis formata; margine ventrali
achaenia alata irregulariter denticulata. Achaenia media cymbiformia
cum alis membranaceis interius involutis, dorso aculeolato-setacea
circumscriptione late ovata vel elliptico-rotundata, achaeniis externis
duplo breviora. Achaenia interna annularia exalata dorso aculeolato-
setacea apice cum excrescentia membranacea unilateralia.

Habitat: in declivibus argillosis jugi Kopet-dagh occidentalis.

T ypus: Turcomania, distr. Karakalensis ad merid. a vic. Kara-
Kala, in declivibus argillosis, 27 VI 1934, n° 266. A. Borissova.

Affinitas: A proxima C. arvensis L. achaeniis externis alatis
circumscriptione lanceolatis (nec cylindricis) differt.

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905

INDEX ALPHABETICUS'

nominum specierum atque synonymorum plantarum
in tomo XXVI Florae URSS commemoratarum

Abrotanum (Bess.) Rydb., subgen. 456

FS SOG Baines ac oragiveorsoss th dees Ue 456
Absinthium, .DC., Sect2n8.22s20.02: 492
=> lalpinum, BESS. (,.....2-...%. 2c! 493
— divaricatum Fisch. ...........0c08 523
== frigidumBesswnnsniiges. gato 497
— grandiflorum BESS. ..........cc 502
— Korotkyi Krasch. ...........cccecee 523
— lagocephalum Fisch................... 504
— €. multicaulis Bess. ............0. 523
=—Inifens Stevia. abotiienss: 502
— peduncularis Stev. «0.00 494
— sericeum BESS. .......ccccccccseeseeteees 502
— Sieversianum BeSS. ........c0ccee 5.17
— splendens (Willd.) Bess. ............494
— viridiflorum rupestre Bess....... 509
— — viride BESS. .....eccceceeceneees 509
— vulgare Lam. .0.......:cceseseeeeees 515
Achillea, L.ipast:.aieaamal...Ani se 70
— abrotanifolia L. wees 329
— abrotanifolia Willd. ...................... 97
— acuminata Frey. ...........:.:cece 109
— — acuminata (Ldb.) Sch. Bip..108
— aegyptiaca S. Gmelin. ..........0..... 96
— albicaulis C.A.M. ...c.cccccceeee 103
= jalpina auct.,..2.tian.. candor LL?
= dal pinasb.. .ica.3. Aeah, BARR 119
— amoena C.A.LM. ....cceccecceceeteeteeees 102
— anethifolia Fisch. ..........ccccccceee 81
SMargentea Lam. :..:-<.scccevonnsePted eZ 361
SASIAUGCA SOU Bite. 2. Riyvaceectacenoviuch .ctts 85
== aSiatica Serg...vack.idddasidok 85
— Biebersteinii Afan. .......000 97
== ibipinnata. Laois. ARM. WE 331
— birjuczensis Kok. .........0.cee 96
— biserrata M.B. uu... cee 106
— — var. griseo-virens Kolak. ..... 107
— borealis Bong. .........:.cccsesseeesesesees 81
— borysteniea Kok. ........0.eeeee 11-2
— bucharica Wink. ..........0.c.e 86
— camtschatica Rupf. .............06 122
SS (Cancrini. Gru. ..:5,600000000. Sedat 96
= lcarpaticarBlaink ine. cae ae 87
— cartilaginea Ldb. ......00. 114

— — ssp. borystenica Sakalo. ...... 112
— — var. latifolia Rupt................. 114
— cheilanthifolia Somm. and Lev. .99
— COarctata POUT. ...... cc eeeeeeeereeeeeees 91
— compacta WAI. ..........ccceeeeneee 91
— corymbifera S. Gmelin. ............... 76
— cuneatiloba Boiss. and Buhse.....99
— decumbens auct. ......ccccceseeeeeeeee 97
— dentifera D.C. ..cceccccccccecesceceseeeeee 87
— — B. distans (Waldst. and Kit.)

FIGS, ......ss-aiensass eters. Cinvereaen eet 87
— distans Waldst. and Kit. .............. 87
— — ssp. stricta (Schleich.) Janchen.

snags snscisnayncenonachavadtcoethe. AUOLE MEET 88
— — var. lanuginosa (Gaud.)

Mansf. {...1) ak taliimtetit. thee. 88
— dobrogensis Prod. .........ccceeeeeeee 92
— dracunculoides Desf..............0 109
— dschimilensis C. Koch. ..........004 89
— CTIOPNOTA AUCL. .......cccceeeceerceeeeeneee 101
— eupatorium M.B, .......cceceeeeeees 90
ae PLLCEOUIG,, NU cis wasceeeseantsinccearevaresien 90
— filipendulina Lam........0... 90
— — var. eupatoria (M.B.) DC. ..... 91
— — var. filicifolia (M.B.) DC. .....91
— — var. leptocline DC............0... 91
— — var. Szowitziana DC............... 91
— iGerberi Willd ).2:000c8...25¥..% 96
— — var. leptophylla Serg. ............. 96
— — a. genuina Schmalh. .............. 96
— — @. genuina Schmalh. b.

tenuifolia Schmalh. .................0++ 96
— — Q.’major C. Koch. ......... 96
— — B. leptophylla (M.B.)

Schima. .........s25 ses. tsseeeeen ees. 94
— — B. minor C. Koch. ............00. 96
— — B. subcristata DC. ........0 96
— glaberrima Kok. ............:.:ccceseee 95
— glomerata M.B. ..........ccccseseeseeseees 91
— grandiflora M.B. ..........:c.scceee 109

906 — griseo-virens AID. ...........:ccceeeee 107

— Impatiens pists ii., mecseest... I... 388 116
— — ssp. euimpatiens Heimer......116
— — ssp. Ledebouri Heimerl........ 117

‘Reproduced from the Russian original. Page numbers of the left-hand margin in

the text—General Editor.

860

— ircutiana Sch. Bip. .........:0++ 114 =——- odorata GrOSSH. \...4::5.:cseosnzesncesenanee 97
— japonica Heimer. ...............000+ 120 mr OMOTAA MOCK: ......cisceceseenvsenenerestscs 77
— kermanica Gand. ...............e 101 — odorata’ Pall. ois. .t0c ree 716
— Krascheninnikovii Afan. ............ 101 =, = RODS, saepessndesks vedsepacapeecctesans cee 83
=a NABI AGIE Ds cuts ccancckeowsesevseatncoseconsscs 89 we tin NNN. sous anc cnapentbnl eh vet svapeeote 83
ma LEAFLOD IE TODD ncscre ene -ncsneterecevenestire 195 Sar ONSICH en OCI oreo create anti enaes 78
— Ledebouri Heimerl. .................... 117 — oxyloba DC. ssp. Schuri Heimer.
ams POMEMISES: WUE Zs. 5s.acsenaxesenseronesrestres il SOR eRe Nore ern eee cre Pont ee reyes) 123
leptopliylla® M:Bivetzsctet...jee 94 — pannonica Scheele. .............:..000 82
— f. desertorum PaszosKi........... 94 — pectinata Willd. «00.0... 93
— f. granitica PaczosKi. ............. 94 ==! ptarmica,.auct.t223...2.0.. eee 109
leptophylloides Prod. ..........0:000+ 94 =iptarmica.L..............2220.. San 108
leucanthema Persoon. .........:..:0++ 109 — — ssp. cartilaginea Heimerl. ...114
lingulata Waldst. and Kit. ......... 105 — — ssp. euptarmica Heimer! var.
macrocephala Ruff. ..............:0 111 acuminata Heimer. ............000.+ 108
MAQNGA. AUC... .2:.[0Ne.. needa see 78 — — ssp. euptarmica Heimer! var.
micrantha. auct....,...A42.e0enes 100 genuina Heimer. .............:0:008 108
micrantha . Willd’ }..2.0aie..2 cx. 96 — — ssp. macrocephala Heimer! var.
micrantha: Willd. ...c.:ap<0.-.1.2tenes 97 angustifolia Heimerl. ....... 109, 111
— B. sulfurea DC... 97 — — ssp. macrocephala Heimer! var.
micranthoides Klok.................. 100 latifolia Heimerl. sent. tans 111
millefolivn AUCt....<..c.cccssessececneneases 81 — ptarmica L. ssp. ptarmicaefolia
millefolium) Lasiveignen.. 22... 78 Heimerl var. grandiflora Heimer.
ssp. eu—millefolium (L.) HayeK.78 = sesseseseseseerencneasevenstcuencecanensseucscanenenes 110
ssp. millefolium (L.) Hayek. ....... 78 — — ssp. ptarmicaefolia Heimerl var.
— ssp. pannonica (Scheele) Ruprechtiana Heimer)................ 110
Hayek...... 2204...) Seen: 82 — — var. Beringiana Trautv. ....... 111
— ssp. sudetica (Opiz) Weiss.... 80 — — var. speciosa Herd. ............+ 111
— var. alpestris Vimm. and Rab.. — — y. speciosa Schmalh. ............ 115
didgduteedcstudendesSanedeavcu PT RR 80 — ptarmicifolia (Willd.) Rupr. ......109
— var. intermedia Serg. .............. 86 — ptarmicoides Maxim. ...............++. 121
— var. macrophylla Serg. ........... 78 — PUbescens AUct. ........cccccseceeeereeees 97
— var. micrantha (M.B.) — romanica Prod. .........0..cccsssseesseeeee 92
Schmalh.....4).2ineigal...% 97, 100 — Sachokiana Sos. ........:.:ccseee 110
— var. setacea Ldb............0c 82 = jsalicifolia .Bess..........<..--a..@@ineae 112
— var. setacea (Waldst. and Kit.) — sambucifolia Desf...i....2...:cs:<scs 194
Koch: ....2250- ee SAVER... EN. caren 83 == santolina auct. 222%. s.unawa 101
—.y. lanatanKochornins....% a. 82 SE BAK. ccccccecsccisces ttt IRR 96
— 6. lanuginosa Gaud. ............... 88 ES ey) EE Rr 100
— 6. occidentalis DC. ..........000 81 — Schischkinii Sosn. ...::2....000i04 103
mollis... ANGiRHSA:. .. 1. OARE...33. 007% 83 — schugnanica Wink. .........:.:.000+ 86
mongolic@. Fisch. wiieiie:...35-20: 119 = /Schuri. Heimer. .........1.. 222000" 123
myriophylla Willd. ........0:c000 329 = Schuri. Sch... Bipsi st. 123
Nabelekii. Heim. ..... 2530008 ..25 «<7: 84 — Sedelmeyeriana Sosn. ................. 113
Neilreichii . Kermisy)i2nint...22.<0; TT — septentrionalis (Serg.) Botsch. .. 115
nobilis: auct.,....:.201%.. GRR. Th = sericea. Janka. ......-S oR). Rieu, 92
NODS Lo. sc csicines nth, BUSI 76 = \setacea. auct......2012. ..eihainae, 85
— var. Neilreichii (Kern.) Beck.77 — setacea Waldst. and Kit. ............. 83
— B. ochroleuca Boiss. .............. 77 90% = sibirica.auct. .2348.. soho 122
— x A. setacea Waldst. and Kit. .. SO —— LAD. oncscncennescess RTA. I. 119
svavns eS, AR NN, ABBA, 208 Vt — — ssp. camtschatica Heimerl...122
Nyaradyana Prod. ........cccccceseeeeeees 93 — — ssp. japonica Heimer. ......... 120
OCHTOLEUCA:'AUCE, <iss.sscsvsceseasnenes cases 77 — — ssp. mongolica Heimer. ...... 119
ochrolenca sBbrh .s.55.:asctetoesss:- 93 — — ssp. ptarmicoides Heimerl. ..121

— — var. discoidea Rgl................. 121
== "SPECIOSA: AUGE:“titiverttes cae et st. 114
— speciosa Spreng. .............:cccceeees 115
— speciosa Henckel f. occidentalis
RUpreyics. Peet ate ee 114
—<spinosay Schurys cst Res, Le 117
+= stricta, Schleichis):..20..04.. 20298". 88
— subhirsuta Gilib. «0.0.0... 19
— sudetica Opize sien... Le sOM ag. 80
— Szowitzii Fisch. and Mey ............ 89
— tanacetifolia All... 88
— tanacetifolia auct. ............:ccccce 87
—fanacetifolias. Ane. Bene Aen. 88
— — var. angustifolia Weihe. ........ 88

— var. a. dentifera (DC.) Koch.87
— var. B. distans (Waldst. and

Kit:)"Rochel™...7.. Hee Ren alg, 87
— — var. Y. stricta Koch. ............... 88
== fanrica: MiBs 22-2288. Re. 94
— — f. villosior Boiss. .................... 94
== tentufolia ‘Lamy int... ce) 103
— — var. albicaulis Trautv........... 103
— teretifolia LOD. ...........cceceeeeeees 101
— LOMENLOSA AUCC. .......ccccceceeseeeeeeesees 97
<=, SLAM OVS) enters tate ieee 92
— trichophylla Schrenk. ................. 125
— vermicularis Trin...................00. 102
— Wilhelmsii C. Koch............0....... 100
Aaenostyles Cassese 8e.. Le 684
Ajania Poljak. ...0.0.0... 398, 880
— fastigiata (Winkl.) Poljak. ......... 405
— fruticulosa (Ldb.) Poljak. .......... 406
— furcata (MB.) Poljak.................. 483
— globularia (Bess.) Poljak. ......... 481
— glomerata (Ldb.) Poljak. ........... 482
— gracilis (Hook. f. and Thoms.)

Polak iss gescnveiccsiv hte 407
— insulana (Krasch.) Poljak. .........484
— kokanica (Krasch.) Tzvel........... 408
— leontopodioides (Bess.) Poljak. 485
— manshurica Poljak. .......00.0..... 401
— Pallasiana (Fisch. ex Bess.) Poljak.

ane ceas daa rte eee, EAVOS 402
— Scharnhorstii (Rg!. and Schmalh.)

T2Ve liye ceen eee ae te. ket ee 409
— Senjavinensis (Bess.) Poljak. ....485

— tenuifolia (Jacq.) Tzvel..............411
— tibetica (Hook. f. and Thoms.)
T2veliveccaseth See se tees 410
— trilobata Poljak. ........00....... 409, 880
— variifolia (Chang) Tzvel. ........... 401
Allardia Decne. ::.Js:c--vrenss: 263, 267
Allardia (Decne.) Tzvel., sect. ....... 267
= plabra Decne? rc.tiets cect rack ocees 270

908

861

— Huegelii Sch. Bip. ...........0...0 271
— Stoliczkae Clarke. .............ccc0000 268
— tomentosa Decne. ..............cc000008 267
— tridactylites (Kar. and Kir.) Sch.
Bipsysesscrptena dle BGI. RRQ 269
Anactdea,-DC...Sectuni ih. 22M 150
Anacyelus | LAts...02is3d. AoE. 68
—=_aureus dbame caciedia tren Gir 151
— Ciliatus Trautv. ...........ccce eee 68
—_ clavatus Perse ck. PR. 70
— officinarum Hayne. ..............c00 69
— pulcher Bess... tRan. Bu. 69
=» pyrethrumy:DGr.:..c: 201. A. 69
— tomentosus DC. .0.........cccceeeeeeeees 70
Andropurpurea Maguire, subgen. ...665
Anthemidaceae Bessey. ..........00:cce 1
ANTHEMIDEAE Cass., tribus. .......... 1
Anthemideae Link, ordo. ...................6+ 1
Anthemidinae Hoffm., subtribus. ....... 9
ya) CE) Oe ee oe 9, 865
Anthemis..sect.,:2¢kat: ASRS. 35
— abagensis Fed. ..........0.0.0 56, 869
—LalpinaiaUuCts ivi. Reale. HABee 123
— — var. B. Willd. ee 18
= altissima «Liss: ARO. ROMER. 58
— Anahytae Woron..............0. 19, 865
— amatolica Boiss. ..........0...cccccesseeeee 24
— argaea Boiss. and Bal. ................ 25
== ARVEDSISy Lan. LEAOK. ROE AOE om 35
— arvensis B. ruthenica Schmalh. .36
= austriaca Jacg.%.1... 2a ca. 59
— Biebersteiniana C. Koch. ............ a1
— — @. Marschalliana Boiss. ........ 31
— — B. Rudolphiana C. A. Mey. ..32
— — ¥. pectinata BOisSS. ...........0.04. 32
— Bollei Sch. Bip. ......... 40
— brachyglossa C. Koch. .......... 10, 45
— caespitosa Herbich. .............0.4. 123
= .calcatea \SOsniiins.. SER. BRA. 30
— candidissima Willd. ...................... 37
— — var. epapposa Trautv. ............ 37
— — var. intermedia Trautv. .......... sir)
— — var. longepapposa Trautv......38
— carpatica Waldst. ..0.... ee 20
— caucasica Adam. ...............ccc008 161
— Biebersteiniana C.A.M. .............. 18
— — jHenckel,/..QRRiA a. <b cee. 41
— — Hormem. o..........cccccececeesseceeseeeeees 46
— chamamilloides Stev. ...........:.0.0.. 37
— chrysantha SoOsn. .......ccccceccceceeseees 41
— ciliata (Trautv.) Boiss. ................ 68
— COArCtALA AUCT. ..........cceeeecdeceteeeees 43
— coelopoda Boiss. .............. ccc 59
—.collina: SOsns sec: AR. OR. 52

862

— — var. intermedia Sosn. ............. 48
SsCcotula LG: cc... WS eee. 63
— f. lithuanica (DC.) Schmalh.......64
— cretacea |Zefir. sats. 2yubins. 43
= CTELACER , ZUDG.. «i nsercscenszenvscnes sachets 29
==/debilis ‘Fed. «.....:cdsce--..cchteh--es 44, 868
— deserticola Krasch. and M. Pop. 65
+=/dubta Stev. .......:c.cscditiee aaa. 51
— dumetorum Sosn..........0..cceeeee 47
— dumetorum var. subtripinnata
SSID. oc occoceocs conte etl Pete eee ts 47
== Emiliae Sosn: ;......-. vce). sates. 25
== E€UXina BOISS’:..\...)090)..tcerel nates: 33
=~ foetida VaMm. 5... MS eictecqe. 63
==AfTUHCULOSA ‘AUCH EE8. swiss 25
— fruticulosa M.B. ......ccscccceeeseeseeeee 28
— fruticulosa M.B. 00... 25
— Grossheimii Sos. ............::.:eeee 38
— Haussknechtiana Boiss. and Reut..
ih Warlenh abbaalehetin a hn nae el OL scree 38
— hemistephana Boiss. ............:0+-+ 39
<=shirtella Wink ......iidsiaewsutx 38
——ihybrida Hampe. «.-...:-ssass-chavatees 176
m= Ayana. SOSI., ..00itSy-s ck -osacacheots=s> 59
= hyrcanica. SOSM...-......0.cserssentss- 59
Ee IPericgd Aalct. .;)iectie aeons 23
== ijberica M(B. <...)...cztt. aneeans 18
— iberica var. Bungeana Trautv. ...23
— — var. elata SOSD. ........csceeeeeeee 18
— — var. minor BOiSS.........:ceeee 18
— — var. minor Medved. ................ 23
— — var. tomentosa Sosn. ..........0.. 18
— jailemsis Zefir. .........cccesceecesseeeeees 47
— 'Kilatbelit DOskkia Aw fed: 20
— Kochiana Sos. ........:ccccceeccceeeeeees 41
— Linczevskyi Fed. ................. 48, 867
— lithuanica BESS. ........c.ccceeceeeeees 64
— lithuanica (DC.) Trautv. .............. 64
— macroglossa Somm. and Lev. .....54
— maris-nigri Fed. ..............++ 41, 866
== ‘maritimay Gere.: aateienees warts. 10
== maritimayPalls ska eins stesso 36
— markhotensis Fed. ............... 42, 867
— Marschalliana Willd. ................ 30
— — var. subglabrescens DC......... 32
— — var. typica Trauty. ............ 30
— — B. Rudolphiana C.A. Mey. ....32
— melanoloma Trautv. ............0.. 57
— microcephala (Schrenk) B.
Fedtsch, ..:.:..\;.:.ne.F awn: 65
== millefolia Wok: upsjeees seawies 348
— monantha Willd. ...........0..cc 43
— monantha var. orientalis Zefir. ..45
—— MONIANG AUCL.. ....00-atervteadospeeyernecte> 25

909

Se BUSI ZS 5 iet bedicgncs tae s-cpeans nanan 20
S= = GEOL BI... cetrasesoticcans onscreen 18
—_ MONLANA Ly werseeeceesseereerseesees 9. 23, 26
— multicaulis Boiss. and Heldr. .....35
— Neilreichii Ortmann. ................. 36
== nivalis! Adams... nif, Bees 18
S= MODIS... covesvcsieephnistetn tan 67
— odontostephana BOiSS. ........:00000+ 64
— ormenidides BOiSS. ........:1ceeeee 60
— pallescens auct. .........ccccecerceseeeeee 41
— pterygantha C. Koch. .......cceeeees 37
— petraed TeMore. ........sccceeseseeseeseeeee 20
— platy glossa AUct. w..ccccccccseseeerersenees 54
— platyglossa C. Koch. ..... 10, 54, 55
= POntica” auctyimecr.c) Grewia 36
— ptarmicaefolia Adam. .........++. 109
— ptarmiciformis C. Koch. .............. 23
— pterygantha C. Koch... a7
— TIZESCENS AUC. .......ccceceereeeeees 48, 57
== ‘rigescens "Willd ys..c.s.0,h3cheu 46
— var. albo-tomentosa C. Koch. ....56
— — var. collina Woron............00 52
— — var. genuina C. Koch............. 46
— — var. latifolia SOSD. .........000 46
— — var. ochroleuca Somm. and
Levis... Ae feu eaiyts 8 Dah oe creates 53
— Rudolphiana Adam. .............::000++ 31
— — var. pumila SOSD. «2... 31
-——ruthenica!: M B.iicisd oitescti tes crectahe 36
— Sachokiand SOSD. .......:cs:esceceeeeees 48
— SAguraMica SOSD. .......ccccseseeseneeeees 42
— Saportana AID. .........eceecesceeneeees 17
SS SANALLISW AUCH tnd. 88 A Sn atecepetpn eee 25
==jsaxatilis (DGS ..2..04.$15..95 apt Sel 26
— Schischkiniana Fed. ............ 55, 870
— Sosnovskyana Fed. ............:.ccec0 31
SeeySterilis Stev...\c..tscrccoscssccnsccennpenenteee 25
=== SHITIACA sVESt. (-:24 coches Sh is casteestean en 20
— subtinctoria Dobrocz ............2.00 41
— talyschensis Fed. ................ 53, 869
— Tempskyana Freyn and Sint. ...... 27
— tenuifolia Schur. .......:..sccceeeeee 123
SA FINCLOT IA ‘AUCtINARG..c.ansessorcassnassestes 41
—=tinctotia Wave. :oekt. Wah the 39
saj——<vara BiuM.B.. 2:.ccnmmta ie 43
— — var. bipinnatisecta Trautv. ....55
— — var. chrysantha Trautv. .......... 41
— — var. melanoloma Trautv......... 57
— — var. monantha DC. ......:s.1000+ 43
— — var. rigescens Schmalh. ..46, 52

var. Triumfettii Trautv. ..........46
var. uniflora M.B. ....c..sseeeee 43
B. floribus pallidis C. Koch. 41
— B. macroglossa C. Koch........ 41

— f. floribus pallide luteis var.

