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Akademiya Nauk SSSR 



FLORA of the U.S. S.R. 

Volume XX 
6.K. Shishkin and S.V.Yuzepchuk, Editors 

Lahiatae 



TRANSLATED FROM RUSSIAN 



62 



BOTANICHESKII INSTITUT IM. V.L. KOMAROVA AKADEMII NAUK SSSR 

Botanical Institute of the Academy of Sciences of the USSR 

Akaje^il^ Vaule SSS^, Bda>]i cli egjeit l^'^T/fl 

FLORA of the U.S. S.R. 



(Flora SSSR) 
(Series initiated by V. L. Komarov) 

Volume XX 

Lahiatac 



Chief Editor B.K. Shishkin 
Volume Editors B.K. Shishkin and S.V. Yuzepchuk 



Compiled by 

A.G. Borisova, I.T. Vasil'chenko, E.V. Volkova, 

S.G. Gorshkova, O.E. Knorring, L. A. Kupriyanova, 

E.G. Levin, V.V. Pis'yaukova , A.I. Poyarkova, 

B.K. Shishkin, and S.V. Yuzepchuk 



Izdatel'stvo Akademii Nauk SSSR 

Moskva -Leningrad 

1954 



JAN18W// 



^^ 



Translated from Russian 



Israel Program for Scientific Translations 
Jerusalem 1976 



TT 75-50001 

The Publication of this Translation 
was supported by 
THE UNITED STATES -ISRAEL 
BINATIONAL SCIENCE FOUNDATION 



Copyright© 1976 

Keter Publishing House Jerusalem Ltd. 

Cat. No. 61209 7 

ISBNO 7065 1560 9 



Translated by IPST Staff 
Edited by Dr. N. Landau 



Printed and bound by Keterpress Enterprises 
Jerusalem, Israel 



Available from the 

U. S. DEPARTMENT OF COMMERCE 

National Technical Information Service 

Springfield, Va. 22151 



TABLE OF CONTENTS 



Systematic Index of Species in Volume XX 

Preface 

Family CXL. Labiatae Juss 

Key to Genera 

Subfamily 1 . Ajugoideae Benth 

Tribe 1 . Ajugeae Benth 

Genus 1240. Ajuga L 

Genus 1241. Teucrium L 

Genus 1242. Amethystea L 

Tribe 2. Rosmarineae Briq 

Genus 1243. Rosmarinus L 

Subfamily 2. Scutellarioideae Briq 

Genus 1244. Scutellaria L 

Subgenus 1. Euscutellaria Briq 

Subgenus 2. Cystaspis Juz 

Subgenus 3. Anaspis Juz 

Subgenus 4. Apeltanthus Nevski 

Subfamily 3. Lavanduloideae Briq 

Genus 1245. Lavandula L 

Subfamily 4. Drepanocaryoideae Pojark 

Genus 1246. Drepanocaryum Pojark 

Subfamily 5. Stachydoideae Briq 

Tribe 1 . Marrubieae Endl 

Genus 1247. Thuspeinantha Durand 

Genus 1248. Marrubium L 

Genus 1249. Lagopsis Bge 

Genus 1250. Sideritis L 

Tribe 2. Nepeteae Benth 

Genus 1251. Agastache Clayt. ex Gronov 

Genus 1252. Lophanthus Adans 

Genus 1253. Schizonepeta Briq 

Genus 1254. Nepeta L 

Genus 1255. Glechoma L 

Genus 1256. Dracocephalum L 

Subgenus 1. Eudracocephalum Briq 

Subgenus 2. Ruyschiana (Mill.) Benth 

[The page numbers of the Russian original appear in the left-hand margin of the 



Russian 


English 


page 


page 




vii 




1 


1 


3 


6 


6 


16 


12 


1^ 


12 


17 


13 


39 


27 


69 


47 


70 


48 


70 


48 


71 


49 


72 


50 


74 


51 


199 


133 


207 


138 


217 


144 


225 


150 


226 


150 


227 


151 


228 


151 


231 


154 


232 


154 


232 


154 


233 


155 


248 


165 


253 


168 


273 


181 


273 


181 


275 


183 


282 


188 


286 


191 


437 


293 


439 


295 


444 


298 


471 


316 



text. 



Ul 



Subgenus 3. Fedczenkiella (Gontsch.) Schischk. . 

Genus 1257. Kudrjaschevia Pojark 

Genus 1258. Lallemantia Fisch. et Mey 

Genus 1259. Hymenocrater Fisch. et Mey 

Genus 1260. Hypogomphia Bge 

Tribe 3. Stachydeae Briq 

Subtribe 1 . Brunellinae Briq 

Genus 1261. Prunella L 

Subtribe 2. Melittinae Briq 

Genus 1262. Melittis L 

Subtribe 3. Lamiineae Briq 

Genus 1263. Pseuderemostachys M. Pop 

Genus 1264. Neustruevia Juz 

Addenda XIX (Diagnoses of New Species Mentioned in Volume XX) 

Supplement 

Index Alphabeticus 