Albidis\ AlDspantineaeSeaueed veces. a5
=" tinctoria © A» Cotulla siii..208. ce. 40
— Tranzscheliana Fed. ............ 27, 866
— Triuumfettii auct. .... 46, 47, 52, 53
— Trotzkiana Claus. .................0 29
— Wiedemanniana Fisch. and Mey. 60
— Woronowii Sosm. ...........:ccccereee 52
— zangeland SOSM. ........csceeesee 59
— zangelanica SOs. .........ssesseee 59
— Zephyrovii Dobrocz. .........::0:0 45
— zyghia Worom. ...........cceeeee 56
Arctanthemum Tzvel., sect. .. 384, 879
Arctica Maguire, subgen. ................ 657
Arctotidaceae Bessey, fam.............. 861
Arctotideae Cass., tribus. ................ 861
Argyranthemum Webb ex Sch. Bip. .

SISA GMb POA EN 129
— frutescens (L.) Sch. Bip. ........... 130
ANF ni ca Tce hss RAS: SERA 655
—alpina.Aucts 2A lAt...s ee 658
Sh ——, Less); Seti RR NL 660
— alpina (L.) Olin and Ladan. ......657
— subsp. genuina Maguire. ........... 657
— — ssp. Iljinii Maguire. .............. 658
— — subsp. intermedia (Turcz.)

Maginire :1..ict ccs... JEX....252 659
ERD a Willd bas. canta. 2 Ren 663
ED liAiGa py TUL Ziyso-..2t0h eA 672
— angustifolia DC. ..........+ 657, 659
See an AAA ed stad.. RNAV ieee cb have ncese 658
— — Vahl B. Lessingii Terr. and

STB pages BS pubtessths-nignnarsevevessoeenseces ont 660
— auriculata Willd. ...............0.08 855
— austriaca Hoppe. .........:.scsseee 676
— Chamissonis F. Schmidt. ........... 668
— — Bf. sachalinensis Rgl.............. 668
ES. Clussit@AMl eens s ARM... 674
SAP THAUSi |: AABRSAG AN. 674
— frigida Meye. ..........scsceeee 660
— helvetica G. Dom... 663
— Iljinii (Maguire) Ijin. ................ 658
—intermedia Turez, ......20..iciie 659
— Langsdorffiana Fisch. ................. 666
— Lessingii Green. ............0..be0 665
— Lessingii Trautv. «0.0... 660
— louseana Farr. ssp. frigida (CAM

ex Iljin) Maguire. ..........cee 660
BEE MATIN Alay Nevavatve- te. ee 723
=l'mollis, Hookantne lene. 667
S“Hmontana - Lin iavsccves ese sses. tte 663
— montana alpind L. «12... 657
— — Bf. stenophylla Ldb................ 658

— B. stenophylla L. «1... 659

910

863

— obtusifolia LESS. .........00...ccceeceees 666
— plantaginisfolia Gilib.............. 663
— sachalinensis (Rgl.) A. Gray. ...668
— scorpioides Adams. ..............0+ 855
Sh BAW. ecco cossscisnentdscnescnssccnees 675
=='Tschonoskyi Ijin. ...2.1....a262¢ 667
— unalaschcensis Less.i....i. ase: 666

— a. minor seu genuina Herder. ..668
— — B. major (pedalis) Herder. ...668

— — f. petiolata Iljin. .............0... 666
-+— f. scaposa B: Fedtsch. ».:...2:. 668
Aronicum: Neck ix..265...1%1-su825 669
(AAronicum) Neck., S€ct.2)-2b.ns0y.:< 671
Aronicum altaicum DC. .........000 672
— carpaticum (Griseb. and Schenk)
Schiuttss.sssncacacas RR eel 675
— scorpioides var. carpaticum
Griseb. and Schenk. .................. 675
Artanacetum Rzazade. ...........:c00+8 $24
Artanacetum (Rzazade) Poljak., sect. .
Lecnunniannosnssadesancdies od RAO <a teehee 524
Artanacetum fasciculatum (MB.)
Rzazade......cbassice-. Shaenosnanad $24
Artemisia) Ls,..4.21524..eeieandahien 425
Artemisia SUDZEM. ............sseseeseeeseess 426
PEP SEC, ionnoiioin< RM tA tees 437
Artemisia sect. Abrotanum subsect.
Polycarpicae. DC, ..:..+...<....ddensas 437
£\fabrotanum) (ist. aiieists <A... see 456
=“sabsinthium, L.c2ett.. 2naeeaeieee 515
=+2Adamsii. BeSS..<.....sti4. sete sreates 463
— akbaitalensis O. Fedtsch. .......... 518
=‘ albicaulis’ Nevskils..i3..asvesteed 614
— /albiceratasKiraschjsssue..aaeony 542
PAP aida ACh wig sunks-asebexaanenassosnndad 582
i2talbida, Lab. anekueeth. Atta 585
+"alpinaPall......issvistscstassede. iiss. .cvee 492
£22 altaica Destth. faswinahe. 28%. <« 461
<Yaltaiensis) KrasGhi isc... cy. .i 514
— amoena Poljak;.i.:......0.4...tii.ces 600
© anethifolia, WeD....2:<....0ae ssw 522
— anethifolia Stelleriana var.
erectiflora DC: ...n:.ncniiinns 523
— anethifolia Fisch. var. macrocephala
F...Schmidt.....:0::208..castnssstt 520
— — Stelleriana var. multicaulis DC.
soshtevckesosasceuibesacadteeetoe ote eee EEE 523
£ annua, L.bewea teeters 489
SSP ANNUA TLAULV ncancidndeshereecteetees 488
==*aralensis,, Kraschitias...2.0i22seeee 594
£35 araraticaKraschtevis...cger sve 553
=2saraxinad Vakbtivictsias.. 2% -csie 620
=< arctica Ress is...28-.citeneets 479

864

— — ssp. subarctica (Krasch.) Hult.. — campestris var. bottnica Hartm......

(Noevopokr:): Poljake.(2.002...05... $54

Pe Oe Lib se as Poa eee 479 snag iaesnantnpsineandnpbiacaanesiht ie: CART EN
— — var. beringensis Hult............ 479 — — var. macilenta Maxim. ......... 550
— — var. ochotensis (Bess.) Kom. .. — campestris var. Marschalliana
ar, Re Se RRB ES A 479 (Spreng.) Poljak. .........uceee 554
—arenania DCi..aii.igdesnndee 540 — — var. sericophylla (Rupr.) Poljak.
—lareliatia.varPschemievianaiii’— od Walt ed. Acer. easiness 554
(Bess) Grossht's2..00 on et.X 541 — — var. Sosnovskyi (Novopokr.)
— arenicola Poljak. ...........0.0000000 619 PON AK. oat sonanne --tghhta. ke 554
— armeniaca Lam. ............cce eee 471 — canescens Willd. ..........ccccesceeeeeee 471
argentatum KlOK. ........:.000ceeee 493 — capillaris Thunb. ...........cssisce 550
Argyi Lev. and Van. ..........0000. 451 — — var. sachalinensis Pamp. .....550
argyrophylla Lab. ......0......cc0 497 — — var. simplex Maxim.............. 550
Aschurbajevii Winkl. ............0..... 503 — capillifolia Fisch. «0.01.0... 513
astrachanica Poljak. ............000.. 579 = caspica Kell. 232332 anes 579
athanasia* Bess). ee 406 — caucasica Willd. 0.0... eee 492
auratawKome tn. 18s. hen eis 492 — chamaemelifolia Vill. .........00..... 458
austiriaca: Jaeqeviac.. ian ..daaint 498 — chamaemelifolia y. armeniaca
— var. barabensis Serg. ............ 499 Hohenack!s...f52..1...15.203.seekeie 458
— var. neglecta Krasch. ............ 499 — chamissoniana BESS. ..........-.0006 479
badhysi«Kraschiwineteteae 619 — chamomilla Wink. ................... 490
— var. arenicola Poljak.............619 — chevsurica Somm. and Lev....... 493
balchanorum Krasch. .................. 617 — chrysostachys Rech. f. and KGie. ..
baldshuanica\Krasch:andeZapri624° ==)... 2. Se, cee. ee 317
balsamita Willd. .........000.:c0000 461 SICING, Berga iiigsct.shcih0l.. Jaco 601
bargusinensis Spreng. ................. 555 — cina Berg ex Poljak..............0 601
Besseriana var. integrifolia — — var. mogoltavica Poljak......603
Ledbsit8.. 22h)... HL ae 504 — ciniformis Krasch. and M. Pop.603
— var. triloba Ldb. ...........004.. 504 —cimoides |Pavl.........scvsdict-.-sesttstae 603
Billardierana’ Bess. ..cwisenient 622 — commutata BESS. ............:csseceses 551
bipetata. Gilib. 2A eee 515 — — var. bottnica Krasch.............. 553
borealiskPanl fs. ..2e. cae es $57 — — var. dolosa (Krasch.) Poljak. ...
borealis Willdenovii Bess. ........ S95" OOO ee eee ace ee TS!
— ssp. bottnica (Lundstr.) Hult. ... — — var. Gebleriana Bess. ......... $51
BRE RE LOE LE 553 — — — var. Gebleriana Bess.......552
— var. Ammaniana Bess........... 551 — — — var. Helmiana Bess...............
— var. Ledebourii Fedtsch. ...... 559° > BOe....2, SA gracile, 550, 551
— var. Mertensii Bess............... 551 — — — var. Helmiana Bess. ........ S52
— var. Purshii Bess. .................. 557 — — var. pubescens (Ldb.) Poljak.
—= Var sspithamacar(Pirsh)yTomey® WO ocecarcasroapereossnenacnarsan threes §5,159552
anid} Gray 20. ....5 ae eee 557 — compacta Fisch. ............cceeeeee 585
— 68. Ledebouri Ldb. ................. 555 — contra Schachm. ..........cceceeeee 601
Boschniakiana (Bess.) DC. ....... 573 — corymbosa FASCH. .........cccceeeeeeee 482
brachanthemoides Winkl........... 424 — curiliensis Spreng. .........:00000 482
brachyloba auct. 00... 448 — cuspidata Krasch. .........cccssee 500
cdespitosal dbo emia 499 — Czekanovskiana Traut. ............. 502
camelorum Krasch. ............c..0+ 595 — daghestanica Krasch. ................-. 535
camtschatica Schlechtd.............. 557 911 — demissa Krasch. .......cccceeeeeeee 562
campestris D1 .23..2. ARR 552 — dentata WiIId. 0.0.0... 508
campestris Maxim. .......ccccecee 551 — depauperata Krasch. ............00 559
— ssp. borealis Hei. ................ 537 =—Ndeserii KraseM. ....cs..-sa!-. acess aaceues 626
— ssp. spithamaea Hall. ........... 557 — desertorum Spreng. ............00000 549
campestris var. araratica — — var. macilenta Pampan. ....... 550

— var. macrocephala Franch. ..530

— — var. Sprengeliana Bess. .......549
— — var. Sprengeliana Bess. f.

Gebleriana (Bess.) Pampan. .....551
— — var. Sprengeliana Bess. f.
Helmiana (Bess.) Pampan. ........ 551
— desertorum Takeda. ............0006 546
= ldiffusanKirascht inane. Lhe 612
— Dimoana M. Pop. ...........:ceee 536
— divaricata (Grossh.) Rzazade. ..576
=dolosa. Krascht 22 R2ehan...0. 2 551
— dracunculus L. ou... cece 529
— — var. humilis Kryl. ................. 533
— — var. pilosa Krasch. ................ 533
— — var. pratorum Krasch............ 530
— — var. Redovskyi Ldb. ............. 533
— — var. turkestanica Krasch. .....533
— dracunculiformis Krasch. ........... 534
— Dubjanskyana Krasch. ................ 587
— dumosa Poljak. ..........:ccceeeees 621
— eldarica Rzazade. .......0......c08 553
— eranthema Be. ...........ccseseeeeres 635
— eremophila Krasch. ..........:::000008 612
= leriOCArTpANB Re. seine ake 631
— erivanica Grossh. ..........::seesee 576
— Falconeri Clarke. ............:0c0c008 505
= fasciculata) MBYS)..2220 ans 524
— — var. armeniaca Bess. ............ 524
— — var. iberica BESS. ........:c008 524
— fastigiata Wink. ...........0...00000 405
— Fedorovii Rzazade. ............:00+6 576
— Fedtschenkoana Krasch. ............. 600
— ferganensis Krasch. .............0+ 607
— fragrans BOIrMM. ..........cccseeeee 608
Ss fragrans, WilldM..2icneeen: 576
— fragrans Willd. ssp. gurganica
Keraseht. ...2.0ee es. A Le 576
— — var. erivanica Bess. .............. 577
— — var. gurganica Krasch. ......... S77
— — var. phyllostachys Boiss. .....577
— fragrans vat. ramaracandica
Boissses... Sl Se aes: 608
— — Willd. B. plyllostachys Boiss....
oaats edad Sete GRA heer: 576
— Freyniana (Pamp.) Krasch. ........ 467
—ufrigida) Eichwegs.® anak ee. 498
SSifrigida), Wilds c.ccccccsncomcctat 494
— — ssp. Gmeliniana (Bess.) Krasch.
, saabda cde Tasos SO La asec neeee Nee 497
— — ssp. parva Krasch. ............ 497
— — ssp. Willdenoviana (Bess.)
Kiraschis..iicciieteie kee ee 497
— — var. argyrophylla (Ldb.) Kryl...
a asasbcetousne sos tRO cc RU RMON MRE MES 497

— — var. argyrophylla Trautv. ....497

865

— — var. foliis magis dissectis Ldb.

wea despecceusteieasndtan same atest Meee 498
— — var. Gmeliniana (Bess.) Poljak.
sutnenetatdavestueettetexeceretceetecarcse RC 497
— — var. intermedia Trautv. ........ 497
— — var. parva (Krasch.) Poljak. 497
— — var. typica Trautvy. ............... 497
— — a. Willdenoviana Bess. ........ 497
— fruticosa WAM. «0.0... ceeeeeeee 540
— furcatayMB®)...c2inkt... 483
— gallica Willd. ..........cccceeeeeeseeeee 573
— gigantea Kitamura. ...............0.. 452
— — var. shikotanensis (Kitam.)
PALE WAI osc sescccennseecds-tceatnackabeacsous 452
— glabella Kar. and Kir. ............... 510
— glanduligera Krasch. .............0 604
=elauca Krascht i532. Ants tncsis 598
= eglaucina: Palle vcnctce,.1geeneee 535
— globularia Bess. ...........ccccceeeeee 481
= glomerata: Ldbicctsd...32.. cites 482
— glomerata Rydb. .......:.ccsseeeeees 485
— — ssp. curiliensis (Spreng.)
Krasch. 22 22d. aes, ARG. 482
— Gmelini Turezinn neath... 465
— Gmelinii Web. ............. ccs 464

— — var. Biebersteiniana Bess. ...465
— — var. intermedia (Ldb.) Krasch. .

inc ARES SOON. ize 464
— — var. legitima Bess. ..............+. 464
— — var. Turtschaninovianum
BR ESS. .ccccsbaisc.-..deeeeeeee. See ae 465
— Gorjaevii Poljak. ........ccssee 596
— gracilescens Krasch. ...........:.0+ 591
— Griffithiana BOISS. ..........:00 518
— Grigorievi Krasch. ........0csce0 598
— Grossheimii Krasch. ..........::0000+ 493
— gypsacea Krasch. ..........c:sseeees 610
— halodendron TurczZ. ............2:000+ 539
— halophila Krasch. ............cee 581
— Hanseniana Grossh........... 575, 576
— — var. divaricata Grossh. ........ 576
— — var. phyllostachys (Boiss.)
GFOSSHS..:...0....50. See Ate eee 576
— Henriettae Krasch. ...............0 558
— heptapotamica Poljak................. 592
— herba alba ASSO. ....0......ccceeeeeees 622
— — ssp. Kemrudica Krasch. ....... 614
— — ssp. saxicola Krasch. ............ 621
— — var. densiflora Boiss. ........... 622
— — var. laxiflora Boiss............... 621
— herbacea Willd. ............cccceceeeees 457
912 — heterophylla BesS. ........2...::0000+ 483
— hololeuca MB. ...............:sccssseenees 507
— holosericea O., PB. ......s.ceseees 501

866

—humilisvMB: oye MaRA ee 579
=Shncana: (Liz) Drees esecscdieseecccee 524
— INOdOTA AUCL. ........cecceeeeteeteereeeee 553
28 {e=— Gab: 2 ee ee 540
— — MB. ssp. tomentella (Trautv.)

Kiasehit.:é <tacwx2h). gk ese: 554
SS Wilde cee oss 530
— insulana Krasch. ..........c.eee 484
= inteorifolia Lc... 21a 449
— — var. stolonifera (Maxim.)

Pam paris ..0.0.12,.. Se all: 446
— — var. subulata Pampan. ........ 450
— — var. typica f. genuina Pampan.

SES EEE Pest atiaenc PASM. 449
— Iskenderiana Rzazade. ..... 575, 576
— Issaevi Rzazade. 0.0... 621
— issykkulensis Poljak. .........:.:000+ 600
— iwayomogi Kitamura. ................. 464
+3 jacutica: Drob. <2). ie 522
— japonica F. Schmidt. ........00000... 488
— japonica Thunb. ..........ccce eee 545
— — var. desertorum Maxim........ 549
— — var. japonica Maxim............ 545
— — var. macrocephala f.

sachalinensis Pampan. ............... 546
— — var. rotundifolia d. vestita

Pa iaapraits Giest.t. eacercweeeoe tet inccetpiee- 222 546
— — Q. japonica Maxim............... 545
— — B. manchurica Kom.............. 549
— — f. typica Nakai. ........0.0. 545
— jeniseensis Spreng. ........:.c00 497
— juncea Kar. and Kir.................. 629
— — var. macrosciadia Poljak ......629
— karatavica Krasch. and Abol....598
— kaschgarica Krasch. ..............+ 599
— Keiskeana Miquel. .............00005 487
= Kelleri, Krasch). inne anne 537
— kemrudica Krasch. ...........0.000 614
— Knorringiana Krasch. ................. 588
— kobstanica Rzazade. ..............0 574
— kochiiformis Krasch. ................. 615
— Koidzumii Nakai. .............c 447
= \Koidzumit Nakai. .....0..2:. Ree 439
— Komarovii Poljak. ...........0.00 449
— kopetdaghensis Krasch. .............. 617
— Korovinii Poljak. .........ccccsscees 589
— korotkyi Krasch. «0.0.00... 523
— Korshinskyi Krasch. ..............0: 622
— Kotzebuensis Bess. ...........00000 324
— Krascheninnikoviana Bess.469, 472
— Kruhsiana BESS. .......:.sccseeseeseeee 504
— Kryloviana Steinb. .........00..000. 518
— kulbadica Boiss. ............ccceeeeee 519
— Kuschakewiczi Winkl................. 544 913

— laciniata Willd. «00.0... 473
— — var. glabriuscula f. marginata

Pampantei 2 aats canes 474

— — var. latifolia f. Maximoviczii

Pamipane ice 4 22st. Qe 469

— — a. glabriuscula Ldb. .. 469, 473
— —6. Turzaninowiana Bess. .......474
— laciniata » Turezaninowii DC. 474

— — f. paniculata Kryl. ............ 473
— — f. racemosa Kyl. .......:c0000 474
— — f. tomentosa Kry\l. «0.0... 474
— laciniatiformis Kom. ..............0. 477
— lagocephala (Bess.) DC. ............ 504

— — var. Besseriana (Bess.)

Pampan. f. a. integrifolia (Ldb.)

Pampanterd 2 AciennIus Reh 504
— — var. Kruhsiana (Bess.) Glehn. ..
SRY RES SRR 504
— lagopus Fisch. ...........ccsceeseeee 486
— — ssp. abbreviata Krasch. ........ 486
= lanata*DC...2.5.. 2g 8 2a 492
= ilatifolia! Ldb 2easiin hee 472
— — f. pilosiuscula Krasch........... 472
— latifolia Maxim. ...........0ccccce 468
— lavandulaefolia Nakai................ 450
— lavandulifolia DC. ..........c cee 453
— Ledebouriana Bess. .............00006 555
— Lehmanniana Bg¢e. ..............00+ 622
— — ssp. leucotricha Krasch. ...... 622
— — var. Korshinskyi (Krasch.)
Poljak. (2.2020. eee 623
— — var. Skorniakovii (Winkl.)
Poljake.scc.:ésscscccescc ROG. PRE 623
— leontopodioides Fisch. ................ 485
— Lercheana auct. ..........:cscceseeseeeee 580
SS Edbis.. J ae ae 591
— Lercheana WeD. .0.....::c0ccc0ceeeeees 579
— — var. astrachanica Poljak. ...... 579
— — var. humilis DC. .........c 580
— Lessingiana Bess. ...........::0cseee 594
— Lessingiand Ky). ........ccccceeeees 596
— leucodes Schrenk. ............:::00 630
— leucophylla auct. ......:.ccccceeeeeee 451
— leucophylla Turcz. ...........0:c: 443
— — var. typica f. genuina Pampan .
EE a aia 443
—= ‘limosa’ Kotdz: sania. ikea 559
SAPapsleyi. Paljake) cissutevienssvesesesvoite 544
— lithophilla TurezZ. ........ccccccceees 504
— littoricola Kitam. .............:::ce8 546
—Slobelii, AMES tasks oeee 458
— lobulifolia Boiss. ...........:cccc 625
— longepedunculata Bess. ............. 479

— macilenta (Maxim.) Krasch....... 550

macranthawLdbwitl eee 462
macrobotrys Lab. ............-. 469, 471
macrocephala Jacq. ........::-1:1+ 518
macrorhiza’ Ture7 yt ee... tee..:. 478
macrosciadia Poljak. .............+++ 629
maikara (Krasch.) Pavl.............. 590
maracandica Be. .......ce 608
maritima Bess. terrae albae
| acto) Ee aE CELL A Be Bk 592
— ssp. incana (Kell.) Krasch
bikcbereesaa chs tresstee ce rete Meee 579, 580
— var. n. Lercheana f. dahurica
CMT CZ ek tececesshasdeeoeteeh eee 580
— f. fragrans Rgl ....... eee 601
aE 7: |; er Pe ee tie ke A 582
Aiaritimiaets.<).. 2S EE. Svs
maritima Stechm ..............::-0+0++ 573
— astrachanica Kazak. .............579
E—.« pernimealsd bias: szsreacesss=tsesncseree 573
— Linnaeana Bess: ...........:0:...... 573
maritima salina Kell. ................ 574
— f. citriodora Kazakevicz. .....574
— ser. sublessingiana Kell....... 596
— ssp. Gmeliniana (Bess.) Krasch.
tbat jsdbeetavgapenaebabwcabadesaannaacatoeerees 580
— ssp. maritima (L.) Hegi. ...... 573
— ssp. monogyna (Waldst. and
ISIS) WED@ Olt teasycaepccrscusenasceasetzeasesast> 574
— var. Blumiana Bess..............- 574
— var. Boschniakiana Bess. .....574
== var. erivanica BESS. .........:.... 576
— var. fragrans Ldb.................- 576
— var. genuina Kryl........ 573, 580
— var. Hanseniana Bess. ........- 575
— var, iIncana Welly 22.0...0e 8... 579
— var. Kitaibelliana Bess. ....... 574
— var. Lercheana Bess. ...........- 579
— var. Lercheana Ldb. ............. 579
— var. Lercheana f. Gmeliniana
Led Decay sisteecccts etn ee 580
— var. monogyna Fedtsch. ....... 574
— var. monogyna Ldb.............+. 574
— var. palmata Lam. ............4... 574
— var. pauciflora Ldb. ............. 590
— var. persica f. Hanseniana Ldb.
soctibathea seaestecee he aR a ee 576
==" VanwRossica Bess. \ieice-.ee 579
— var. salina (Willd.) Koch. ....574
— var. Szowitziana Bess........... 55
— var. Wilhelmsiana Bess........ 574
— var: Vv. taurica Lab: See 578
— — var. €. genuina f. tt. suberecta
Lb.) ..0t. AO eae Re 580
— — var. X. Gmeliniana Bess......580 914

867

Artemisia maritima var. v. Meyeriana

BOSS es idssessveasecsagterootesteerieree ee 576
— 6. Stephaniana Bess. ............ 585
— €. compacta LAD. ............+ 585
— 7. Fischeriana Bess. ............ 585
— 1. Helmiana Bess. ............... 585
— wu. Boschniakiana. Bess. ......573
— v. Stechmanniana Bess. ....... 589
— «s». Lercheana humilis Ldb. ....
MaDe sectetccetsveavansssteetlds foe RMIT AES 580
Marschalliana Spreng. ............... 553
Martjanovii Krasch. .............006 O13
matricarioides LESS. .........::0++ 150
Meximovicziana Krasch. ............ 469
madioxima Krasch. ............:.0++ 468
megacephala Rup. ........:0008 413
Messerschmidtiana Bess. ........... 464
— var. viridis. BESS. .....c:0..08. 464
Meyeriana BESS. .........:eceeeee 486
Ee ACGTOSSH ss cous santastestectstich cotcseseeve 576
— var. divaricata Grossh. ........ 576
mogoltavica Poljak. .........:10-+ 601
mongolica Fisch. ..............ssscs++ 444
— var. tenuifolia f. genuina

Pam pan fii. bite see tsistsaee chewceneen oes 444
mongolorum Krasch. .........0-+++ 580
monogyna Waldst. and Kit. ...... 574
monostachya Be. ......seseeeee 551
montana Pampan. ...........sseee 452
montana Schlecht...........ceee 479
mucronulata Poljak. ........000+ 598
multifida Kar. and Kit. .............. 629
mutellina S.G. Gmel. ...........0++ 494
multicaulis LD. .........0ccccsceeeees 523
nachitschevanica Rzazade.575, 576
namanganica Poljak. ............0++ 611
Nitens (BeSS.) St€V....0i2...secevendees 502
Mitrosa WEBMAG «02s ccissthercesths teens 580
— var. gobica Krasch...... 580, 581
—— var. subglabra Krasch........... 580
nivea Redowsky. .......:.s:ceseeeeee 498
NOTVEZICA AUCL. ....ccecceserereesesesenees 480
NOTVEBICA FTIES. .......s0sseseeeeeererseees 480
norvegicad Halll. ........ssseceeseeees 479
NOTVEQICA SPTENG. «....-seeereeseeeesees 479
norvegica globularia Hall......... 481
— glomerata Hall............1s-00+ 482
— pacifica A. GTay, «1... 479
— ssp. heterophylla Hall. ......... 484
— var. uralensis Rupt. .........-.+ 480
— var. villosula Trautv. ............ 480
mutans Nakai acco cine se tncrerest 451
nutantiflora KOM. .......:.:-0eeee 446
nutans WANG. cceeses.<. tse eee 574

868

— obtusiloba Kyl. ......ccececeeees 514
— obtusiloba Ldb.............. eee 509
— — var. glabra LAD. ......cce 509
— — Q. gracilis Ldb. «0... 509
— — B. fruticosa Lab. 0.0.0 514
— Olliveriana BeSS. ........ccseeseeeee 622
— opulenta Pampan. ............c:ccee 439
— opulenta Pampan. f. laciniata

PAM Panhys cevsish.-retecactatratcih.oscaidea.v- 440
SA DTLCTGUS. WU ora ssensnttascnnre ess 524
— Pallasiana Fisch. ex Bess......... 402
= (Pallasit Willdtssasok.s6ssGus.8-.8sc02s 555
— pallida Poljak. ...... cece 612
— palmata Lam. .......cccceceereseeeeee 573
ssipalusttis. L.. ....ccmen ak. -nesrtstcacese-s 491
— pamirica Wink. ...........cccsee eee 534
— — var. Aschurbajevii Winkl.....534
— paniculata BESS. .........cceeseeeee 560
5 EAM, «ccacennnca ns sczsatpee teciventcsvath 456
— pannosa Krasch. ..........cccseeeeees 556
— pauciflora MB. ............4.. 537, 540
— — B. cernua BESS. «0.0... 538
S35 -— Kryl. .......:...-stsiteheometniareren 591
— pauciflora Web. .........ccceeees 589
— — ssp. maikara Krasch. ............ 590
— — f. maikara (Krasch.) Poljak. 590
— paucifloriformis Rzazade........... 574
== pectinata. Pall. soi. gp tvaayessonk.y 637
— — var. yunnanensis Pampan. ...638
— peduncularis MB. ..........:.:00++ 494
— pedunculata Steudel. .................. 494
== Persica, BOISS. tars.-.2558-oh-tahans 506
— phaeolepis Krasch. ...........:.c0000+ 473
— phyllostachys (Boiss.) Woron. ..576
— polysthicha Poljak. .........:.:cc08 596
= (pontica’ L. .AeteS, caeearncree a 461
= potrecta Krasch)) ..:.ss083. cases 607
— potentillaefolia Fisch. .............. 471
— prasina Krasch. ........ccccseseseeee 620
— Prilipkoana Rzazade. ................. 575
— procera Willd. .....cccceceeseceeeeee 456
— proceraeformis Krasch............... 457
— prolixa Krasch. .......:..csssssseseeseees 587
= ipubescens LAD. -..41 aid swe beevaieess 551
= ipunctata Bess. «iisysed--95%yscpenweenee 469
— — B. stricta Bess. ween 472
— punctigera Krasch. .........cesce 480
— pycnorhiza Ldb. ...........ceeeeeeee 559
— quinqueloba Trautyv. ........c. 540
— rhodantha Rup. ...........ccseeeeees 586
— Redowskyi Ldb. .......cccceccceeeeeees 530
— TEMOSA SUZAW. ......ccsssesessereasencseee D357
— remotiloba Krasch. ...........:ee 470
= repens, Pall; ....cc:ssostsbeees «aseteys 498

915

rubripes Nakadi.......it..1.2daaeenns 450
Tupesttis .L...,.:.<ctd seen one 508
— var. oelandica Bess. .............. 508
— var. thuringiaca Bess............ 508
— var. Willdenoviana Bess. .....509
— var. v. Willdenoviana Bess. 508
— f. alpina Kryl.......cccccee 509
— f. salsuginea Kryl. «0.0.0... 509
=— QO. FE. Muelll,., .jeoccds.2s.-s0tteeeben 480
Se ls ee ee ee 493
Ruprechtit Polak: wisis.ccssssscseescerss 480
Fulifolia <Stepli.wdcis,)...ce-.tGiesnnn 505
sabulosa,, Willd, ..:.s.4ciet-+..eptee 540
sachalinensis, Tiles: wisunvs..4--00 550
SACKOPUM MAD: ssencepeysriceanseen a 464
— var. intermedia f. incana Ldb.
ais sacuancssidpacedhs cic eaRvEmee \acemeieaeale 464
— var. intermedia Ldb. f. nivea
| ee er eo, ane 464
— var. latiloba Ldb. f. e.
Freyniana pampan. ..........0:-..0+ 467
— var. latiloba Ldb. f. incana
Pampan yt .teseiecee tights. tpgete ans 464
— var. latiloba Ldb. f. intermedia
Pam pan 1.0250, Fann: Rea Rieck ani 464
— var. minor Ldb. .......0......000006 465
— var. minor Ldb. f. vestita Kom.
ee ete ee Bec 464

— var. santolinifolia Pampan. .465
var. viridis f. minor Ereyn. .467

— B. intermedia Ldb. .............. 464
Saitoana Kitamura. .............:00. 448
Salina Kell. te... cictcm ee --camiine 574
= Wd et cctaxesd teeta NE cc ayeivns 573
— ssp. monogyna SagorskKi. .....574
— ssp. monogyna var. Kitaibelii

SA OTS Kise, Meuse eta hcesan dp roces 574
SalsoloidesWilld..<4.<%.2.-ay--:, 538
SQINGNUSIGA AUCH ia .fos05..5...006oussens 447
ee ee. SEES 438
samoiedorum Pampan................. 520
santolina Schrenk. ..............::0.+++ 625
santolinifolia Turcz. .............0.+ 465
santonica Waldst. and Kit. ....... 574
— var. monogyna Fritsch. ........ 574
Saposhnikovii Krasch. ................ 542
Schischkini Krasch ys \c.,i.-.7 5015-000 580
Schmidtiana Maxim.............0.+. 501
Schrenkiand Edbycsvcsscit:.-/sseus:eu. 582
scopacformisn LaDy iy isees. cvises acc. 581
SCOPATIA Maxim, ......0...00ceseeeeeees 550
— W. and K......... Ie is vie ais 560

— var. scopariiformis (M. Pop.)
Poli abies f--s:scaie ion es tei ayia son 561

scopariaeformis M. Pop. ........-.- 560
scoparioides Grossh. .................;560
scotinavNevski.Wiveais.. Runa 614
selengensiseTurez.c:918tis ORE 454
selengensis lusus umbrosa Rgl. 453
— B. umbrosa Lab. .........0000 453
senecionis Jacq. ex Bess. .......... 414
senjavinensis Bess. .............0.++ 485
sericea F. Schmidt. ..............00. 501

— Stechm. var. Steveniana Bess.
Jee douenuaadbeint BAeaeetaievesscS AT 502
sericea) Web). Ba, lh 501

— var. Gmeliniana Bess. .......... 502
— var. grandiflora DC. ............ 502
— var. Ledebouriana Bess. ...... 502
— var. nitens (Stev.) DC.......... 502
— var. nitens f. parviflora
Pampa ny (2:2... thd. CHB he 502
— var. Pallasiana Bess. ........... 502
— var. parviflora DC. ........00... 502
— var. turkestanica Wink. ...... 503
sericea Web. f. Czekanovskiana
(Drantv:) -Poljakfi2 2282... 2802 $03
— f. nitens (Stev.) Poljak......... $03
— f. Pallasiana (Bess.) Poljak. 503
Sericophyla, Rupr. ..0.s.0+-.-mbies 553
sefotina Bee: 36. .sdnnamadetads 608
shikotenensis Kitamura. ............. 452
sibirica,(L.), Maxims. s2ccacenhss 417
Siebert NOSS, ....:-:-.c1s,.5cd etnies 621
Sieversiana, Willd. ..:...cdsscivissees SZ
— var. pygmaea Kryl. ...........+ 518
— var. septentrionalis Pampan.

wi tihelt. eehlnchs i ahnluainan 522)
silyaticayMaximacancrs tees... 445
Skorniakowi Winkl. ........:cccee 622
Sondiana Bee. anus Reise 616
songarica Schrenk. .............:0 543
Sosnovskyi Krasch. .........:.0000+ 553
specigera). CaK ochisstaiae, ..¥e...00 620
spithamaea Pursh. ........:.scsee Bie /
splendens )Willdiiicstt.. een ictincs. 493
Stechmanniana f. armeniaca Bess.
svaamtauiad. Siawk-- gilebonartees 458
Stelleriana, Bess 4 2s.:%.seyess ..f-s5=: 455
— var. sachalinensis Nakai......455
stenocephala Krasch. ................. 616
stenophylla Kitam. ...........0s000+ 450
stepposa Kell. and Kom............ 574
Steveniana) Bess\s.i05.ss2....cr2r--se0e 482
stolonifera (Maxim.) Kom......... 446
— var. laciniata Nakai. ............446
suaveolens Poljak. .........c.:escee 592

subarcuicatKrascns «s..ccscscccssesesases 479

916

869

sublessingiana (Kell.) Krasch. ..596
— var. Gorjaevii Poljak. ........... 596
— var. polysthichum Poljak. ....597
subulata Nakai: ¢....0.c280.2..208les 450
subviscosa. Turezainne.. Reawae 510
succulenta(Edbaiaus- sceestelse 521

superba; Pampanwalt 2nntiinn a: 438
Szowitziana (Bess.) Grossh....... 575
tanacetifoliasl........x tlw. sktinm 468

tanaitica Klokiac.. eas 538
CAUTICALESS..cocoinses sete he RE 594
taurica: Willd. £2.ctal. RoR 578
— var. erivanica DC. ............... 576
— var. Hanseniana DC............. 576
terrae-albae Krasch. ................... 592
— var. heptapotamica Poljak. ..593
— var. kurdaica Poljak. ............ 593
— var. Massagetovii Krasch. ...592
— var. semiarida Krasch. and

LAE escwcnaascas Athi tet ke Re caeee 592

— — var. suaveolens Poljak.......... 593

tenuisectasNevskionwin...20%.cc0 598
— var. glaucina Poljak.............. 599
— var. karataviensis Poljak. .....598
— var. mucronulata Poljak. ...... 599
tianschanica Krasch. ...........06 585
‘Tilesiiwsdb s2atnr2teks. Ren. .aves 443
togusbulakensis B. Fedtsch....... 506

tomentella Trautv. ..................... 554
Tournefortiana Rchb. .................. 488
; transiliensis Poljak. ...............0 597

— var. boamensis Poljak. ......... 597
Trautvetteriana Bess. .............066 $37
trifurcata ‘uct. 1n%..ciitiienam 484
—=2 Stephin: sienna see 483
Trinigna* Besstain nik... 88. vee 487
Tschernieviana Bess. ........ 540, 541
tufanicaKraseh:h:...001. 2p 612
— var. diffusa Krasch. .............. 613
— var. eremophila Poljak. ........ 613
— var. pallida Poljak................ 613
turcomanica Gndg. .............c..08 618
Turczaninowiana Bess. ............+ 505
— var. altaica Kryl...........000 505

— — var. Falconeri (Clarke) O.

FédtséhacraraunnWis...¥..acneglas 505
— B. dasyantha Schrenk........... 505
turkestanica (Rgl. and Schmalh.)

Franchitn..cicts...2eeedl. walk 317
Turschaninoviana Krasn............ 465
umbrosa Tureg2i). zits 453
— var. typica Pampan. .............. 453
unalaskensis Rydb. ..............0000 440
Ursorim TULA cccciscsesessscsceseses 440

870

— ussuriensis Poljak. ............::000 446
— uzbekistanica Poljak. ...........0+ 611
— vachanica Krasch. .........::ssee 623
=——PValida Krasch. <.. 0.622.742: 610
— venusta Pampan. ..........ceee 450
— verlotorum auct. ........:ccceeeeeeeeeee 439
— verlotorum Hull. ........cccceeeeee 440
— verlotorum Lamotte. ................. 440
sSfvestita Wall. .........). ates 465
= yiolacea. Lab. ...... 0. 557
ES=CVIFIGIS. Willd. oo. .ia..:stessesee- Sesetaler 508
— viridifolia LAD. .........cccccereeees 508
— vulgaris Kom. .........ce+8 447, 453
aL Vl paris Feehan ee. ER acs 438
— vulgaris mongolica Fisch. ......... 444
— — stolonifera lusus b. glabrescens
Reg le eesecseteeesitee - ReOReOhenOe aN odrat 488
— — Wrighti (Gray) Hall and
Clementsy st: wh tveiernin tei 440
— — ssp. Tilesii Hall. ..........0.00.. 443
— — var. communis LOD. .............. 439
— — var. incana Maxim. .............. 451
— — var. incanescens Franch....... 451
— — var. indica f. montana Nakai. ..
wnoyne Saal es ASAE TEE MEN F528 452
— — var. integrifolia Ldb............. 449
— — var. kamtschatica Bess. ....... 439
— — var. kamtschatica Nakai. .....452
— — var. leucophylla Turcz. ........ 443
— — var. parviflora Maxim. ........ 451
— — var. samamisica Maxim. ...... 447
— — var. selengensis Maxim. ...... 454
— — var. selengensis lusus
Serratifolia, Rel. sous titres: 454
— — var. selengensis typica Rgl. 454
— — var. stolonifera Maxim. ....... 446
— — var. stolonifera lusus incana
11k) Ua Ree etn ae AME age st 446
— — var. tenuifolia Turcz............. 444
— — var. Tilesii Lab... 443
— — var. umbrosa BESS. ........6..006 453
— — QO. minor LAD. ...... eee 444
— — a. selengensis f. serratifolia
KOM. 5. 050.0 Seta eT, 3 de ovees 454
— —-y. coarctata Forsell. ............... 444
— vulgaris y. vulgatissima Kom. .451
— — €. leucophylla Turcz............. 444
— — Maxim. ...... eee 447, 453
——, Welbyi. Hemsl: *. 0. 2222....: deat: 539
— Wightii Gray. ........eceseseseereeeeees 440 917
— yunnanensis (Pampan.) Krasch. 638
Asterotricha Tzvel, sect. ........ 358, 878
Athanasia maritima (L.) L.............. 128

Balsamita Mill., gen. ............ 184, 196 .
Balsamita (Mill.) DC., sect. ........... 196
Balsamita balsamita (L.) Rydb......201
=imajor Desf." ath Rania 198
— suaveolens \Pers. ich vavetserint ess 201
— vulgaris Willd. ..........ccccceeeeeeee 201
Balsamitopsis Tzvel., sect.....212, 871
Brachanthemum DC. ............ 390, 880
Brachanthemum S€Ct. ..........:ccceseeees 395
Brachanthemum ser. Eubrachanthema
Kerasche ts of Bote. eu vcacsueaie 395
— ser. Procumbentia Krasch. ........ 395

— ser. Pyrethroidea Krasch. 394, 880
— Baranovii (Krasch. and Poljak.)

Kraschsis :scasradeied spins 394
— fruticulosum (Ldb.) DC. ............ 396
— kasakhorum Krasch. ................... 397
— kirghisorum Krasch. ...............+. 396
— Krascheninnikovii Rubtz. .......... 293
= TitO Vii KraSc hin sautt iavecatiesstepey- «sens 395
Brachyglossa Tzvel., sect. ..... 240, 872
Branicia insignis AndrZ. ...........:000+ 733
Brevilobae Kuprian., sect. .............. 651
Gacaliaiiauct. 2425.) ROR ee 697
Cacalia\L: ssc thm. Bis 683
— sect. Eucacalia DC. ..........0... 683
— aconitifolia Bge. .....ccccceeeeee 698
— aidzuensis KoidZ. ..............::.cs0008 689
Val pinae Vos sesisssssacscssnshivtra deere 684
—atriplicifolia V0 iie%.. aR etae ae 685
— auriculata auct. ..........:..+ 692, 696
— auriculata DOS YaIAak...W SS 694
— auriculata var. alata Nakai. .....692
— — var. kamtschatica auct. ........ 693
— — var. kamtsch. f. alata Kitam. ...

aibssacdunsgsiseduaentacoarede ten te tM 693

— — var. kamtschatica Matsum. .696
— — a. ochotensis Kom. .... 692, 694

— — a. ochotensis Maxim. .......... 688
——— Bultensiccuisaiks.... oars 696
— auriculata Hulten x C. hastata L.
FAL REE ET REREAD EL PRP OR UE 688
— farfaraefolid AUct. .......cceeee 691
— — Q. FAMOSA AUCL. ......ccecceeeeeeee 691
— glabra ‘Labs Wann. Aa 687
a) hastata-auctonic. tena 689
ma haStata- Liv-issscceveorsen.teavcenureente tts 687
— hastata ssp. hastata Hara. ........ 687
— — ssp. orientalis Kitam. . 690, 691
— — var. orientalis Ohwi. ............ 690
— — a. pubescens Ldb.................. 687

— — Bf. glabra auct. ..........e 689

— — B. glabra Ldb. ................8 687
— — x C. kamtschatica (Maxim.)
Kudoutw a. Lee ela 688

— kamtschatica (Maxim.) Kudo. ..695
— Komaroviana (Pojark.) Pojark. .691

— macrophylla M.B. ..........:000c0 739
— pinnata Willd... 741
— praetermissa (Pojark.) Pojark. ...692
— rhombifolia Willd. ............:000+ 739
=—trobusta. Tolm; jn... 22s. Aa 689
— sagittifolia Mertens. ..............-.. 687
— suaveolens auct. ...........0.4. 687, 696
— suaveolens L. ..................... 684, 687
— Tschonoskii Koidz. ........... 689, 682
Calendula, Lin 2k). 228 0G..288. x: 857
arvensis slost,. 22nanow,..t84..c 859
== arvensisoMB., GAtth.t.). Waeseans 858
— brachyglossa Rup. ........:.:10+00++ 860
<> byzantina\DCY ARRAN 860
echinata) DC). A828). Wah 860
— ‘gracilis DG@....28¥,.. anRNaVaKs 859
— — var. firjusensis Vass. ............ 859
— karakalensis Vass. ............. 860, 895
— officinalis AUCt. .....ccecccecceeeeeees 858
BS) officinalli§ Masiasicdocsysasrotencntcesy exeh os 860
2>ipersica! @sAiM. wtiean...Abs ere 858
— persica C.A.M. var. gracilis Boiss.
Lee iat cioc IANS. LAAN, che 859
Calendulaceae Cass., trib. .............. 857
+= Bessey, ifamiA..ntniaisianiaik 857
CALENDULEAE Cass., tribus. ...... 857
Cancrinia Kar. and Ki. ....... 293, 876
GANGSTA SC Cbireccenieveeworcsenvesef dh iidaees 314
— angrenica Krasch. ..........:0c0e8 272
— Botschantzevii (Koval.) Tzvel. .309
— brachypappus Wink. ...........0. 313
— chrysocephala Kar. and Kir. .....314
— chrysocephala ssp. tianschanica
Kpaseh jescocivioves PENS. POR 315
— discoidea (Ldb.) Poljak.............. 313
— eriobasis (Rech. f.) Tzvel..........308
— ferganensis (Koval.) Tzvel. ....... 304
— Goloskokovii (Poljak.) Tzvel. ...301
— karatavica Tzvel. ...............cceee 306
— karataviensis (Rgl. and Schmalh.)
Po liaks. crs icopenpenimrmorengast 278
— Krascheninnikovii (Rubtz.) Poljak.
AE SAR Pe). SN, 293
— lasiantha Wink. ........0....:.c000-20+ 278
— lasiocarpa Wink. .........:cccceee 313
— Litwinowii Krasch. ...............00+ 283
— mucronata (Rgl. and Schmalh.)
T2VEh. i cccosezavsssensvats SAR Oe 302
= Nevskil. TzVell) o<::cct. UWE Nee 309

918

871

— pamirica (Hoffm.) Poljak. .........286
— pamiroalaica (Koval.) Tzvel......307
— paradoxa Winkl. ex Poljak.......281
— paradoxos Winkl............:cce 281
— paropamisica (Krasch.) Tzvel. ..310
— Pjataevae (Koval.) Tzvel. .......... 306
— platyrachis (Boiss.) Tzvel.......... 311
— santoana (Krasch., M. Pop. and
Vved:)) Poljak:.ensunernwanm 311
— setacea (Rgl. and Schmalh.) Tzvel.
wansaamannosinapanansa steadied. SoM eke eae 304
— submarginata (Koval.) Tzvel.....305
— subsimilis (Rech. f.) Tzvel........ 302
— tadshikorum (Kudr.) Tzvel........ 299
— tianschanica (Krasch.) Tzvel.....315
— tripinnatifida (Oliv.) Tzvel........ 303
— urgutensis (M. Pop.) Tzvel. ...... 308
Cancriniella Tzvel. ................ 292, 876
— Krascheninnikovii (Rubtz.) Tzvel.
ibe, dates ae Ahan fname 8a <8. h) 293
Carniolici Schischk, sect. ...... 748, 883
Centipeda, Lour:.........-.5...- ase 421
= orbicularis Lour. .nibers-seave8 422
Chamaemelum Mill. ...................00. 66
Chamaemelum Vis. ..........:.:cccsccceeeees 157
=S7Casss(Seet) Mines Aten ees he.. 16}.35
— ambiguum (Ldb.) Boiss. ............ 168

Biebersteinianum C.A. Mey........31
caucasicum auct................ 162, 167

Ra BOISS catcrctcacnanckesansctchsassaseeeateas 161
— — var. pumilum Boiss. ...........+ 162
— colchicum Mand. ............0:000008 174
— cota G.A.2 Mey. {2h Seene karen. 58
=| comavAll, mensasezee es: 63 -
— daghestanicum (Rupr. ex Boiss.)
BOSS, ..Ass-4. Bese th-dattiaaee erat caete 222
— — var. glabrum Boiss. ............+. 222
— decipiens (Fisch. and Mey.) Boiss.
adsQel Laks. Pod ddabas- See Deee eds ay 182
— disciforme (C.A.M.) Vis. ........-+ 181
— — B. quadrilobum Boiss........... 181
— — f. radiata SOD. .........cccceee 181
— elongatum BOISS. .........ceeeee 173
— Grossheimii An. Fed. .............0+. 165
— heterocarpum BOisSS. ...........+6+ Lz
== inodorum (L.)) Vis: sacians.... 175
— — var. B. maritima Vis. ........... 177
— Karjaginii Mand. and Sof......... 167
=| Kofschyt auch. ..0.cr an rnats.. 167
— lamellatum Boiss. ...........0000006 155
— limosum Maxim. ................:00000008+ 177
— maritimum BOISS. .........000-0 ee 177
— melanolepis An. Fed. ..............+ 162
Ee mODile ML) PALL. £3. Stee. 67

872

——), Praecoxe Vas) CK AOS LR 183
— rupestre Somm. and Lev. .......... 173
— sevanense Mand.| ............c:0c00000 182
— silaifolium (Stev.) Trautv. ex
Grossh?’. Xataeax). catenins 220
— Szowitsii Boiss. .........ccccceseceeee 166
— Tchihatchevii auct. ...0....cccc. 174
=§—={Boissw1:....c0net)..nnokene. 174
— tetragonospermum F. Schmidt. 170
— transcaucasicum Manden. ......... 167
— trichophyllum auct. ........c.cc000000 182
Chamomilla DC., gen. .....cccccccccseee. 148
=—t_ SOCL AE ERR SI LR 175
— inodora C. Koch. ...i.0.0...ccccs 175
—=.— Gilid. ..dnasih). aimee s 175
—. praecox'C; Kochynatiiisiwie: 183
Chionogeton Pobed., subgen. ......... 161
Chlorocephala Pobed., sect. ............ 172
Chrysantheminae Hoffm., subtribus.
Se ree...) ea ee ee 129
Chrysanthemoachillea Carmen-Sylvae
Prodan. ...ncnit. Bel. Aes 83
— Borzae Prodan. 2.0.0.0... 84
Chrysanthemum L. ...............00.0..0.. 13]
— Chrysanthemum sect................... 134

— sect. Argyranthemum (Webb. ex Sch.
Bip.) Benth. and Hook. f. 129, 390

Chrysanthemum sect. Argyranthemum
subsect. Brachanthemum (DC.)

Lid Bie in nosed ivi Soil de OAC scree 390
— sect. Brachanthemum (DC.) B.
Fedtsch...2cicaghmedt Mental 390

146
134
325

— — Coleostephus (Cass.) Boiss.
— Euchrysanthemum Clarke. ...
— — Eutanacetum (DC.) Kitam...
— Glebionis (Cass.) DC. ..........134
— — Gymnocline (Cass.) Hoffm. .193
Gymnoclines (DC.) Ling. ................ 330
— — Gymnoclinia Benth and Hook.
FocssizethRA MORE. BEAR RM 193, 330
— — Leucanthemum (Mill.) B.
Fedtsehisistet cere. Rds 140
— — MagarsasDC@i intl. .cmna kes 129
— — Pinardia (Cass.) Boiss. ........131
— — Pyrethrum (Zinn) Gaud. ...... 232
— — Pyrethrum subsect. Dendranthema
(DC.); Kitaimeetitaen oe: 364, 398
— — Pyrethrum subsect.
Leucanthemum (Mill.) Kitam. .. 140
— — Pyrethrum subsect. Richteria

(Kar--and:Kir.) Tongs enn 248
— — Richteria (Kar. and Kir.) Clarke.
daditeacd ie IR RR Rae 248
— — Tanacetum (L.) Hoffm. ........ 325

919

— abrotanifolium (Bge.) Kryl. ...... 250 *
— absinthiifolium Fisch. ex DC. ..370

achilleifolium (M.B.) Kuntze. ..351

— adustum Fisch. ex Trautv.......... 386
— alatavicum (Herd.) B. Fedtsch. 249
— aplinum MB. ..41.20...:csennsatess 161
Se en iia 228
— — f. tatrae Vierh. «0.0... 228
— arassanicum Winkl. ............000000.. 259
— AFCHICUM AUCK. ......cccccceseeecceeeeneeeee 376
Sah ILS... erate pie 386
— — ssp. alaunicum K. Pol.......... 376
— — ssp. Gmelinii (Ldb.) Kitam. 386

— ssp. Maekawanum Kitam. ....386

—='— Ssp. polaris Hult. -.t..ciahes 387
— — var. yezoense Maekawa........ 385
— argenteum (Lam.) Bornm. ......... 361
— — (L.) Willd. «000... 239
— artemisiaefolium Klatt. .............. 256

athanasia (Bess.) B. Fedtsch....406

Aucherianum var. Roopianum

(Bordz)) \Bordzi-3siesit ae 239
— aulieatense B. Fedtsch............... pat,
— balsamita auct. .......0c0cccccceeeeseeeeee 201
SG, Bid sossconcdvceossnecd eee are 196
— — var. radiatum Schmalh......... 196
— — var. viride Bordz. ........:.....0. 197
— — f. glabrescens Nabel............. 197
— balsamitoides Nabel. .................. 198
— Biebersteinianum Adam. .............. 31
— Dipinnatum L, o.....cccceecesetenne 324
— boreale (Fisch. ex DC.) B.

Fedtschiita. 23.5...\...: 38s. 670 eee 327
— budjnurdense Rech. f. .......00..... 326
— canescens (DC.) Rech. f. .......... 336
— capusii (Franch.) B. Fedtsch. ...413
— Carinatum Schousb. ................0... 183
— carneum (M.B.) Steud. .............. 217
— CAUCASICUM PETS. .........0...000cce0eeee 161
— chamomilla Patze. ...........ccccc00 148
— chrysostachys (Rech. f. and K@ie.)

Kitano a). idee Ginn eae 317
— cinerariifolium (Trev.) Vis. ......213
— cinerarium Steph. ex Ldb. ........ 370
— Clusii (Fisch. ex Rcehb.) Kreutz.

1 np btbctiits «fi OM witantcetica en 234
— coccineum Willd. ............cc0000008 218
— coreanum (Lev. and Vant.) Nakai

eX: Moris. 4:c2:... hears. Beas! 380
—Lcoronarium iki neeiaknk 134
SS AUCH) ki cee. ees 135
— coronopifolium Willd. .........006 2107
| COFYMDIETUMM Licks scccccnees tne 232
= corymbosum. Li... enet.. wkeeee 232

— b. Clusii (Fisch. ex Rchb.)

Halacsy csscseics,.c0n lat RAs 234
— crassipes (Stschegl.) B. Fedtsch. ...

Seesascdasecesss ORM OU 338
— darvasicum (Winkl.) B. Fedtsch. ...

siicdes cee. KiMtab ets cdoid ce. Mote ohve 415
— daucifolium Pefs. .........ccceee 221
— dentatum Gilib. ......0...cccccceeee 143
— disciforme C.A.M. «0... 181
— discoideum (Ldb.) B. Fedtsch. .313
— djilgense Franch. ..........cccceeee 260
— eriobasis Rech. f. «0.0... 308
— erubescens Stapf. .......:ccccceecees 374
— Fischerae (Aitch. and Hemsl.)

RRO GN fe seccccccctteeetaeconpusszses a 287
— floccosum Kitam. .......0.0..ce 363
— fruticulosum LAD. ........ccccceeeeeee 396
— — (Ldb.) B. Fedtsch. ................. 406
= TUtesCens Lis cccsscasccascesssadecas etek 130
— Galae M. Pop. 0.0... ccc 242
— glaucum PETS. .....ccccesesereeeeeeees 197
— glomeratum Saposhn. ................- 413
— Gmelinii Ldb. ex Turcz. ............ 376
— gracile (Hook. f. and Thoms.) B.

FEGtseh?')..). 202002. OUR 407
— gracillimum Winkl. ex O. and B.

PEGtS CMs 2:502:..: 8088. Oe 259
— grandiflorum Hook. .........:00000 171
— Griffithii Clarke. «0.00.00... 262
— Grigorjewii (Krassn.) B. Fedtsch.

Se ie Ale 409
— hakusanense Makino. ................. 383
— Herderi (Rgl. and Schmalh.)

Minkw. and Knorr. .................... 414
— hissaricum (Krasch.) Bornm. .... 241
— Hookeri Kitam. ................0::00c006 407
— hwangschanense Ling. ............... 376
— INGICUM ANCE. ......ccceccceseceecesteeeeees 373
Sa NAICUMN Lay si0i8e. I BRO 371
— INOdOrum L.,. .....cccceeeeeeeees SAT 7
— integrifolium Richards. .............. 388
— ircutianum TUICZ. .......ceceeeeeeeee 143
— isabellinum (C. Koch.) Rech. f.

Rh ree a nr 360, 361
— Kelleri Kryl. and Plotn. ............. 212
— Kotschyi (Boiss.) Nabel............. 229
— Kozo-Poljanskii Golitz. .............. 377
— laciniatum Gilib. ........0..cceee 136
— Ledebourianum Ling. .........0 313
— leontopodium (Winkl.) B. Fedtsch.

Licata CORTETeTagvease MT 260
— leucanthemum Li. ..........:ccccccesees 143

— ssp. ircutianum (DC.) Sukacz.
sscuslasedscssctesdedaad COs ee 143

873

— — var. auriculatum Peterm. ..... 144
— leucanthemum var. discoideum
KOCHI. . LLL. Meveraneee teeters 144
— — var. hispidum Boenn. ........... 144
— — var. ircutianum (DC.) Kryl. 143
— — var. subalpina Schur. ........... 145
— leucophyllum (Rgl.) B. Fedtsch. ....
ihotogp Filynts aeiebeaetead Rds reaamsas) Alans. ¥ 414
— lineare Matsum. ...............:.000006+ 139
— — var. manshuricum Kom. ....... 140
— littorale Maekawa. ...................... 384
S=HUEIAUM, Nakai .....-s.ceccecves.stettee ee 383
— macrophyllum Waldst. and Kit. 194
— majus (Desf.) Aschers. ............... 201
— Marschallii Ascherts. .................. 218
— Maximoviczianum Ling. ............. 374
— — var. dissectum Ling. ............. S79
— Maximowiczii Kom. .................... 379
— maximum Ramond.................00 145
— millefoliatum L............. 83, 84, 348
— — var. achilleifolium (M.B.)
SCHMAUNS 2h. TAT Na .. S88... 2 351
— — var. Kittaryanum (C.A.M.)
KRraschis.4.. 2098... 0. see ey 345
— var. tanacetoides (DC.) Kryl. ...
ciecavias etl RATE RTs ROE ee SEY
mongolicum LIANG. «1... 378
monspeliense GeOTgl. .......20.000+ 183
morifolium Remat. ............0 373
mucronatum (Rgl. and Schmalh.)
Minkw. and Knott. ................04 302
multifidum DeshA0...2..... 28022 197
TRY CONUS. VaNi 05. AHN. «2d sa. TOs 147
myriophyllum (C.A.M.) B. Fedtsch.
Hiesecoweswcprasacwoes aii colts SOAS 332, 333
myriophyllum (Willd.) Nabel. ...331
NAKLONGENSE AUCC. .......ecceceeeeereeee 374
=S"Nakais 2068). one 375
— var. dissectum (Ling.) Hand.
IMA LZ io seccas cnn case ad ncatnvesnste OER 379
NGRUNE AUCH. D....cVHTA ch 410
neofruticulosum Ling. ..............+ 406
Newesskyanum (Winkl.) B.
MeOGtSGhts 22. .2005..063. 20. b, ARR 328
— orientale Adam. .............c:cccccseeeeee 32
— oxylepis BOrdZ. .........:cscseesereeeee 238
— Pallasianum (Fisch. ex Bess.)
Komen... See eB oe 402
— palustre (Willd.) Pers. ............... 215
— pamiricum Hoff. ..........0.c0 286
— parthenifolium (Willd.) pers. ....203
— parthenium (L.) Pers. ...........00 204

— — var. parthenifolium (Willd.)

Schmath....8.60i.88.1. eth 203

920

874

— polycladum Rech., f. ..........0.606 288
— porphyrostephanum Rech., f. ...262
— poteriifolium (Ldb.) Bornm. .....236

— praealtum VED. «20... 207
— pseudoachillea (Winkl.) B. Fedtsch.
. scaaphaniaaastOAeh o> « SGubts aathaecs cary 328
— ptarmicaefolium Mus.-Puschkin. ....
aa lanai uncannits satiate 109
— pulchrum (Ldb.) Winkl.............. 253
— punctatum (Desr.) Bordz. .......... 215
— pyrethroides (Kar. and Kir.) B.
FedtSchh, «..cscisineacaeottt Rie Se 256
— reflexum UEKI. .......cccccsceseeeeeeee 375
— richteria Benth. ...........:...:cceee 256
— richterioides auct. ...........:0000 254
= — Winklisx:.). sna ea 255
— richterioidies var. virescens Winkl.
ssnacdaceeadacnceanasnateebonh «Etec eee eeeERTS Ue aaa 254
— Roopianum Bord2z. .........00c000 239
== /roseum “AdaMm..... ..4-.0dubseeadins veces 217
— — ACO) sia. Es 219
— — var. carneum (M.B.) Schmalh.
avsSeolViesthoasdsh. BUA EEO NING Ms oe REO avg 217
— rotundifolium W. and K............ 142
— Roxburghil Cass. ............ceeee 135
— — var. breviradiatum DC. ........ 136
— — var. discoideum DC.............. 136
— rupestre (M. Pop. ex Nevski)
RREGR,.. Bo ancscuan.akieee stress betes testes 287
— santoanum Krasch., M. Pop. and
NIVEG. sc cnsesccssas oS etIetoot- sided +e MAES 312

— santolina (Winkl.) B. Fedtsch. .343
— saxatile (Kar. and Kir.) B. Fedtsch.

DRS, a Bho kootiescdencc ele UT ee 344
— scapigerum C.A.M. ......:cceeeees 388
— Scharnhorstii (Rgl. and Schmalh.)
B...Redtscit.......0cu8. secteur 409
— schugnanicum (Winkl.) B. Fedtsch.
DeAE 1 eae T) ae Re Nie” x 416
SE ESE REMUTIL Lo... casacincenseasacssscatepaces ereteey 136
— Semenovii (Herd.) B. Fedtsch...244
— sericeum Adam. ........ccccceceeees 237
— serotinum oie) ewineisain. 139
— setaceum (Rgl. and Schmalh.) B.
FReGtseh .....socnaces-=dttSetn ay. Svleeeense 304
— sibiricum Fisch. ex Turcz. ........ 376
— — Turcz. ex. DC. .....eeeee 376

— — var. acutilobum (DC.) Kom.376
— — var. alpinum Nakai. .... 380, 879
— — var. latilobum (Maxim.) Kom.

aceaceR Baiveudes did t evel: elena eee 375
== |SIMENSE, AUCH) 2.4.2. d bs cteesnndabaitonacs 374
— — Sabiliesiissccmeiin tt. Riders Sais
= | SENUALUIM, LGD... .cccacaneceeso.theebeeetnese 370

— Stoliezka@exauctai2hs. issih hike 262
— Stoliczae Clarke. ............cccsce 263
— suaveolens ASChETS. ..........01.00006 150
— subcorymbosum Schut...............- 234
— subsimile Rech. f. i..i..cccccsccsescsees 302
— Szowitsii (C. Koch.) Bordz. ...... 329
— tabrisianum (Boiss.) Bornm. ....335
— tadshikorum KudQr. ...........0....:000 299
— tamrutense SOSD. .........ccccccseceeesees 334
— tanacetoides (DC.) B. Fedtsch. 337
— FANACELUM VIS. .....cccceesccceseseeereees 326
— tenuifolium auct. ......cccccececeee 410

— tenuissimum (Trautv.) Winkl. ...355
— tibeticum (Hook. f. and Thoms.)

Hoffim; 22 seen Se ot 410
— transcaucasicum (Sosn.) Bornm. ...
annctea Larne eee ei ERE cin 353
— transiliense (Herd.) Minkw. and
Kinorr, ols coheed. epee 258
— trichophyllum (Rgl. and Schmalh.)
O. Kize. ine a. .c ee eee 363
— trinioides Hand.-Mazz. .............. 417
= ‘Turreanum-Ni8 accidents 213

— uliginosum (W. and K.) Pers....139
— umbelliferum (Boiss.) Hoffm. ...363

— uralense Krasch. ..........::ceeeeee 346
— variifolium Chang. ........:cceeeee 401
— — var. ramosum Chang. .....-....- 401
— vulgare (Lam.) Parsa............000 143
— (LL) Bemhij ngs wks 326
— — var. boreale (Fisch. ex DC.)
Makino. ..::.ssi¢e34-sancknaween B27
— Walteri Winkl.............. cee 359
— Weyrichii (Maxim.) Miyabe and
Miyakeiih....2cc0ts oneal 383
— — var. littorale (Maekawa) Kudo.
Leinadage eR A er REARS weet 384
— yezoense Maekawa. .........:00000 386
— Zawadskii Herb. ...........:cesceeseeee 376
— — ssp. acutilobum (DC.) Kitag.....
eee de. bah eusanineanls 379
— — ssp. latilobum Kitag. ............ 375
— — var. alpinum (Nakai) Kitam.
CONS RO he Ok YE 380, 879
— — var. latilobum (Maxim.) Kitam.
RAR or ied Gros bras 374, 375
Cineraria sect. Holoseris Rchb. ....752
— sect. Tephroseris Rchb. ............. 753
= alpina All Aiken 768
—+ sHREIZ8. 4 tds canines 753
SSlaligicdg™ CiAsMe ticciacnuacnns 843
— ambracea Hornem. .............000+ 715
— atropurpured LAD. «0.0... 771

— aurantiaca Hoppe. ..........0 768

921

—VAUTCG Lincs eee ee 736
=~ lauriculatayLdbsmstiereiiien 733
— I GAINDESITIS REUZ. PAG AA. eats 753
— — var. longifolia Trautv........... 759
—'—, &.,.dentatasi DG. .t:.2.0ena kt 759
——".capitata, Wahlbi ities), iene 768
== (CON DICH: DEUS) sts. <stesssschecs «tsseseens'oe mil
= caucasica MBas2.duiksxn wads 855
S=aerispa-lacge sieve. sccewshs 9. ged accde 765
— — 7. Schkurii Lbs .h.:..adcsce 753
epluschery Id ys... 0s 5 eiesaeaees 802
— flammead TurcZ. ....ccccccccccecccseeeeees 767
— frigida Richards. .......,.....s0.. 771
— — Q. atropurpurea Herd. ......... 775
— — @. eriocephala Herd. ............ Vas

— — Q. genuina a. arctica Herd. 771
— — genuina b. taimyrensis Herd.....

— frigida &. genuina c. uralensis

| Lote Seer pepe eer eee me Ti
— — Richards. B. robusta Herd. ..775
— — robusta a. atropurpurea Herd.

eicsssos ete Devavstut tara: tsiThe oxtedbmebeneslie 777
—— ULV AMSION juett... Aes 0 3. caediaeeics 769
SAGEM AIG CAM, ..snonanpsocsensisinebéoase 853
ee Ola ella TMUNGZ Et. «haat. derstoestes ves 762
wR LADIala ded). cieceadihs sivsscestccdesvacess 733
—— PIANC Ay Lo. gievciciostaseeatita syesitn-e 847
= ==" s/0) 000) Aiea eee 808
— heterophylla Fisch. ..................... 750
ALO EM OE ISCH S21 use sesesonsoaicydevescccacass 767
Sag LOM GLLOUIG, AMES. 2s5s<0s00ctsscatysanesacuen 759
ee TPE LAD rans dglissnnad sinh esidancsibis a 750
— macrophylla Lab. ............0.000000 837
MLTR TTT (Ye) EE oe eee 720
—— NANA NUT GZ ohetarcogtyontcseeiaeentees 802
—— MON POUCA mL Zs: <hes. tact scts<c cess 849
= LUST ES it avrses ascascs sofocne nae eee ts2
=— PAPDOSG. IWCUD: <.cccccccccncee-sseesgesteas 760
—— DOT VIULONA. NUIB, cote cacsaottccessensees 744
— racemosa. M.B., j.....:.-.cceseese00sc000s 731
——— FENUOUG. CAM -ccrsscscneyesstceizessetee 853
—- rivularis Waldst. and Kit. ......... 765
a= FODUSIO LGD. es-coacctse tite cates 820
— sibirica auct.....810, 812, 818, 889
some Rees NLA sey su Sev dest tcoanctate covers corettnte rrr 807
cen OPI, «case eece ehog eeenreens 808
sm SDE CIOS NCHTAG.*...scscs.scteeteetteerse 802
— Subdentata Bge. ..0........cccscecseacees 786
— taraxacifolia’ M.-B. ...........:.<s0++- TPS)
— thyrsoidea LAD. ..0.......ccccceeeeeeee 827
—— URCTHOSA AGUMD. .ccsccetfeantecstsecsitecs T52
Cinerariifolia (Heyw.) Tzvel., sect. 213
Coleostephus Cass. ............:::cccc0e 146

875

—- myconisy(L.)) Cassia .ieee 2 147
Corymbosae (Fr.) Hand.-Mazz., sect.
wbnes stb eMTss cde N tee ed 798
FA ENA essa cenpretscnlerepecivinlincitives 9, 39
Cota (J. Gay ex Guss.) Rupr., sect. 39
—laltissima. J... Gigs 1, Beaks Ries. 58
—laustriaca..5: Gays. 6.2). Sees. 59
— coelopoda BOSS. .:...........ccsseeies 59
—OFMENIOIdES. BOISS. w....0sc0eseusets0sss0s0 60
RAN oo cvaievuineneeniinnecacehas OTA 418
SIR UP as SCC. gecrarecenvind ets invssthhlovdes 63
WUT EAN SE Sich MO EN 151
=— coronopuoliay Ls. \:.....62A0th...i1n 421
— foetida S.C. Gmel. ..0.......cccceeeees 63
— matricarioides Bong. .................. 150
Crassocephalum valerianofolium Less.
ewecat eH eee d. STA EOL Ss os EERE 683
Crociserides /DC.,: sectiict..aigigit 726
Crossostephium auct. ..........ccccsee 315
Crossostephium Less. ..................... 316
— sinense (L.) Makino. ...........0...... 316
— turkestanicum Rgl. and Schmalh.
sabes Os ceetireeteevadeeve EIA S Tnedt RR 316

Cyathocephalum Nakai. 788, 850, 894
Cyathocephalum (Nakai) Kitam., sect.

avneEMITtenntn Renee eee Roe RAN 836
— Schmidtii Nakai. ............00.cc000000 851
Cynareae Less., subtr. VI Arctotideae
eS 94 Meanie RRMA ee 861
— subtr. Calendulaceae DC. ......... 857
Cynarocephalae Less. div. II
Gundeliaceae DG@i.0.U8...keh 861
Cynoclinés;:-DC.,iSe6R 22. 168
Decaneurum Sch. Bip. ..........c00008 137
— serotinum (L.) Sch. Bip. ”........... 139
Dendranthema (DC.) Des Mooul. ........
ee ces aN e baaaiy ap ee 364, 879
Dendranthema Sect: .:.....-02--nte 370
——dreticum (L:)) Tzvel.2..0.2 en. 386
— erubescens (Stapf) Tzvel. .......... 374
— Hultenii (A. and D. Love) Tzvel.
cree ite oe ea pier be ee a ca id 387
— indicum (L.) Des Mooul.............. 371
— integrifolium (Richards.) Tzvel. 388
S="KUMMIENSE’ TZVEl.. ccetesst.c--ceoteee: 385
— littorale (Maekawa) Tzvel. ........ 384
— Maximowiczii (Kom.) Tzvel. ....379
— mongolicum (Ling) Tzvel. ......... 378
— morifolium (Ramat.) Tzvel........373
— naktongense (Nakai) Tzvel........ 375
— sichotense Tzvel. ............... 380, 879
— sinense (Sabine) Des Mooul. ...... 373

— sinuatum (Ldb.) Tzvel................ 370

922

876

— Weyrichii (Maxim.) Tzvel......... 383
— Zawadskii (Herb.) Tzvel. ........... 376
Dendranthemopsis Tzvel., sect. ............
Spe See 394, 880
DiotisabDests &. £:23itek: te eee cee 127
— candidissima Desf. ............::c000006 128
— maritima (L.) SM. .......ecceeeeeeees 128
Dolichorrhiza Pojark., subgen. .............
casaaxkvananeanndeaasenat atest sta yeros 853, 895
IDET RI (aT 11 Wel CR ee a 669
Ee Cavalli SOC bans cuccacasncs Geant. uc SEES 672
— sect. Aronicum Hoffm., sect. ....672
— sect. Doronicastrum Cavill. ...... 676
— sect. Doronicum Tausch. ........... 671
— sect. Eudoronicum DC............... 676
— subsect. Cardyophylla Cavill. ...674
— subsect. Grandiflora Cavill. ..... 674
=—— altaicum Pallieus, 0) shee 672
——V arnicd, Desh... tee ee 663
— atlanticum Chab. ..............c..00. 679
— austriacum M.B. ..........ccccccc0c000 678
— austriacum Jacq. ...........seseeeeee 676
=—— Balansae: Cavill. c.cc.scessscccseserencenns 678
— bargusinense Serg. ..........0c:0000 673
— cacaliaefolium Boiss. and Heldr. ...
Sg a ec an eck fee 678
— carpaticum (Griseb and Schenk)
INyimiis. 3. . Lbu...2eetbiads -qasdetavee cases 675
— carpetanum Boiss. .............eee 677
— CAUCASICUM M.B. ...cccccceeceesseeeseee 677
— Clusi [(All): Tauschiesdiasate 674
— — var. villosum Tausch. ........... 674
— cordatum C. Koch. .................... 677
— dolichotrichum Cavill. ............... 678
===» Falconeri Clarke. ..c...s0-ssssesnoacncsone 680
— Haussknechtii Cavill. ................. 678
— hirsutum NAM e. 9c asks cass toss odes 674
— hungaricum Rchb. f. ..........008 680
— longifolium Griseb. .....................680
— longifolium Rchb. ...........cece 676
— macrolepis Freyn and Sint. ....... 678
— macrophyllum Fisch. .............0 678
— maximum Boiss. and Huet. ....... 678
—— NONIANUM) AM. 5 snow Seo sasscacsiteass 663
— oblongifolium DC. ..................... 681
— oblongifolium DC. var. leiocarpum
SM AUOV A bis eet esi. els hee coi 681
— oppositifolium Lam. ..........:.:. 663
——s OFLEN tale JOPLIN) thes «sa tnaescstecsseacsnse 677
— pardalianches auct. «0... 677
—« Dardaliancnes Uh sci. -o0a) sauces sashes 679
— plantagineum L....................-0s0s0 680
— reticulatum Boiss. ............:.ccc000 679

== Roylet DG. oo meet sta his -scaasteespsaets 679

— Schischkinii Serg. .............-sssseses 673
— scorpioides Lam. ..........::c::0:e008 674
==) Thirkei .Schultz. 22. 8..2%4:72-nae 679
— turkestanicum Cavill. ......0...000.... 681
— vaginatum C. Koch. ............0.000 678
Dracunculus Bess. subgen............... 525
Elichrysea Ain, ordo. 3). 2a 1
Endocellion Turcz. ex Herd. .......... 653
Endocellion (Turcz. ex Herd.)
Kuptlan. vceaegivine ith. Pee 653
Endocellion boreale Turcz. ............ 653
Erechtites Raf... e248 Rae 682
— hieraciifolia (L.) Raf. ..........0..... 683
— valerianifolia (Wolf) DC. .......... 683
Eriopappus Dumott., sect. ............... 752
Erucifolii Rouy, subsect. ................. 710
Erythrochaete Sieb. and Zucc..............
SS RE oA: Se et ed 788, 798
Euanthemis DC., sect. ........ 16, 39, 63
— ser. Annuae BOISS. ...........600:cc00000 35
— ser. Perennes BOisSS. ..........c000c0000+ 16
Eugastrosulum Sch. Bip., sect. ...... 183
Eugastrosulum (Sch. Bip.) Pobed.,
SECO nce Manco itis. Me eee 183

Eumatricaria Sch. Bip., sect...........148
Eusenecioneae Benth., subtr. .........640

Extremiorientales Schiskh., sect. .........
ides tvs agradecer tee 746, 883
Farfara-Gilibscn cairn 641
Filavotmaritima Ws. Wiican 128
Filifoliom Kitamit... cence 416
— sibiricum (L.) Kitam. ................. 417
Filipendilinae*DC. sectt ite 990
Gastrosulus praecox Sch. Bip. ...... 183
Glebionis” Cass." .<.8- state 131, 134
— Roxburghti Cass. .........:.csceesee 135
Glossanthis Poljak. .......:::000 PART
— aulieatensis (B. Fedtsch.) Poljak.
Piri eies 2 See ba eae Seas i: 277
— aurea (Krasch. and Vved.) Poljak.
Ree A tales pads ate Chere 2 281
— radiata (Karsch. and Vved.) Poljak.
esvegaandtae sath setaeas Cases Meena e 280
Glossophyllum Pojark., sect. 852, 894
Grammarthron Cass. .......:.1scecees 669
min SUP QONUS. ccactnennctpareeMlceessveseasiatat 671
Gundelia"lie gee cone 862
BLADE ee ccccsevesrepan ter scnet ce eee 862
— purpurascens Bornm. ..........-..0++ 862
==, POUrMeLOrH! Vi i.e eee: 862

923

— — var. asperrima Trautv. ......... 862
— — var. tenuisecta Boiss. ........... 862
GUNDELIEAE Boiss., trib. ............ 861
Gundelinae Benth., subttr................. 861
Gymnocline Cass. ........+: 184, 193, 330
— sect. Anactis C. Koch. .............. 325
— sect. Chrysactis C. Koch........... 330
— sect. Leucactis C. Koch............. 193
— achilleifolia (M.B.) C. Koch. ...351
— argyrophylla C. Koch. .............. 3334 |
— armena (C. Koch.) C. Koch. ....352
— chiliophylla (F. and M.) C. Koch.
Se iets, RR RRA RRR 352
— leucocephala Cass. ..........seee 194
— macrophylla (W. and K.) Bluff
and) Fingerhs).:. A220... 2eeas 194
— myriophylla (C.A.M.) C. Koch. 332
— Szovitsii C. Koch. ........c.ceeee 329
Gymneclines (Cass.) DC., sect....... 193
Handelia Heimerl.............0.......08 124
— trichophylla (Schrenk) Heimerl. 125
Haplophylla Tzvel., sect. ....... 388, 880
PREESICOMO RAL | cccnassecsertssstcsn tees 683, 685
— auriculata Pojark. 00.0... 694
i FISTAEA POJATK, 2....-2..cecessrscsesenesoon 687
— kamtschatica Pojark. ..............04 696
— Komaroviana Pojark. .............+ 691
— praetermissa Pojatk. .........00 693
— robusta Pojark. ...........ccsseeee 690
Hemipappus C. Koch. .................. 359
— argenteus (Lam.) Tzvel. ............. 361
mm CAMS Ce NOCH. cin -c.ccrersocsccsescesccere 360
— isabellinus C. Koch...............004 361
Herbichia carpathica Steud. .......... 750
Hippolytia Poljak. .......00.... eee 411
— Aphanassievii (Krasch.) Poljak. 408
— darvasica (Winkl.) Poljak. ......... 415
— Herderi (Rgl. and Schmalh.)
PON] ake seccsrettvsersssticescessscssecsesosseeses 414
— leucophylla (Rgl.) Poljak. ......... 414
— megacephala (Rupr.). Poljak. ....413
— Scharnhorstii (Rgl. and Schmalh.)
| eo | fl 3 eer he ra ee 409
— schugnanica (Winkl.) Poljak.....416
— senecionis (Jacq. ex Bess.) Poljak.
sande Potansatattses eee ttecicnecaseseeretehontes 414
— tomentosa (DC.) Tzvel. .............. 416
Homogyne Cass. ...............:cscseseenees 654
— alpina (L.) Cass... 655
Hopped RhbD. ..........ecceeeereeee 788, 801
=="SIDIFICAMRCMDY ts s.ctsectcesetccseces este 807
——"SPECIOSAURCND i isionittescsatscschectsastseesee

877

IndanivDE@s- sect: .ins.. Mee 720
Usmediat Cassini sesteccsscorseetats 131, 133
Ismelia (Cass.) DC., sect............000 133
Ismelia versicolor Cass. ..........00008 133
Jacobaea" DCS SECE «.. iii Ane cescestdeeess 710
Jacobaea carniolica Schrank. ........ 748
== PRCHStt™ Cu Mie secs cseccitoceseseotceces: 745
— erucifolia Gaertn., Mey. and
SHEN: Se eo eh AEs Boeerecerace streets 710
— Othonnae C.A.M. ..........cccccceeeeeee 741
— sarracenica Moench. .................- 746
STC Cd GT D is eos taescoccescssccacucsstee 715
— squalida C.A.M. uu... 783
— tomentosa Moench. .................006 720
——— VES COSED. COULD ceiecte ceuectesscaxteorrevzces 7179
==" VULDGFIS (ClBUS., te..cctec.c-veccetceonenses 780
Br 6 Vos 9 11 Be oer ee 715
Junceum Poljak., S€Ct. «0... 626

Karelinoidei O. and B. Fedtsch., sect.

Pe ee eT ee ee 751, 883
Kaschgaria Poljak. ..............:00008 423
— brachanthemoides (Winkl.) Poljak.

— Komarovii (Krasch.) Poljak. ..... 424

Leianthus (Novopokr. and Sidor.)

TTZV GI SE CT a accsdesstdsdexesstacegstoeeee 280
Leiospermum Sch. Bip., sect.......... 181
Lepidolopha Wink. ..................00+ 288
— Fedtschenkoana Knorr. ............... 291
== ‘filifolia. Pavilii2 200. . steed ctteout 289
— karatavica Pavl. ........cccccesseees 291
— Komarowii Winkl.................:0+ 289
— — var. mogoltavica Krasch. .....290
— mogoltavica Krasch. ..............++ 290
— nuratavica Krasch. ..........::-00+ 290
Lepidolopsis Poljak. .........::scee000 293
Lepidolopsis Poljak. ..............:00000 315
— Goloskokovii Poljak. ........:.:0+ 301
— Kotschyi (Boiss.) Poljak. ........... 333
— Krascheninnikovii (Nevski) Poljak

ORM ee ec fee ae 309
— mucronata (Rgl. and Schmalh.)

Pol pike) scaidisediiteind nal eR 302

— paropamisica (Krasch.) Poljak. 310
— pseudoachillea (Winkl.) Poljak. 329
— scopulorum (Krasch.) Poljak. ...342
— setacea (Rgl. and Schmalh.) Poljak.

pee trie An ea eS. 8 04
— Tadschikorum (Kudr.) Poljak. ..300

924

878

— turkestanica (Rgl. and Schmalh.)

PORK YS: costes tocsestcnnnseeee eee 316
— umbellifera (Boiss.) Poljak. ...... 363
Leptanthemum Tzvel., sect. ..210, 871
Leucanthemella Tzvel...........00.000.... 137
—serotina (L:) T2Veliiscstussctcees 139
-— linearis (Matzum.) Tzvel. .......... 139
Leucanthemopsis (Giroux) Tzvel.,
SECU aa cvdncts gestae eee eon tas 227
Leucanthemum Mill. ...........0.......... 140
— alaunicum (K.-Pol.) Worosch. ..377
— algidum Fisch. ex Trautv. ......... 388
— alpinum (L.) Lam. oe 228
— — var. cuneifolium Mutt. .........229
== /ATCH CUT! AUCENS. .5.i0hisecatsasdepsereaese 387
sae = (LDCR A Airs ascot) 386
— balsamita (L.) Over. «0.0.2... 197
sy Stank. eee ibs hasguecaseeaseaesese 201
— corymbosum (L.) Gren. and Godr.
PIRES CRIDER once tanedee taren earieots 232
— ZeOrgicum SOSD. ........scseseeeerees 143
aa (Ge LNT AD ores. ccdesstecsteccdieee ieee 386
— Hultenii A. and D. Live. .......... 387
— integrifolium (Richards.) DC. ...388
—Vircutianum DEA eee 143
— leucanthemum (L.) Rydb. .......... 143
— maximum (Ramond.) DC. ......... 145
— parthenium (L.) Gren. and Godr.
BETIS aco acacia pete A aa eee COPIERS 204
— Raciborskii M. Pop. and Chrsh.
pa cht aliei DR aed CE 145
— rotundifolium (W. and K.) DC. 142
— segetum (L.) Stank. 0... eee 136
— serotinum (L.) Stank. ..........0.... 139
== P sibinicum/auct. 4. iw. eee 377
=o DG ahs eee 386
— — ssp. alauncum (K.-Pol.) Golitz.
£1 eS a tet aes nite 377
— — var. acutilobum DC. ............. 377
— — var. alpina F. Schmidt......... 378
— — var. latilobum Maxim. ......... 375
— — var. peleiolepis Trautv. ........ 378
— sinuatum (Ldb.) DC. ........... 370
— subalpinum (Simonk.) Tzvel.....145
vulgare) Lan iiscatarantideaicts 143
— — var. ircutianum (DC.) Kryl. 143
— Weyrichii Maxim. .........:ccccee 383
Leucoglossa DC., sect. .......... 168, 172
Ligularia O. Hoffm. ........c.cceeeees 797
Ligularia®‘Casswiik:.eniidcnene 788
Ligularia subgen. ...............004 797, 886
Ligulatia sectasc.ts cadena 801, 886
— sect. Cyathocephalum (Nakai)
Kita Soft 2, mechs, tees 797, 894

— sect. Eulgularia Kitam. ............. 7197
— — series Corymbosae Kitam. ...798
— — series Racemosae Kitam. .....801
— sect. Senecillis (Gaertn.) Kitam. ....
RARER, et ee ea 797
— II. Racemosae Hand.-Mazz. ...... 801
— abakanica Pojark. .............. 810, 887
— afghanica Pojark. 0.0.0... 837
— alpigena Pojark. ..........cccceeees 841
—=(altaica: auctias.2kk. Waianae 842
== jaltaidacDC rtd awe 843
— altissima Pojark. «0.0.0... cee 835
— angusta (Nakai) Kitam..... 844, 850
— arctica Pojark................ 817, 891
— Aucheri D@ks...0e indeed 725
— auriculata TUrcdZ. occ 694
— bucovinensis Nakai. ...............0.. 818
== /BungeiM:..Popewiast..o).ntskena 834
— cacaliiformis Rchb. ..............0 806
— calthaefolia auct. .......:ccccceeeeee 798
— calthifolia (Maxim.) Maxim......800
— carpatica (Schott. Nym., Kotschy
POjabk tists wares stooge 8
— caucasica (M.B.) G. Doon. ............. 8
— clivorum Maxim. .........cccsceeeeseeeeees 7
— Correvoniana (Alb.) Pojark. ........ 85
— deltoidea Nakai. ........... cee 827
— Fauriei (Fr.) Koidz. ................0 849
— Fischeri auct. ..........c0000c00 804, 805
— Fischeri (Ldb.) Turcz. ................ 802
— — f. diabolica Kitam ................ 805
<== PO LOUG DIC. taecapcsaascosvsiouesnacssnveanes 853
— glauca (L.) O. Hoffm. ................ 847
— — ssp. carpatica Hayek............ 848
— — var. altaica Kryl......csce 843
— — var. typica Kyl. ........ceeee 847
== NAStAA) LESS. .cstsbscsitsrnstasarenteanbuarn 687
— heterophylla Rup. «20.2... 838
— — a. alpina Rupr............. 840, 841
— — B. subramosa Rup...
casgee ied a neers eat 839, 840, 841
— — y. thyrsoidea Rupr. ..... 839, 840
— Hodgsonii Hook. ............:c:ecse 798
— Hodgsonii var. calthaefolia Koidz.
acssseaghiednecd¥osdbeh aes tactasvunatemacann seas 798
— — var. sachalinensis Nakai......800
— Jacquemontiana (Decne.) Kitam.
satiwnnpeesestv ads angis Satna MaEa es Soesanne sete coer 844
— jaluensis Kom. .........::csscseeeeeeee 826
— karataviensis (Lipsch.) Pojark. .824
— Kareliniana Stschegl. ................. 821
— Knorringiana Pojark. .................. 832
— longipes Pojark................. 816, 893

== Lydiae MimGen. cccccscceccrcusecermeenes 806

925

macrophylla auct. .......eeee 838
macrophylla (Ldb.) DC. ............. 837
+ var. minor Winkl. «00.0004... 839
mongolica (Turcz.) DC. ............. 849
narynensis Winkl. .........:++0e0 822
narynensis (Winkl.) O. and B.
FedtsGlis scossesecacseasar. cettits ntact ronteess 822
Povlovii (Lipsch.) Cretz. ........... 852
PETSICA AUCL. .....scceseseerccseeceereeeceees 838
persica BOISS. .........-..+:+00 837, 840
Potanini Pojark. «0.0... 841
pulchra Nakai. .........sssseseeees 826
Putjatae Hand.-Mazz. ..............+. 849
renifolia (C.A.M.) DC............. 853
TOPUSIG AUGCE, ssccarsacecc-teiotendterecbioeees 822
robusta ((Ldb.) DGC.-.220i.c2 820
— var. Kareliniana O. and B.
edison: secescssecce coca 821, 822
— var. typica O. and B. Fedtsch.
eT OE, ELSE 822
— f. turkestanica Krasch.......... 822
sachalinensis Nakai. .............060 805
Schischkinii Rubtz...................... 825
Schmidtii (Maxim.) Makino. .... 850
sebennensis ROWY. ...:.c.ceeee 806
SIDIT ICQ AUICK, 4... gerceases ecctacerseteet eases Zaseae
803, 808, 812, 816, 817, 818, 889
Sipirica, (L..)) Cass. wags.s.-sssassteneert 807
sibirica speciosa DC. ........-01+ 802
— ssp. Fischeri Kitam. ............. 804
— var. alpestres minor turcz. ..816
— var. araneosa Nakai. ............ 803
— var. bohemica Rupt. ..........-.- 808
-— var. fallax Nawal. .:.i0t..censets 803
— var. gigantea Serg. .......+.04+ 807
— var. glaberrima Ser. ........... 807
— var. gracilis Serg. ....... 816, 817
—— var. hirsuta Serg. ...............-.- 810
— var. minor Nym........... 817, 891
—- var. SPECIOSA AUCL. «0... eee 805
sichotensis Pojark. ............ 811, 888
songarica (Fisch.) Ling. ............. 831
SPECIOSA AUCC. .....seeeesereseeees 803, 805
— Fisch. and Mey. ............---.-« 802
— var. araneosa Kom. ............... 805
splendens (Lévl. and Vaniot)

INAK AL. ccsscccspseiscten-n----<s enemies 803
subsagittata Pojark. ........... 812, 889
talassica Pojark. ............s..s0-e+0 840
tangutorum Pojark. .......--.s.0:es 841
Taquetii (Lévl. and Vaniot) Nakai.
Bra pa pete sewn os Joneses hn 849
Thomsonii (Clarke) Pojark. ....... 834
thyrsoidea (Ldb.) DC. ...........++ 827

879

— — var. alpina BE. ..........:0 834
— trichocephala Pojark. ........ 800, 886
= Stucrainica’ Minder..1.2. tise. 818
— virgaurea (Maxim.) Mattf. ......... 852
Ligulata Krasch., sect. ........... EAR 277
NATE TRESS soe cotcccctscetes strseceecesonnas 9, 63
Maruta (Cass.) Boiss., S€Ct. ............ 63
Bi ailiiisin | Crea eerrecree sean cae 63
— — fi. lithuanica DC. ................0. 64
— fOetida CASS. ........sceerceceececesseerseese 63
— microcephala Schrenk.................. 65
— vulgaris Bluff and Fing............... 63
ps Eyre CoE) gr) Bie ate ha etn Pet ai ere 147
— aplina (L.) DEST. .........eseeeeeeeees 228
— ambigua auct. ................-.- 170, 172
— KI. once ceceserceresecececeeneeneone 168
ae TP CIIC sc) DCSE yccttace- cones sucestesses 386
— _ AF ZENE L..........0.-rcssesncerseonseeeerere 239
= aurea (Lin) ES OlSS: cremcrceresaateeswunrss 151
= hal saniiia: (1s.)) WeStycere-seccenttsnee 197
— bipinnata (L.) Dest. ...........00 324
a CHAMONITUG. Varin ctstavigereneesess 148, 175
— coreana Lev. and Vant.............. 380
== ‘coronaria. (10) WEStn ccrcsacttte esses 134
— corymbifera DC. .........sceeseees 181
— corymbosa (L.) Desf. ..........+++ 232
— deflexa GiliD. ...............c-c--cneseeeees 148
— ‘disciformis DC. .......ccieccnrceeersenes 181
se ISCOLAE AT DC iis. ceeect nerecaes Oh ennereses 150
— grandiflora AUct. .......-sreeceresereee 172
pe SD hy | pooner Wii
— heterocarpa C. Koch. .............++ 173
Se INGICE (s.) DESks cecetysrcecee scares St
we TAOMOTA AUC Tne ipeetecre retre otee 177
A A) Onetireirbe stir ris Pvetrekn or 175
=> = maritima’ LNG. '..1-.20s:200 177
Sek VAT: VEETICE ISLS teenie coett earn nce 171
— — B. maritima Wahlenb. .......... 177
— — ¥. praecox LAD. ...... eee 183
=="—"€. breviradtata Lab! --...i.0.:. 178
— — A. monocephalum Sch. Bip. 171
— — f. phaeocephala Rupt. ......... 171
— lamellata Be. .......ccseeeeevees 155
— — var. discoidea Rgl................. 156
— — var. discoidea Wink. ........... (BB)
— — var. turcomanica Wink. ...... 155
— latifolia Gilib.. .......ccc.cc.ccesseesees 204
— Ledebourii (Sch. Bip.) Schischk.
saiesvicsdveriepancch dodanssere enters eaten Tae ENT 313
— leucanthemum (L.) Dest. ........... 143
— limosa (Maxim.) Kudo............... eg)
— matricarioides (Less.) Porter. .... 150
SS imaritima| Li ccsaccsssscseevenseess decors 177

926

880

— — ssp. limosa (Maxim.) Kitam.

wi SROs ceca cheirtenth 1, Soebtenaeteiasbacts 178
— millefoliata (L.) Dest. ...........0.++ 348
— myconis (L.) DEST. «0... 147
— oleracea Buch.-Ham. ................. 135
— parthentum L...css.ic.-ccésisindscsoronernes 204
— phaeocephala (Rupr.) Stefansson. .
dot hiivis daseeeannesses,nccnuesti enor cnaapl cans 171
=I DFACCOG WIGS .kkicihvessas Pistesetecanee 183
— pulverulenta (Willd.) Poir. ........ 209
=—| DUNCIALA DEST... cisxsszcesusecpsuacbestonns 215
— Raddeana Wink. .............:cee 151
mm, PECUN BA Ligier schsst evayes th becattenaeaties 148
— segetum (L.) Schrank. ................ 136
—=—serolna ((1c;)) DESiAes sssctescsscasss<<> 139
— sive Chemaem. nanocephalum
PRUE WMiceeeetgec aces cece oapee rea veevactssateasacs 181
== SONSOFICA BOC! 5 dacg rreesiereateadss 313
— spathipappus Winkl................. 262
— suaveolens (Pursch) Buch. ........ 150
— subglobosa Krasch. .......:..cccce 156
— tanacetifolia DEST. ...........eee 197
— Tchihatchewii (Boiss.) Voss. .... 174
Matricaria tetragonosperma (Fr.
Schm.) .Harasi.ds:. 5.26 eee 170
— turcomanica (Winkl.) Pobed.....155
— Tzvelevii Pobed................. 149, 871
Matricarioides Tzvel., sect....312, 876
Mausolea Be. .........:.::ccccscsseeseeeeeee 613
— eriocarpa (Bge.) Poljak. ............. 631
Mesadenta: Raf. ji:.0:35<-sscce5--Pecseefores 685
Microcephala Pobed. ...................... 152
— lamellata (Bge.) Pobed............... 155
— subglobosa (Krasch.) Pobed. ..... 156
— turcomanica (Winkl.) Pobed. .... 155
Millefoliatae: DC, Sect: ¥...:00.--t. teases is
Millefolium C. Koch <...ic.scccscnccpscecerss 75
— officinale Giildenst. ...............00 719
— vulgare Giildemst. ...........:cceseeee 719
Montana Maguire, subgen............... 663
Montani Schischk., sect. ....... 749, 883
Myconia Neck. ex Sch. Bip. .......... 146

— chrysanthemum Sch. Bip........... 147
— myconis (L.) Briq. and Cavil. ..147

Moyriagyne: VSS a5 ise sors cscn stro tophecn os se 421
NETL OSS Sec tas tdeci cane ceneaes stannstrecs 422
Nardosmiia Cassy ..:.5...0iissescescarstoness 645
Nardosmia subgen. .........::sccesseseee 647
== "Nardosmiia: /SeCte wciecscsisticdensccdecacs 647
— ‘angulosa\\Cass;...i.:..uttisereencees 648
— Billingsiana Fisch. ..........ccce0ee 654
— Déorfleri (Hay) Kuprian. ............. 647

== FOMIN BOMdZ:,.:.c-cscsc<ne. ote ee 648

— fragrans (Vill.) Rehb. ......00.. 647
= frigida ,aucts Asie Bia cae 649
= = Bodkaetiiectirgies ees 648
— frigida (L.) Hook. ..........cceeee 649
= frigida: B. LAD. idaink...taperteeens 649
— gigantea (Fr. Schmidt.) .............. 653
<= glacialis: LaDy sciscssanenorssdlinnbeal 654
— Gmelini (Turcz.) DC. ................. 653
— himalaica (Kitam.). .................6 653
— japonica Sieb. and Zucc............ 652
— laevigata (Willd.) DC................. 650
— laevigata subfoemina DC.......... 647
— palmata (Ait.) Hook. .............. 651
— populifolia (Smith.) Schischk. .653
—saxatilis, Turczs::.....os.qs.-sneveontens 647
= ispuria Less: :. ened nines 643
— StraMined CASS. .....cscscceseeeseeseeeee 650
NeesiasSpreng)i:.. 52a. 127
— maritima (L.) Spreng. .............. 128
Neaceis) Cass... 2).cicn. i. BEG Hic 682
Neopallasia Poljak. ..............:0:0:00 636
— pectinata (Pall.) Poljak. ............. 637
Obejacae° DG. SECEN. Ait Asienttntres 779
Oliganthi ‘BGiss:}. sectz.:2kreee 739
Oligocephalae Pojark., subsect.

No daeid sible Socata Wee ee nUne Ro 820, 892
Oligosporus. Cass: Xtkse.. Sisk ee 525
—Vaffinis Lessin. Rea Rs 551
— campestris (L.) Cass. .....cccsee $33
— condimentarius Cass. .........00+ 530
— scoparius (W. and K.) Less......561
Omalanthus ‘Bessy 2a Baia wet 324
Omalotes\DGSRReL As, 324
Omalotes (DG.) Tzvel-} sect. Xie. 324
Otanthus Hoffmgg. and Link. ....... 127
— maritimus (L.) Hoffmgg. and Link.

Stevie cde EE 128
Othonna palustris L. ........:ccccceee 752
—‘sibirica Litt: BORO 807

Paniculatae Pojark., subsect. 827, 893

Pardalianches Tausch, sect. ............ 676
Parthenium (Briq.) Tzvel., sect. .....201
— matricaria Gesn. ex Rupr. ........ 204
P@tastieS ROM, iscccscseceyacnaceseteee eee 645
Petasit@s Mine onc gecceeccc coerce 642
== AIDUS (las) CACKEM: vecceecoesr aster eeaee ss 644
— amplus Kitam. ..........:c:ceeceeeseee 652
== {FIR IQUS EEIESs ons ceryscesnnseosctressatt oars 648
— georgicus Mand. .........::0eee 643
— giganteus Fr. Schmidt................ 652
— glacialis (Ldb.) Polunin. ........... 654

927

— Gmelini (Turcz. ex DC.) Polunin

Bd eS A cena 653
— himalaicus Kitam. .........0......00 653
— hybridus (L.) Gaertn. .................. 643
— hyperboreus Rydb. ......... 650
— japonicus Kitam. .............cccc0 652
— — F. Schmidt. wi... ncn. 652
— — var. giganteus Makino. ........ 652
— Kablikianus Tausch. ................... 645
— laevigatus (Willd.) Rchb. .......... 650
SStniveus: Labi Sere ead, Slat 643
— officinalis Moench. .................6.. 643
— palmata A. Gray. .......... eee 651
— Saxatilis Kom. ......c.. cscs 647
— spurius (Retz.) Rchb. ................. 643
— Tatewakianus Kitam. .................. 651
— tomentosus (Ehrh.) DC. ............. 643
Phaeocephala Pobed., sect. ............. 168
Picrothamnus Nutt. gen. ................. 631
Pinardia‘auct... penwats..mciekewn 131
— coronaria (L.) Less. ..............04. 134
— Roxburghii (Cass.) Less. ........... 135
Polychrysum Tzvel., sect....... 299, 876
Pontia alpina (L.) Bub. ................ 228
Pseudo-Chamomilla DC., gen. ....... 181
Pseudohandelia Tzvel. .......... 361, 878
— umbellifera (Boiss.) Tzvel. ........ 363
Pseudo-oliganthi Sof., sect. ............ 742
BRTAIBA DG sasicavssicisszaterninieisien 104
Ptarmica (DC.) Koch., sect............. 104
StachmingiaEdys teases. 108
= Stalpina aucts.....c. Rene. sevcdoes 117
Sf DCist... SAR. Qn) 119
— — f. tenuisecta Kry). ...........0.... 117
S2sbiserrata DCs: A tet. inde 106
— barealistDG)). ira. 81
— borystenica Klok. and Sakalo. .112
— cartilaginea Ldb. ..........0.ccccce 114
— cartilaginea Ldb., ssp.

septentrionalis Serg. «20... 115
=*Serandiflora( DC...o.M2. ees 110
— Impatiens DC. ......cceeceseseseeeeees 116
— integrifolia Gilib. ........... eee 108
=“ireutiana , DCiiee. aki coks 114
— kamtschatica Rup. ..........:0008 122
S2eK rylovit. Sere in. ara toek 117
<= jlenensis) Turez.. 14. Aue ees 114
= ‘lingulata,. DOi.!.0.0): SR ee. 105
— macrocephala Kom. .............00+ 111
— mongolica DC. ...cccscccseceeeeeee 119
— — var. lilacina F. Schmidt. ..... 120
— ptarmicoides Worosch................ 121
=\sibirica LAbIM2 eel aes 119

— SPECIOSA AUCL. 0... eeeceeeeeee 109, 115

881

Sa DG i istnennnteatite-.ccik. avenee eae 114
— — Stank. sa. Ss.. maka’ 115
— tenuifolia Schur. ........:...:cccceee 123
— tenuisecta (Klayl.) Serg. ............ 117
— vulgaria Blakw. ........cccccccceeseeeees 108
— — 6. Cartilaginea DC............... 114

— — Clus. var. cartilaginea DC. 112
— — DC. var. pycnocephala Trautv.

onecneneo RSS, RE AE. BAY ces 112
Pyrethrellum Tzvel., sect. ...... 214, 872
Pyrethroides Tzvel., sect. ...... 282, 877
Pyrethrum Zinn.\ ................... 184, 871
Pyfethrom secln.i0.:(...0s.asatnase 232
— sect. Dendranthema DC. ........... 364
— sect. Chrysoglossa C. Koch......330
— sect. Eupyrethrum Sosn. ............ 232
— — subsect. Balsamita (Mill.) Sosn.

J secsedesteovopasensinegesssess tte, METAS 196

— sect. Gymnocline (Cass.) Sosn.
subsect. Leucogymnocline (Sch.

Bip:) Sosns2vs... shies 193
— — subsect. Xanthogymnocline
(Sch. Bip.) SOSM......s0.-+-:0--s.tor 330
— sect. Gymnoclines DC................ 330
— sect. Leucoglossa DC. ............ 232
— sect. Leucogymnocline (Sch. Bip.)
Boiss 013. isha. lee 193
— sect. Tanacetum (L.) Boiss. ...... 325
— — subsect. Eutanacetum (DC.)
OST. .....cssec--stibee sc. tase ereneeeee eee 325
— sect. Tridactylina DC................ 389
— sect. Xanthoglossa DC. ............. 330
— * Chrysanthema M. B. .............. 330
— abrotanifolium Bge. ex Ldb......250
— achilleifolium auct. .........c0:cccee 29
SEC MLB) ie ctccccacsecge cave tees teh cers 351
— — ssp. Wisockianum Pacz. ex
Cetin, Mi. 90-5. sta eh. tate os 349
— — var. discoideum Kar. and Kir.
nanacthenanccarecengasseldtahtberanree oR eRe See 341
— Akinfievii AleX. ...........:cccscceseeeseees 357
— alatavicum (Herd.) O. and B.
Fediseh. 721;.,\..:.yaastss-.eeetnesaeeontane 249
— alpinum (L.) Schrank. ................ 228
— — ssp. cuneifolium (Murr.) Tzvel.
<astccnncererenssadgasie ste RAeE I. te eee 229
— — ssp. tatrae (Vierh.) Tzvel.....228
— ambiguum LAD. ........ccceeeenees 168
— arassanicum (Winkl.) O. and B.
Fedischises4. 2. Gasvneb- ope ee eeeiet: 259
— arcticum (L.) Stank. ......... 386, 387
— argenteum Adam ex Hoffm. .....209
— — (Lam.) Boiss. «00.0... 361
— — var. tenuisectum Boiss. ........ 360

928

882

— armenum C. Koch. .......:.c.:eeseeeee 352
— aromaticum (Rupr. ex Boiss.)
"FeV lickicctssc6.. SEAT... RE 222
— athanasia (Bess.) Boiss. ............ 406
— Aucherianum auct, .........ccccccce 238
— Aucherianum DC. ..................05.. 239
— — var. oxylepis Bordz............... 238
— balsamita (L.) Willd. ........0.00..... 196
— — var. tanacetoides Boiss. ....... 198

— balsamitoides (Nabel.) Tzvel. ...198
— Baronovii Krasch. and Poljak. .394

— Baumannii Stev. ......ccccecceceee 348
— bipinnatum (L.) Willd. ............... 324
— Bornmiilleri Hausskn. ................. 229
— breviradiatum Ldb. .................0. 178
— Buschianum Sosn. .............00000000++ 205
— canescens (DC.) Boiss. .............. 336
=] GAPE MILB 5 0scecsserovnaasceucesvevsess PANG
— caucasicum Willd. .....0.......ce 161
— — B. Szowitzii DC. ue 166
— ceratophylloides auct. ..........0+ 220
— chamaemelifolium (Somm. and Lev.)
SOSMeis crschvoosvacs. s,s Ae Mapa IR 219
— cheilanthifolium Sosn. ............... 352
Pyrethrum chiliophyllum F. and M.
eo ATC ASE ee 352355
— — var. oligocephalum (DC.) Sosn.
fscvavst REAR. Sid WARREN. 352, 354
— * Chrysanthema M. B................ 330
— cinerariifolium TZV. .......ccccee 213
— Clusii Fisch. ex Rcehb. .........0..... 234
— coccineum (Willd.) Worosch. ...218
— corymbiforme Tzvel.......... 234, 873
— corymbosum auct. ............. 234, 235
— corymbosum (L.) Willd.............. 232
Sete VAT LAD csscnctsriccssesycesttortetee 235

— — var. oligocephalum Lipsky. .235
— — var. tenuifolium (Willd.) Ldb.

LAS cat PRIA D ce. te 234
— crassipes Stschegl. 10... 338
— daghestanicum (Rupr. ex Boiss.)

Rupr .nexsFlerovirit siete nee 222
— daucifolium (Pers.) Ldb. ............ 230
— — Stev. ex Karel. ou... 220
— decipiens Fisch. and Mey. ........ 182
— Demetrii Manden. ..........0.00. 208
— discoideum LAD. .......... cee 313
a me | MB ipsscessaeseesvevalteti de, MUON 349
— ==, Winkl,)... JARI. AAR ies 156
— divaricatum (Sosn.) Sosn. ......... 207
— — var. monocephala (Sosn.) Sosn.

ssenverNGsd giabéceonutee df MUAED.» ROMER 208
— djilgense (Franch.) Tzvel........... 260

— duderanum Boiss. ........00.ccc000000- 332

— dumosum auct. .........0c000 355, 356
— elongatum F. and M. ................ 173
— Fischerae (Aitch. and Hems].)
BUSED a. 0aseses RESIS 8 nd PRONE 287
— flavovirens BOiSS. .......c.ccsceccesesee 334
— frutescens (L.) Willd. ........ eieeehied 130
— fruticulosum Biehl. ..................... 209
— — Fenzl. ex Boiss. ..................4 231
=='Galae.M...Popsnesi... 2a 242
— glanduliferum Somm. and Lev. 205
gracile SOSMs)vivwsp versions. denied 239
— gracillimum (Winkl. ex O. and. B.
Fedtsch.) O. and B. Fedtsch. ....259
— Griffithii (Clarke) Buser............ 262
— Grossheimii Sosn. .....0......0:.:000 206
— Heldreichianum Fenzl. ex Tchih.
snowuusnckaaviney at tOucdealsteed t= BA ae ee 231
— hissaricum Krasch...................... 241
— indicum (L.) Cass. ........ccsscceseeee 371
— inodorum Moench. .............:.0000 175
— — var. nanum Hook. .............0... 171
— jacobeiforme C. Koch................ 215
— Johnstonii (Hemsl.) Buset......... 317
— Karelinii Krasch. ........00.........08 254
— kasakhstanicum Krasch. ............. 343
— — ssp. karataviense Krasch. ....341
— — ssp. saxicola Krasch............. 342
— Kelleri (Kryl. and Plotn.) Krasch.
dates esis apes Gove suunevaneToeneuvercscettets ene 212
— kermanense Bornm. ...............0... 286
— Khorassanicum Krasch. ............. 358
— /Kirilowtt TUYGCZ.:..,-0..0088 0. 390
— Kittaryanum C.A.M. .......:.:000 345
— Komarovii Sos. ..........ccseeeeee 224
— Kotschyi BOisS.............scseseseeees 229
— Krascheninnikovii (Rubtz.) Pavl.
REA Ta Bee te Mee 293
— Krylovianum Krasch. .............0 250
— kubense Grossh. .........cccceeceeeeeeee 209
— lanuginosum (Sch. Bip. and Herd.)
TZVEl ss sccccesdtsscccvocettvcsn tesseneentte 252
— latilobum (Sosn.) Sosn. ............. 203
— leontopodium (Winkl.) Tzvel. ...260:
— leptophyllum Stev. ............00+ 211
— leucanthemiflorum Boiss. and
Huet. Ce 239
— leucanthemum (L.) Franch. ....... 143
= \Lilae.. Bord2Z:. :siicinc ye AEA 215
— longipedunculatum Sosn............ 354
— macrophyllum (W. and K.) Willd.
1s see A RE AR IAN od 194
— var. angustifolium Ldb. ............. 195
— majus (Desf.) Tzvel. .......0...006 198
— Marionii AIbov. .........00...ccccee 216

929

— maritimum Smith. .........0..00.0. 177
— Meyerianum Sos. .........:.:ccce 220
— Mikeschinii Tzvel. ............ 243, 873
— millefoliatum auct. ...337, 346, 352
— — (L.) Willd. wo... eee 348
— — var. achillefolium (M.B.) O.
and) Bi: Fédiseht 2... cca 351
— — var. macrocephalum Ldb...........
coed bepacettond BAL ee, WY, seats 345, 348
— — var. microcephalum Ldb. .....348
— — var. oligocephalum (DC.) Boiss
sh Aiitha tbat UB E. DAL AI 354
— — var. tanacetoides (DC.) Trautv
PE elt. EBS. MA REM 337
— mindshelkense B. Fedtsch. ........ 341
— mucronatum auct. .... 304, 305, 306
— — Rgl. and Schmalh.................. 302
— multifidum (Desf.) DC. .............. 197
— myconis (L.) Moench. ................ 147
— myripohyllum auct. .......0cce 331
== .CAcMietiieeieth). ahead 332
— — var. eriocephalum Boiss. .....332
— — var. subvirescens DC. .......... 331
— — var. variegatum Boiss. ......... 833
— neglectum Tzvel. ............... 257, 874
= NUVGLE AGAM, on corsapsccaspesesces tase senenaie 18
mee UTEVEUND Wek Beira o22. ct onscnsaett tonne ate 209
— niveum var. divaricatum (Sosn.)
RS OST Aub Pre it See RR is act 207
— odessanum KIOK. .........0ccceeees 348
— oligocephalum DC, ..........:000000+ 354
— ordubadense Manden. ................. 230
— orientale Willd. ................:ceeee 32
— oxylepis (Bordz.) Tzvel. ............ 238
= PACZOSRIULELELMS. ..ccdsscss-n0resenesseeee 349
— Pallasianum (Fisch. ex Bess.)
ESO ire ae 402
— palmatum (Vent.?) C. Koch......207
— palustre Willd. ...........ccccceeeeeeee 215
— pamiricum O. Fedtsch. ............... 518
— paropamisicum Krasch. ............-. 310
— parthenifolium Willd. ................. 203

— — var. anthriscifolium Sosn. ...204
— — var. canescens Boiss. . 207, 209

— — var. discoideus Boiss. .......... 206
— — var. latilobum Sosn. ............. 203
— — var. macrophyllum Sosn. .....204
— — var. peucedanifolia Sosn. ....202
— — var. pulverulenta (Willd.)
Keanelty. Steerer cestints titesateaeescharsetteg 209
— — var. sevangensis Sosn. ......... 207
— — f. divaricata Sosn. ................ 207
— — f. monocephala Sosn. ........... 208

— parthenium (L.) Sm. .................. 204

883

— — var. aureum hoft...............06 205
— — var. breviradiatum Sch. Bip.205
— var. discoideum Sch. Bip. ......... 205
— parthenium var. flosculosum DC. ..

nanstangsveansyreeteeticteth «fH SAIN OT HI 205
— — var. laciniatum hott. ............. 205

— — var. longiradiatum Sch. Bip.205
— — var. parthenifolium (Willd.)

Lip sk yussicewseicssovansscdtetteeeeseeecandeeaetees 203
— — var. selaginoides hott. .......... 205
— parviflorum Willd. ............ 175, 183
— paucifolium C.A.M. .....cccceeees 220
— pectinatum Fisch. ex Turcz. .....251
—=\—Hausskn..........0205:..R8 0 358
— peucedanifolium (Sosn.) Manden.
seGhxnnsisunanaacoisxckanse titers «ME CE ae 202
— pinnatifidum Willd. .................... Lat
— platyrachis BOSS. «0.0... 311
— polyphyllum Boiss. .............0006 329
— ponticum AIDOV. ........:ccceeeeeees 236
— poteriifolium Ldb. .............00.00. 235
— pracaltum (Vent.) Tzvel............. 208
= Spraecox. .MB2.c/0048.. 2aR Ree 183
— pseudoachillea (Winkl.) Bornm. ©
a RRORE ARE. RE 328
— pseudoanthemis Boiss. and Huet.
ee, eee 239
— ptarmicaefolium Willd. ...... 18, 109
— pulchellum TurCz. ............::cc08 251
— PUlChrUm AUC. ........cccceccereeseeseeeees 254
— pulchrum LOD. ............c ee eeeeeeees 253
— pulverulentum Willd. .............000 209
— punctatum (Desr.) Bordz. ex Sosn.
wi geswindacensidonapton ete ES Oeics RENN 215
— pyrethroides (Kar. and Kir.) B.
Fedtsch...ex.._Krasch. t2s2a\x-ane 256
— — ssp. songaricum Krasch. ...... 255
— — var. glabrum B. Fedtsch. ex
Krascht,.2....1Rk. Wah Been 247

— richterioides (Winkl.) Krassn....255
— richterioides var. virescens

(Winkl.) O. and B. Fedtsch. .....254
— Roopianum (Bordz.) Sosn. ........ 239
=froseum.auct. ¥.1i8%1... eas 218
— roseum (Adam) M.B. ..............05 217
— roseum var. Adami Trautv. ....... 218
— — var. album Grossh................. 219
— — var. carneum (M.B.) Trautv.

Banke. Aho... 18.. SSE 217
— — var. chamaemelifolium Somm.

and. Levis. aakt sn... SRY 219
— — var. leucanthum C.A.M. ...... 220
— rotundifolium Waldst. and Kit. ex

M...». POD. sssssensavscdnesveinass 00. teeeteee 142

930

884

== Roylet, DC, dyed terest.» 288 av 263
— santoanum Krasch., M. Pop and
Vive, ceri ..thcea. .irusin teat he soe cee 311
— saxatile Kar. and Kir. ............ 344
— sclerophyllum Krasch. ........+0 347
— scopulorum Krasch. ........00000068 342
— segetum (L.) Moench. ................ 136
— Semenovii (Herd.) Winkl. ex O.
and. Bi. FEdtsCh ....c0<.0:s0000eetee 243
— sericeum (Adam) M.B. ..............237
— serotinum (L.) Willd. .............. 139
— setaceum Rgl. and Schmalh......304
—‘setifoliunt Rel. ixcit..cnaealss 304
=="sevanense SOsn. .......s:cenetet..avs 207
— sibiricum (DC.) Stank. ............... 377
— silaifolium Stev. .............ceeee 220
== {simulans. Pavliisikt..tageetisneeeet 283
— sinense (Sabine) D.C. ................ 373
— — a. sinense Maxim. ............... 374
-— songaricum Tzvel. ............. 255, 874
— sorbifolium Boiss. ............0....00 195
— Sosnowskyanum Crossh. ............ 354
— Starkianum AIbov. ..................-.. 236
— — var. foliosa Albov. ............ 236

— subcorymbosum (Schur) Schur .234
— suffruticulosum Fenzl. ex Tchih. ...

Sci sil a etl a tava Pee ier 8 25)
— svanicum M. Pop. ....cccccceceeerees 202
— Szowitsii BOISS. .........cccccesseeeeees 331
— — (C. Koch.) Grossh. and
SCHISCHK ., ..cccscesocssdbeeatas terrors 329
— tabrisianum auct. ...........cccceeees 355
—si== Boissievaik.).eaeak LA. ceuceeoeees 335
— tadshikorum KudT. ............:0000000 299
— tamrutense (Sosn.) Sosn. ........... 334
— tanacetoides DC..........cccccccccceeree 337
— tanacetum DC. ........cccceccecceeeeeeeeees 198
— tauricum ZeleM. ............cccccceeeeeees 232
— Tchichatcheffii hort. W. Robinson.
Me Met A Ms ihe 174
— tenuifolium Willd..............0..0.... 234
— ENUISECtUM AUCL. .......eeeecceeeneees 230
— tenuisectum Boiss. ............00.05 229
— tenuissimum Trauty. .............0068 355
— tianschanicum Krasch. ...............247
— transcaucasicum Sosn. ............... 352
— transiliense (Herd.) Rgl. and
Sehmal hy, 44. iL ARR. ARE evs 258
— transiliense var. glabra Franch.
PEEISIS.. MCS LATTES, RR, 28 260

— — var. subsessile Rgl. and
Schmalhi,,.).cixmininiet. ket 256

— — var. subvillosum Rgl. and
SCHMAIMN, oi, sancaacennsesasaasnes MMe Rod 258

— — var. tomentosum Rgl. and
Schmalh. ...2:..2i sae wea 256

— tricholobum Sosn. ex Manden..223

— trichophyllum (Rgl. and Schmalh.)

Bornm:, and (Sint. 20277. 62440 363
— —=) Sosnis...dewd wantin. 223
— turcomanicum Krasch.................358
— turlanicum Pavl., .........c:ccccccececeee 341
— uliginosum W. and M............... 139
— umbelliferum (Boiss.) Boiss. .....363
— uniflorum F. and K. ............64 356
— uralense Krasch. ..........::cs0cee00es 346
— — var. tomentosum Krasch....... 345
— urgutense M. Pop.\ ............ 308, 874
— velutinum Fisch. ex Trautv. ...... 324
— vulgare (L.) BOiSS. «ccc 326
— Walteri (Winkl.) B. Fedtsch. .... 359
— Woronowli SOs. ...........ccccccceceees 334
— xanthomelanum Krasch. and

Grossh..,.......220uitee eee 32
— Zawadskii (Herb.) Nym. ............ 377
Quadridentati Boiss., sect. .............. 725

Racemiferae Pojark., subsect. 826, 893
Rhytidospermum Sch. Bip., sect. ...175

— inodorum Sch. Bip. ........0:0c0 | Wf)
Richteria Kar. and Kir. ......... 184, 248
Richteria (Kar. and Kir.) Tzvel., sect.
EET Se oteetten siaceeiee eare 248
— leontopodioides B. Fedtsch....... 260
— leontopodium Wink. .................. 260
— pyrethroides Kar. and Kir......... 256
— — var. djilgensis (Franch.) O.
FPOGUS CHES stercterscace: creceerevaconryecens 260

— — var. subvillosa (Rgl. and
Schmalh.). O. and B. Fedtsch. .258

Richteriopsis Tzvel., sect. ..... 237, 872
RumataFed® ‘sect. cc 16, 865
Santolina Vo. 22 n.ssitc5.-sgsachctsceatoncss 125
— chamaecyparissus L. ............... 126
INCOR Lea orev saon cass satan cn s*-nnzene +s 126
— maritima (L.) Grantz. ..........-...... 128
— tomentosa Lam, ..........0....scceeeeeees 128
——= 4 VITIGIS . WANG oss. cctsanccascdnnans aoapuner ase 127,
Santolinoideae .DC..,. Sect..,..........--.-. 100
SCGDOSY BIOUID oi hectare as 750
Scaposi (Hoffm.) Schischk., sect.
ePeaepiiviebin te Maid sorceress FS 750, 883
S@ne cis GAC Ci gek acta ca teoreni tess he -cace sa
plesete 788, 836, 844, 893, 894

931

Senecillis (Gaertn.) Kitam., sect. .........
senstoade cba avarice ch. tydhes dente 836, 893
— (Gaerth.) Pojark., subsect. 844, 893
— Gaertn...... 788, 836, 844, 893, 894
— carpathica Schott. «00... 750
— — and others... eee 848
=" Fischert, Kitamius...20:2.)...108..3: 802
— glauca auct. wo... 843, 848
> (i — Gaeringteis tee wiihewas 847
— — (podolica) Schott and others.
wp ES. ces ctans tae itl Gave 848
— — ssp. carpatica Hayek............ 848
— — var. transsilvanica Schur.....848
— heterophylla Schur. .................... 848
— Hodgsonii Kitam. ............cce 798
— jaluensis Kitam. .......0..cccceeee 826
—- Schmidtii Maxim. ...........cccee 851
Senecio Benth. and Hook. f. .......... 683
Sétieeio, L............d06. eens... 699, 881
— subg. Cacalia Maxim. ................ 683
— sect. Cacalia Benth. ................... 697
— sect. Heloserides DC. ................ 752
— sect. Ligularia Benth. ...... 788, 797
— B. Euligularia Fr. 0... 797
— gr. Ligularia B. Euligularia 3,
Corymbostikr, <nstsex xt its. ti... 798
— — — 1. Racemosi Fr. ..... 801, 893
— abrotanifolius Wahlb. ................ 750
— — var. carpaticus Hayek. ......... 750
— aconitifolius Turcz. «0.0... 698
— acrabatensis Franch. .................. 728
— aksuennsis Pav. ....0...cscseeeeee 135
— alabugensis Wink. ..............00. 786
— alpestris LAD... cece 764
— altaicus Sch. Bip. ............c0 843
— ambraceus TurCzZ. ..........0ccee 715
— amphibolus C. Koch. .........:000. 728
— amurensis Schischk. ......... 757, 883
— aquaticus Hill. ow. 788
— aquaticus Huds. a. barbareifolius
Wimm. and Grab. .............. 717
— aquilonaris Schischk......... 763, 884
— arcticus RUpf. ........cccceccsesseees 752
= Garenarius .M.B....<cis..hbiah see 711
— arenarius Thunb. .......00.0.. cee W112
— argumemsis TUICZ. ......... cece 716
— argunensis f. angustifolium Kom.
SRN ES A LA oa 716
— — f. latifolius Kom. ............0... 716
— armenius Jaub. and Sp. ............. 721
— asiaticus Schischk. and Serg. ...762
— atripurpureus (Ldb.) B. Fedtsch. ....
sovennsde tba dec AML CER Ata, oo. uoeea ctl 771

— — var. aurantiacus Trautyv. ....... A712

885

— — var. discoideus Trautv.......... 772
— — var. flosculosus Trautv. ........ 772
— — var. glabratus Trautv. ........... 772
— — var. monocephalus Trautv. ..772
— — var. pyroglossus Trautv........ 772
— — var. radiatus Trautv. ............. 772
— — var. tomentosus Trautv. ....... 772
— aurantiacus (Hoppe) Less. ......... 768
=4'—. Grosshee iin a). 2a 769
== Kryhn.. iil. angle Tt
— — b. lanatus Rchb............0.0... 768
— — a. glabratus Turcz. .............. 766
— — B. flosculosus Bess. .............. 768
— — B. flosculosus Turcz. ............ 766
— — B. leiocarpus Boiss............... 769
— — B. tomentosus Ldb. ............... 757
— — Y.- tomentosus DC. ..........0 768
— — ¥.- tomentosus Turcz. ............ T75
— — 6. capitatus DC. uu... 768
— — 6. capitatus Turcz...........0..... 775
+ \auratus. DC: tehs...ined. Ree 736
— barbareae folius Krock. ............ TTF
— Besserianus Mind.................008 768
— — f. radiatus Mind................... 768
— Biebersteinii Grec. ........:cceeceeee 733
== \Blinii’ Leveillews. 2A 716
— Bogdanoviczii Kom. ..........0...00 777
— borysthenicus Andrz. ................. 718
— [= 1Giossh. O..aiike ene). as 719
— brachychaetus DC. ........... 759, 760
— — var. longifolius Grossh......... 760
— — var. mesophilus Grossh. ....... 760
— — var. xerophilus Grossh. ........ 760
— bucharicus Winkl................00 729
— Bungei Franch. ...........cccceeseeeee 834
— Buschianus Sosn. ............0ccc00 726
— cacaliaefolius auct.

edhe dce nano 802, 808, 812, 818
<2) Sch.: Bip..:.... deel anced 807
— — Q. speciosa Franch. .............. 802
— cacaliifolius auct. ......cccceeee 803
— var. speciosa Franch. ...............+: 803
— cacaliiformis auct. ..............e 808
= = Rehb.. fill...........:..s0.0bcsdiavctivasee 806
— calthaefolius Maxim................... 800
—) Calvertin Boiss) issiianititts...30e8.c0 780
— campester (Retz.) DC..........0... 753
— — var. aurantiaca Trautv......... 776
— — var. brachychaetus Schmalh.

eo i So SCL TT Rete Ope 760
— — var. fulva Trauty. ........... 776
— — var. pyroglossa Trautv. ........ 769
— CAMpeStris AUC. 1... 754, 757
=} =. Boiss. t2ebe een eee 757

932

886

— compestris var. pyroglossa Trautv.

paioneganctete ica denctcuutscesbeccusn te scemcrcreen 770
— — b. capitatus Schmalh............ 768
— — Bf. aurantiacus Schmalh. ......768
— — Y. capitatus DC. uc 757
— — 6. glabratus DC. ...... ce 762
— Candolleanus Sosn. .........00000000 785
— cannabifolius Less. ............0008 747
— capitatus (Wahlb.) Steud. .......... 768
— carniolicus Willd. .........000.. 748
— carpathicus Herb. ...........cee 750
== 'caspicus. LoeSs: %...2ceiiernteots 731
== \CauCcasicus IDC. eenaas..ds oe 855
— caucasigenus Schischk. .............. 769
— chrysanthemifolius Ldb.............. 783
— chrysanthemifolius Poir. ............ 788
==\cinerana. DES. sere 720
— cladobotrys Ldb. ....0......0. eee 759
— — var. pseudorivularis Cuf......760
— — var. typicus Cub... 759
— conipes Somm. and Lev. ........... 724
— congestus var. palustris (L.) Nyl...

davecunsussuseasensvonareetadtntste: toe 752
— — B. *. intermedius DC. .......... 752
— cordifolius AUct. 0... 726
— coronopifolius auct........... 783, 787
— coronopifolius Desf. .............0... 787

— — var. pinnatipartita Trautv. ..787
— — var. sphacelatus O. Hoffm. .783

— — var. subdentatus Boiss. ........ 786
— Correvonianus Albov. ................ 856
— crassifolius Sibth. and Sm........783
— crispa @ dilatata Ldb. ............. 764
= crispatus.. DC. 22082 amenaeen 765
= erispus. Kittel, \.....dsecd. stuns 765
— cryophilus Schischk. and Serg. 775
—"cyri C. Koch. tote. enna 732
— Czernjaevii, Mind. ........00. 754
—dahuricus Fisch. .......0i2.00420% 747
= 2 Sch Bipyieet.seieas 8% 694
— — Q. ochotensis Maxim. 092, 694
— — B. kamtschaticus Maxim. .....696
Senecio dentatus y. parvulus Ldb. 786
= DOria AUCH 5... ,.ccecseceies teense 731
SS) ED ea See 733

— — var. macrocephala Trautv. ..733
— — var. microcephala Trautv. ...733
— — B. Biebersteinii Lindem. ......733
— — B. macrophyll ; Schmalh. ...733

— — y. umbrosus Schmalh. .......... 734
— doriaeformis auct. .......cccccceee AF
— — var. thyrsoidea Franch. ........ 735
— dubius Lb...) sat: Aa ateen 781

— erraticus (Bertol.) Hegi. ............ 1) BZ

— erraticus Bertol. «0.0.0... cece 717
— — Bertol. ssp. barbareifolius
(Wimm. and Gr.) Hegi. ............. ii wy]
— erucaefolius LAD. .........ccccccceee 715
— erUucifolius AUCt. .....c..ccccceeeeeeeeee 719
== erucifolius. Lis... RR RRR 710
— — var. (flor. majoribus) Ldb. ..715
— — var. mollis Trautv. ........000... 711
— erucifolius var. viridis Trautv. .710
— — Q. canescens Trautvy. ............ 7A
— — B. latifolius Boiss. ...... 411,078
— — Y. grandidentatus Boiss. ...... 711
= leuxinus! Minden OR aes 783
— Fedorovianus Schischk. ............. 778
— ferganensis Schischk. ....... 720, 881
— flammeus Turcz. ....0...... cece 767
— — ssp. glabrifolius Kitam. .....767
— — var. glabrifolius Cuf........... 767
— — var. typicus Cuf. 0... 767
— — f. glabrescens Hara. ............. 767
— flosculosus Gilib. 0.0... 780
— fluviatilis Wallr. ou. 745
— Franchetii Winkl. «0.0.0. 729
— frigidus Less.,..25. 023228208. 771
—~ ——fToreziainn.. th. SRR ah 115
— — var. discoidea Trautv. .......... 771
— — var. radiata Trauty. .............. ag
— Fuchsii Ke Gmel!..22nc eae 745
— Fuchsii Grosst. .......ceceseeeeeeeee 743
— — y. Turczaninovii DC............. 742
— fulvus (Stev.) Schischk. ............. 770
— glabellus (Turcz.) DC. ............... 762
— glacialis Marco. .........:cscsceeees 784
=Gmelini LD. .<csnsse: i ttes. teehee 788
— grandidentatus Ldb. ................ 711
— Gundelius C. Koch. .........000cee 721
— hajastanicus SOF. .......:ceeeeeeees 745
— hieraciiformis Kom. ..........00 765
— hypoleucus Lab. ...........0ceceseeee 736
— Igoschinae Schischk. ........ 163, 885
— Iljinii Schischk. ................. 743, 882
— incanus SCOP. ......i.. cise 748
— — ssp. carniolicus (Willd.) Br. Bl.
and. Dostal <0... Re ZR 748
— — var. carniolicus Heat............ 748
— integrifolius AUct. .........cceccees 159
— — GTOSSD. oo. eeeeeeeceeseeeeeeseeeeeeees 760
— — (L.) Clairv. ssp. atropurpureus
(Ldb.): Caf: ne 2a. TT
— — — var. robustus (Herd.) Cuf.
venue oclhls. Satte c Meee, . 2 775, 776
— — — var. taimyrensis Culf. ...... 775
— — — — f. Nekrassovii Cuf. ....777
— — ssp. capitatus Cuf. .............+. 768

933

— — ssp. capitatus (Nym.) Cuf. var.

aurantiacus Cuf. f. Gmelini Cuf.

scvteucnsesoieteevesiavds edt EOE ER. SEEMING 766
— — var. pyroglossus Cuf. .....770
— — var. pyroglossus f.
primulifolius Cuf.........20ikite.. 0 770
Vacobueaaledb. xt oe.i3...cek..22 710
Jacobaea’ Wasi... ces BAS 715
— L. var. ambracea Trautv. and
i CS RSS a ESAS 3 SRR SL ATS 715
— var. borysthenica Trautv. ....718

— var. erraticus Fiori and Paol. ...

Betande hi eiaideiatavesceascevuewsadh «sod PS 717
— — var. typica Trautv. ........00... 715
— Jacobaea {. grandiflorus Turcz. ....

BRIA. SN, ne EU 715
— — y. farinosus Lmdem. ............ Apis
— — Y. grandiflorus Turcz. .......... 716
— Jacquinianus Grossh. ...........0... 743
— jacuticus Schischk. .....0........ 779
= jailicola Juz.228.....)..20siils. 757
— kamtschaticus Kom. ............0000+ 776
— karataviensis Lipsch. ..........000+ 824
— karelinioides Winkl. ................. TED
— Karjaginii Sof. occ 770
— Kawakamii Makino. ............0.0.. 761
— kirghisicus DC. ....cccccseeeeeeee 733
— Kochii Lab. «0.0... 753, 765
— Kochii a. Schkurii Ldb. ............788
— Kolenatianus C.A.M. ...........00. 728
— Korshinskyi Krasch. ............0.0+ TZ
— Krascheninnikovii Schischk. .....783
— Krylovii Schischk. ..........0...0. 715
— kubensis Grossh. .............0.:e0 724
— lanatus S.G. Gmel. ......00c 736
— lapsanoides DC. ........cccccsseeeeees 744
— Ledebourii auct. 0.0... 838
mm SCN BAP. Asses. Lobloeiseddscsoses 837
— — B. minor Franch. ...........00.. 839
— lenensis Schischk. ............. 758, 884
— Lipskyi Lome.i....3....G0ae0Nae 722
— Litvinovii Schischk. ..........0.00.0.. 747
— longeligulatus Lev. and Vant. ..767
— longiradiatus Trautv. ................. 728
— Lorentii Hochst. «0.0.0.0. 721
— lugens Richard. ........00 ee 788
— lyratifolius Rchb. ........cceee 716
—— \Lyratus, TUrez. ....ce 2-1. 750
— macrophyllus M.B............c08 730
— Malaisei Hult. ....... ee 777
— Massagetovii Schischk. .... 722, 882
— maritimus RChbD. ..........cceeeeees 720
— microphyllus (errore typogr.)

SPLEN Bir vvvecsvrse. dont eeneec eee 731

887

mollis. auct...M. wasinenths...B..ves 711
mongolicus Sch. Bip. ............0. 849
monocephalus Schur. ................. 750
Memorensis ‘Likert yek...see..0 742
— ssp. macer Printz............0... 742
— var. polyglossus Rupr. ......... 742
— var. Turczaninovii Hara. ...... 742
— B. octoglossus C. Koch. ...... 742
— y. Fuchsii Ldb. ue 745
Noeanus(Rupr. tiads!svun..228..00 787
ODSCUTUS DRISChIS eee 742
actoglossusiDC. isintraheans 742
— B. macer DC. ou... 742
Olgae Rgl. and Schmalh............ 728
orientalis \Willdet8). ici e eae 727
Othonnae4yMeB: .22-2..xietats 741
— var. genuinus C. Koch......... 741
— var. rariflorus C. Koch........ 741
otophorus Maxim. ...............0:0000+ 747
ovatus Willd? °2.8.225.asnatiats 745
palmatus: Pally’. snetscvs. casts 747
paliidosus 1.. NEA stg: teet ves 735
— a. nudiusculus Ldb. ............. 735
— B. auratus Schmalh. ............. 736
— B. glabriusculus DC............. 735
— B. vulgaris Ldb. ........ cee 735
— y. hypoleucus Ldb. ............... 736
— Y. grandidens Rupt............... TES)
paluster var. genuina Trautv. ...752
palustris (L.) Hook? .<:i.23sce 1p?
— yY. congestus Hook. ............... 2,
pandurifolius C. Koch................ 724
papposus (Rchb.) Less. .............. 760
paucifolius Gatek .a5)iue:..&..... 733
pauctlobus! DG sick. cst cheeses 741
Paulsenii O: ‘Hoffim..:::ic8ste-.2 730
PavlovitWLipscht 2s);.0.1..2ene ieee 852
peduncularis Griseb. ...........:++ 783
pedunculatus Eg. .........:00000 783
pedunculosus Trauty. ....... (PING P92
Pierotin Miquelam...itars. avisconsteess 757
DiNnnatuse SCHSPBI1p: yest. -.te.-0x 741
platyphylloides Somm. and Lev. ....
avetacnesiéstavee:+ssesetetia es PeMEe vette Ore 740
platyphyllus BOSS. .........:0s000 740
= DCo.:..::... 08. Aee. ates 739
— var. platyphylloides (Somm.

ana ev.) Grossi). «scdivn: Asse. 740
Pojarkovae Schischk. ..............+ 09
polycephalus Ldb. ................:0004 785
ponticus, GrECESCUS ferns senses 711
porphyranthus Schischk. ............766
pracaltus UdD.,......tces- esses 2/10
— f. borysthenicus DC..........4. 718

934

888

— — y. davuricus DC. .......c ce 716
— pratensis. LdD....£.3.0..62.55, ene 762
= '— Maxim: aedokaeles 757

— — var. polycephalus Rgl. .........757
— praticola Schischk. and Serg. ..762

== \Pricei Simps:i). 0G. ee ex TIS
— primulifolius Somm. and Lev. ..770
— propioquus Schischk. ................. 743
— pseudoarnica Less. .....0..........04 723
— — var. kurilensis Kudo. ............ 723
— pseudoaurantiacus Kom. ............ 777
— pseudoorientalis Schischk. ......... 727
— pseudoracemosus Grossh........... 731
— Putjatae Wink. ..........ccc cee 849
— pyroglossus Grossh. ..........0.0:00+ 769
— pyroglossus Kar. and Kir. ......... 770
— — var. macrocephala Lipsky. ..770
— quinqueligulatus Wink. ............. 744
= iracemosus) LADRI EA. eR 733
— racemosus (M.B.) DC. ............... 731
— — Var. araxina Trautv. ............. 7132
— — var. glabra DC. .......cecceeee 732
— — var. Schelkovnikovii Crossh. 732
=i... puberuseD Gav cnniies. Abies 731
— racemulifer Pavl. oe 135
= \apistroidés’ DG J,.At ins... Ree 783
—rariflorus C. Koch. .....ccceeeeeeee 741
— Renardii Winkl. ......00. ee. 749
— renifolius Sch. Bip. .......ccecee 853
— resedifolius Less... 750
— — q. genuina Rgl. «0... 750
— — B. lyratus DC. wo. 750
— — Y. multicaulis Rgl. «0.00.0... 750
— — 6. multiflora Rgl. oe .. 750
— Reverdattoi Sobolevsk. ............... 778

— rhombifolius (Willd.) Sch. Bip. 739
— rivularis (Waldst. and Kit.) DC. ....

dace Sees 765
— var. typicus Domin and Podpera. ..

aavbiowaraceh Ma sole ACHAT 1a. ARR at 765
— robustus Sch. Bip. ..........0.c00 820
— — var. Kareliniana Trautv. ...... 822
— — var. typica Trauty. ............. 822
— — QO. LYPICA AUCL. .....ccesscercssceseee 820
— rupestris Waldst. and Kit. ......... 788
— sagittatus Sch. Bip .ii0...5..800.0: 687
— — Q. pubescens Maxim. ......,....687
— — B. glaber Maxim. ........ 687, 689
— salsuginosus Schischk................ 787
— Saposhnikovii Krasch. ................ VAT
—) SATTACENICUSU LE EAS. BAA 745
— — Bf. sclerophyllus C. Koch. ..745
— saxatilis;.Lom.........10.2...28tnwne 722

— Schelkoynikovii Grossh. ............. 132

— Schischkinianus Sof. .................. 719
— Schischkinii Lipsch. .................... 825
— Schmidtii Franch. ............0:0ccc000 851
— Schvetzovii Korsh. .......0...0...000. 733
— senecillis aucti ncn. itn. ens 848
= ==Maxkim: i... aaa ae 847
— — var. carpatica Janka. ............ 848
— serawschanicus Wink. .............. 729
—)sibiricus Ln filha a 827
— sichotensis Kom. .............:c:c0008 764
— similiflorus Kolak. ............0000006 740
— SONQATICUS AUCL. ........cceeseseeseesees 834
SS RISC TSR ccskcaes cee 831
— — var. hastifolius Wink. ......... 831
— Sosnovskyi Sof. ...........ceceseeee 784
— spathulifolius y. praticola Rouy.
Steet RRS RE RE OM 762
— splendens Lévl. and Vaniot. .....803
— squalidus M.B. ......::cccccseceeeeee 783
— stenocephalus BOisSS. ..........0.04 739
— subalpinus C. Koch. ........00.00.. 726
— subfloccosus Schischk. ............... 760
— subfrigidus Kom..............ccee 7
— subscaposus Kom. ..........ccceeeees 766
— subdentatus Bee. ...........cs sce 786
— subdentatus (Bge.) Turcz. ......... 787
— — var. fasciculatus Litw........... 786
— — var. glabellus (Turcz. ex DC.)
Cufs.2'.. Stetina Sites 762
— — var. pinnatipatitus Bee. ....... 787
— — a. ramosus LAD. ..............000 786
— — B. simplex Lab. 0... 786
— succisifolius Kom. ...............0. 758
— Sukaczevii Schischk. .................. 762
— Sumneviczii Schischk. and Serg.
RE eR LY ee citeece: er ttc 1)
==tsylvaticls Li... 23aee eet 782
— taraxacifolius (M.B.) DC. ......... 725
=HataricusiBessome ss sien 736
— Thomsonii Clarke. .................0.. 834
— thyrsophorus C. Koch. .............. f3z
— tianschanicus Rgl. and Schmalh.
2 REE ee et 749
— Tichomirovii Schischk. .... 772, 885
— tomentosus Host. ...........cccceeeee 736
— triangularis Hook. «0.0.0.0... 788
— tubicaulis Mansf. ...............00000 752
— tundricola Tolm. ................c 7176
— tundricola Tolm. .........0cccceeeee 175
— Turczaninovii DC. ........ ee 775
— turkestanicus Winkl..............0+ 831
— umbrosus BESS. «0.0.2.0... 730, 733
— umbrosus Waldst. and Kit......... 734
— uralensis Gorodk. ...........ccccceceeee 771

935

— valerianifolius WoOIf. ...........0+ 683
— Veresczaginii Schischk. and Serg.
ie Reaate ke Vie HERE Re Sel tia a Ane 764
== VETRALLS HAUG. tei iiatensdsececetsevatan des 785
— vernalis Waldst. and Kit............ 783
— vernalis var. leiocarpa Trautv. .785
— — @. genuina Lindem. .............. 783
— — f. caucasicus DC..............4-+ 784
— — B. pauciflorus Lindem. ........783
— — Y. Nanus BOSS. ........0.eeee 784
———VINGOSUS AUC ssearsereseverersealteattecse 780
Be Nam PDS Acave Ale foo cceeeeseseaecteosent cece 780
Se AVISCOSUS Mis tiecstk etre fcuseserscs-cnctduvaet es 779
— vulgaris L. ......cceccceseecsseeeeeseeeeners 780
Be VAL. OAUDIA) LTAULV 2 \iee-navennnseset 781
— — var. parvula Trautv. ............. 781
— YEFOENSIS FI. .......seseresesesesersesereees 798
SENECIONEAE Cass., tribus......... 638
Senecionidaceae Bessey, fam. ........ 638
Senecionideae Benth., trib.............. 638
— subtr. Artemisieae Less................+- 1
— subtr. VII Artemisieae Less. 4
ATICINISICOE LESS Jeti eel st 129
=='— —.2. Cotuleae Less............... 129
—)— —" 1. Santolineae Less: :::.:::....: 9

— subtr. Chrysanthemeae Less. ........ 1
— subtr. VI Chrysanthemeae Less. 1

Anthemideae: Wass. \v:...c20-sisteer.-t-s- 9
— — 2. Chrysanthemeae Less. .....129
— subtr. Senecioneae Less............. 638
Senecioninae Hoffm., subtr. ........... 640
Sericochaetae Pojark., subsect.

Rte Ae eet eee tots Bee 837, 894
Seriphidium (Bess.) Rouy, subgen. 562
Seriphidium, SEC. ..............-.:ssseeen 573
Seriphidium Bess., S€Ct. .........000 562
Solidago palmata Pall. ...........::00+ 747
Spathipappus Tzvel............... 261, 875
— Griffithit (Clarke) Tzvel............. 262
— porphyrostephanus (Rech. f.)

MTPZVEl es csceerecvatavsckinscesees sens eeaeteteveseras 262
Sphacelatae SOsM., SET. ......-:.s:0ss00 16
Stellerianum (Rydb.) Poljak., sect. 455
Stenostegia Pojark., sect........ 820, 892
Stereochaetae Pojark., subsect..............

BAS SRM RR 849, 894
Subbulbosa Tzvel., sect. ........ 279, 877
Syneilesis Maxim. ..............::000008 697
— aconitifolia Maxim. ............. 698
Synosma Raf. .......scscesesesessseneneneseess 683
at hastata PGjankeeen.Atin.snisene-cets 687
— Komaroviana Pojatk. .............0+ 691

— robusta Pojark. v.0ic..cieeee cesses 690

889

Tanacetopsis Tzvel., sect....... 300, 876

TANACEtUIM AUC iii isess-.nciecssesous 398, 411
Tanacetum L. ...........:.:csee 317, 878
==> ML ANACEMIMINSE CE. sveyuescvecvonivesaeanssss- 325
— sect. Balsamita (Mill.) Sch. Bip.
beastie k See ER 196
— sect. Eutanacetum DC. ............. 325
— sect. Leucoglossa § 3 Alpina Sch.
BID. oscscecatsceceasth berets utoneeeee ae 227
— sect. Leucoglossa § 2
Macrosperma Sch. Bip..........-... 232
— sect. Leucoglossa § 1.
Microsperma Sch. Bip. .........+.-+ 201
— sect. Leucogymnocline Sch. Bip.
so cael RNs Rte attetoton aaettat tis acetate dete 193

— sect. Pyrethrum subsect.
Cinerariifolia Heyw. ....,......0+ 213
— — subsect. Eupyrethrum Briq. .232
— — subsect. Leucanthemopsis
GITOUK 5) 25.2 svcteebtes sts zecetrttsersneetetoees De
— — subsect. Parthenium Briq. ...201
— — subsect. Pyrethrum (Zinn.)

ROY Wierensdvenccssctosnacsxpalbsutbseserethauaiens 232
— sect. Xanthogymnocline Sch. Bip.

savase dace tecvents &cbeha sees aeons 330
— abrotanifolium (L.) Dreuce. ...... 329
— achilleifolium (M.B.) Sch. Bip. 351
— Akinfievii (Alex.) Tzvel. ........... 357
== alatavicumuerGy-...f--n2:1-sceuvaccnese 249
=) alaunicumuKs-Poly.nts.cstepeccec 377
— alpinum (L.) Sch. Bip. ............++ 228
— angulatum Willd. .........ceeee 90
— Aphanassievii Krasch. ...........++ 408
— arcticum (L.) Sch. Bip. ............. 386
— argenteum (Lam.) Willd. ........... 361
=='— var. glabratum DC, nuns. 361
— Aucherianum (DC.) Sch. Bip. ..240
Ss halsamita Lis Arica seen 198
— balsamitoides Sch. Bip.............. 197
— Barclayanum DC. .......ccsssreensees 341
— bipinnatum (L.) Sch. Bip. ......... 324
==tboreale Fisch. exsD@tencee 327
— Botschantzevii Koval. .............+ 309
— brachanthemoides (Winkl.) Krasch.

Be Mis Meteo, EGov ca scasteweomaateonane 424
— budjnurdense (Rech. f.) Tzvel. 5325
2+Vcanescens: D@ixtss PAU 336
— — ssp. erivanense Tzvel. .......... 337
wet NCANESCENS) DC. \ig kbs cpeike aleadetccnere $24
— — var. parvifolium DC. .. 337, 879
— Capusii Franch. ...........-1eeceeee 413
— carneum (M.B.) Sch. Bip.......... 217
-—— caucasicum Sch. Bip. ..........00 161

890

— chiliophyllum (F. and M.) Sch.

BUDE sccacasnsnscootsisuaeseakteee CaaS 352
— cinerariifolium (Trev.) Sch. Bip.
sasseay-ecceiuavsiiinedicasnavesenGee <eee eaMean ee 213
— corymbosum (L.) Sch. Bip. ....... 232
— — ssp. subcorymbosum (Schur) B.
Pav, .....<cssndhcpendtes hae aticrtbelos. osteo 234
— crassipes (Stschegl.) Tzvel. ....... 338
— darvasicum Winkl.................000+ 415
— discoideum Rchb. f. ..................- 313
— duderanum (Boiss.) Tzvel. .......: 332
— dumosum Boiss. ...........:::cccseceeees 356
— fastigiatum (Winkl.) Krasch. ....405
— ferganense Koval. .........:cceceee 304
— Fischerae Aitch. and Hemsl. ....287
— flavovirens (Boiss.) Tzvel. ........ 334
— fruticulosum Ldb. .................0.... 406
— Galae (M. Pop.) Nevski. ........... 242
— Gmelinii Sch. Bip. ............ 377, 386
— gracile Hook. f. and Thoms. ....407
— Grigorjewii Krassn. ............0+ 409
— Heldreichianum Fenzl. ex Tchih.
jataas Minin sales Dtagstuns bea canscy saves Oe 231
— Herderi Rgl. and Schmath. ....... 414
— heterophyllum Boiss. ................-. 333
— INCANUM AUCK. .....cccccceecesseceeseeeees 348
— — F. and M. ex DC. ............... 336
Se Linc MOT bs Sates). Sees $24
— indicum (L.) Sch. Bip. ............. 371
— integrifolium (Richards.) Sch. Bip.
wcctasel salaaspeetatranctt htt chock naa 388
— Johnstonii Hemsl. ...............0.0000 316
— karataviense Koval. ..........0.0000 306
—=iKarelingl TP2Vele22.,.. nA). seeeek 344
— khorassanicum (Krasch.) Tzvel. 358
— Kittaryanum (C.A.M.) Tzvel.....345
— Kochii Sch. Bip. ........c eee 352
— kokanicum Krasch. ............:0:0-00+ 408
— Komarovii Krasch. ...............06 424
— Kotschyi Boiss. (.......ccccceeeeecees 333
— Kotzebuense (Bess.) Bess.......... 324
— Krascheninnikovii NevskKi. ........ 309
— Kuschakewiczii O. Fedtsch. ...... 286
— lanuginosum Sch. Bip. and Herd.
nA EPA i RN PN dO Ort a ad te py!
— Ledebourii Sch. Bip. .............. Bis
— leptophyllum (Stev.) Sch. Bip. .211
— leucanthemum (L.) Sch. Bip..... 143
— leucophyllum Rg. ......ccccceeeee 414
— longifolium Wall. .....ccccccceeeee 412
— longipedunculatum (Sosn.) Tzvel.
sswonivicnad evedvagdie teens dh hdc eee ee OS 354
— macrophyllum (W. and K.) Sch.
1S FA PRs Sar baht aS wo sor om 194

— Meyerianum Sch. Bip. ..........0 337
— millefoliatum F. and M. ............ 329
— millefolium (L.) Tzvel................ 348
— morifolium (Ramat.) Kitam....... 373
— mucronatum (Rgl. and Schmalh.)
Mis Pool. .ciieinich-.oae aknoune 302
— — var. corymbulosum Winkl. ..303
— myriophyllum Willd..............0... Ja
— — ssp. borshomicum Tzvel. .....332
== \NanumM, alict,,... sade. distiae sche 410
3 — Clarke. i... .0c...:tbatee- een 412
— Newesskyanum Winkl................. 328
— niveum (Lag.) Sch. Bip. ............ 209
— odessanum (Klok.) Tzvel. .......... 348
— oligocephalum (DC.) Sch. Bip. 354
— orientale Willd. ..........cccccceceeeeee 524
— Paczoskii (Zefir.) Tzvel. .......... 349
— Pallasianum (Fisch. ex Bess.)
Trautv.;and Meyer. jaa itemn yas 402
— palustre (Willd.) Sch. Bip. ....... 215
— pamiricum (Hoffm.) Bornm. .....286
— pamiroalaicum Koval. ............+ 307
— paropamisicum (Krasch.) Koval.....
aasedsassagavestatshaden SAV EER Cia neve ed 310
— parihemifolium (Willd.) Sch. Bip.
ns agncBidp Boake sted sUaT eT aR ah OR he aE 203
— parthenium (L.) Sch. Bip. ......... 204
— Pjataevae Koval. ...........ceceeeee 306
— pseudoachillea Winkl. ................ 328
— parthenifolium (Willd.) Sch. Bip...
asenonqaucoenclacasiubeaneleveredaetcs Gatere akan 203
— pulchrum (Ldb.) Sch. Bip......... 253
SSPRESENI. OVA, .ccc,cassecesscesesuesearesees 304
— roseum (Adam) Sch. Bip. 217, 219
— rupestre M. Pop. ex Nevski. ....287
— santolina Winkl. .......... eee 343
— saxatile (Kar. and Kir.) Sch. Bip.
ask tyoasocassvenoisavae eet: TRIE eran 344
— saxicolum (Krasch.) Tzvel......... 342
— Scharnhorstii Rgl. and Schmalh. ...
sc visistvuceapais eaasenben Sulsssaecaer ie Ee 409
— schugnanicum Winkl. .........006 416
— sclerophyllum (Krasch.) Tzvel. .347
— scopulorum (Krasch.) Tzvel. .....342
— Semenovii Herd. ...........:ccscceseeeeee 243
— senecionis (Jacq. ex Bess.) J. Gay.
safevWnssenenen Ghee acta eee 412, 414
— serotinum (L.) Sch. Bip. ........... 139
— sericeum (Adam) Sch. Bip........ 237
— setaceum Tausch. ..........ccccccseees 329
— — (Rgl. and Schmalh.) Oliv. ...304
si DIFICUnt, Liweg ie. ayseus ection 417
— silaifolium (Stev.) Sch. Bip. .....220

— sinense (Sabine) Sch. Bip.........373

937

— sinuatum (Ldb.) Sch. Bip.......... 370
— suaveolens Hook. ..........:.cc000c00008 150
— subalpinum (Schur) Simonk. .... 145
— subcorymbosum (Schur) Simonk. ...

— submarginatum Koval. ............... 305
— subsimile (Rech. f.) Koval. .......302
— Szowitsii (C. Koch) Sosn. and

MARR cisco teen ees cassecouaseeees 329
— tabrisianum (Boiss.) Sosn. and

Takhic shew ee, veneer, 555
— tadshikorum Kudr. ...............0:0++. 299
— tamrutense (Sosn.) Sosn. ........... 334
— tanacetoides (DC.) Tzvel. .......... 337
— tauricum Sch. Bip. ...............+ 348
— tenuifolium auct. «0.00... 410
= OAC. ceah caterer ettescerttecsestere =e: 411

— tenuissimum (Trautv.) Grossh. .355
— tibeticum Hook. f. and Thoms. 410

——"fOINENIOSUMN™ AUCU. 1... cntieuscctestserssns 414
3) 0] CR CE RS ee ee 416
— Tournefortii Sch. Bip................. 240
— transiliense Herd. ..............0000008 258
— trichophyllum Rgl. and Schmalh.
CRG LETS ck 363
— tripinnatifidum Oliv. .......0.0000. 303
— turkestanicum (Rgl. and Schmalh.)
Polyjakscott..J208.. A2tih 3. RRRaRe 316
— turcomanicum (Krasch.) Tzvel. 358
— turlanicum (Pavl.) Tzvel. ........... 341
— ulutavicum Tzvel............... 350, 878
— umbellatum Gilib. .................00. 326
— umbelliferum Boiss. ..............:0++ 363

— uniflorum (F. and M.) Sch. Bip.
356

— uralense (Krasch.) Tzvel............ 346
— urgutense (M. Pop.) Koval. ......308
== VU GATE: ACU vsccscc<ccsécsescoscetoescoceses 327
= Villigp are ts.) sc.ccenrstesesst arses.ceesenreress 326
— — var. boreale (Fisch. ex DC.)
Mrauty.. and Mey:.-...2:..¢-.t<00ee 327
— vulgare var. genuinum Trautv..326
— Waldsteinii Sch. Bip. ..........0.... 142
— Walteri (Winkl.) Tzvel............... 359
— Weyrichii (Maxim.) Kitam. ....... 383
— xylorrhizum Krasch. ex Stanjuk.
Be ead soda deed Ree tes Se Cm 410
— Zawadskii (Herb.) B. Pavl. ....... 377
Tephroseris (Rchb.) DC., sect. ....... 753
— aurantiaca Griseb. and Schenk.
dears esis Seventeen ae een eee eee 768
= CAMPESITIS: IRCHDs.<..<.5-ccvorccsc0ss0e3s f/e%s:
— cladobotrys Criseb. and Schenk. ...
Sages gute seseeseist esses ee oet Panesar a eee 759

891

==! Crispa "REND: ii 2..1. tae 765
—' papposa ‘Schur... aun keane 761
Trichanthemis sect. ...........0..000 278
Trichanthemis Rgl. and Schmalh.

PUT. Lok, AK A 27331877

— sect. Eutrichanthemis Lipsch. ...278
— — subsect. Leianthus Novopokr.

and SidOficcz. ete ae 280
— aulieatensis (B. Fedtsch.) Krasch. .

saveaeneciwwen TRE A, Sevetth st REE 277
==) aurea’ Kiraschyn.)..2. 0 ee 281
— Butkovii Koval. ..............ccceeeeeee 282
— glabriflora Novopokr. and Sidor. ..

DONGITL LANL. SEO ARRAY 281
— karataviensis Rgl. and Schmalh.

tS eh A. FE 278
— — ssp. dshungarica Rubtz. ....... 279
— Litwinowii (Krasch.) Tzvel. ...... 283
— paradoxos (Winkl.) Tzvel. .........281
== radiata. aucth 2.1. RARER 281
— radiata Krasch. and Vved.......... 280
Trichanthemopsis Tzvel., sect.

hiv evesamimteriirencternaterte eae 244, 872
Tridactylina (DC.) Sch. Bip. ........ 389
— Kirilowii (Turcz.) Sch. Bip....... 390
Tridactylina Turczaninovii Krasch.378
Tripleurospermum Sch. Bip. ........ 157
Tripleurospermum subgen. .............. 168
SUSCC be scsascascacaccesstvaessnsece. ahs ME 175
— ambiguum (Ldb.) Fr. and Sav. . 168
— ambiguum Fr. and Sav............... 170
— aromaticum Rupr. ex Boiss. .....222
— breviradiatum (Ldb.) Pobed. ..... 178
— caucasicum (Willd.) Hayek. ...... 161
— colchicum (Mand.) Pobed. ........ 174
— daghestanicum Rupr. ex Boiss.

EERE Ses tor maintey ccoeaece 222

— decipiens (Fisch. and Mey.)

BBV OURIMI: ccs sos. secestbcdaeste acces cbeeneeeouanes 182
— disciforme (C.A.M.) Sch. Bip. .181
— elongatum (F. and M.) Bornm. 173
— Grossheimii (An. Fed.) Pobed. . 165

— inodorum (L.) Sch. Bip. ............ 175
— inodorum maritimum Sch. Bip. 177
— — var. ambiguum Rchb. ........... 170
— — B. limosum Rl. ........0cceee 177
— Karjaginii (Mand. and Sof.)

RODEO koe asivescitesnacasszatenens seen et eae 167
— lamellatum (Bge.) K. Rech....... 155
— limosum (Maxim.) Pobed. ......... 5/7
— maritimum (L.) Koch. ................ WOGATE
— melanophyllum (Nabel.) Pobed.

Se ORM EN ae MON RP nee en 161, 165

— nivale Rupr. ex Boiss. ...........0+ 223

892

— parviflorum (Willd.) Pobed. ...... 183
— phaeocephalum (Rupr.) Pobed..171
— praecox (M.B.) Bornm. ............. 183
— pulchrum (Ldb.) Rupr. ..........0+ 253
— rupestre (Somm. and Ley.) Pobed.
stscibcnzpsvvadeadnaneswaditeeeisiztae ce bee 173
— sevanense (Mand.) Pobed. ......... 182
— subnivale Pobed. ................:000+ 162
— subpolare Pobed. .........::cc:cccecesee 172
— Szowitzii (DC.) Pobed. .............. 166

— Tchichatchevii (Boiss.) Bornm. 174
— tetragonospermum (F. Schmidt)

Pobe Giges: s5hcesteg teem 170
— transcaucasicum (Mand.) Pobed.

Sansdatl.. past shane ine: 167
Tripterospermum caucasicum’ Owerin.

sachevannthethixelhe athe ners aumeelivdes . steers avs 161
Turaniphytum Poljak. .......... 632, 880
— eranthemum (Bge.) Poljak......... 635
— kopetdaghense Poljak. ...... 636, 881
Tpssila go Mess Ak. Gucet ceuccntatok:- Aeageee 645
Tussilago Ly... tii: aiccnaieeiseesh 641
SADA snpncasbxcssacsieianeasistbaasasver: costae 644
= alpina Ly... dicd..as:tet aes 655
— corymbosa R. BI. .......sccesseeeereees 649
== farfara yi arcg scented cenieeee 641
a= frigida: Ls) .tiad..uinaraniemaiesie 649
== iGmelini Tted: ic... caacionrencceeso> 653
Sa VIE Lo innnssqpecssunnsksvsneyencx ee! 643
— laevigata Willd. .........:ccccceeeee 650
— palmata Ait. ...cccccccccsesesseseseeeees 651
com (petasites MB. viis5:..vevissstsaseecaree? 643
== ISPUFTUSHREtZ.025...2)..szessusdenesecnasecnse 643
— tomentosa Ehth. .........:cceceeeee 643

938

Tussilagineae Benth., subttr............. 640
Weamial Pave iii pics een! 271
— angrenica (Krasch.) Tzvel. ........ 272
Ugamia trichanthemoides Pavl. .....272
Velutini’ Boiss: icici 720

Waldheimia Kar. and Kir. ... 263, 875

Waldheimia, SECt. .......-:-:-.:.ceccnashenestl 269
— glabra (Decne.) Ral. ........:cccse0e0 270
— Huegelii (Sch. Bip.) Tzvel. ....... 271
— Korolkowii Rgl. and Schmalh. .268
— Stoliczkae (Clarke) Ostenf. ....... 268
— Stracheyana Rg. oo... 271
— tomentosa (Decne.) Rel. ............ 267
— transalaica Tzvel. .............. 269, 875
— tridactylites auct. ........:cecceseeeees 270
— tridactylites Kar. and Kir. ......... 269

Xanthoglossa (DC.) Sch. Bip., sect.
diserbeadsseunsis odpasaancdhtcetcess CIRO SES TMD 330

Xanthophtalmum Sch. Bip. ... 131, 136

Xanthophtalmum (Sch. Bip.) Tzvel.

SECEs).2....200 nathan ayer 136
Xylanthemum Tzvel. ............. 284, 877
— Fischerae (Aitch. and Hemsl.)

TZveléh..jaeare tes esa 287
— pamiricum (Hoffm.) Tzvel. .......286
— polycladum (Rech. f.) Tzvel. ....288
— rupestre (M. Pop. ex Nevski)

Tzvel;)..0.-- 22 eine 287

Xylopyrethrum Tzvel., sect. .. 229, 872

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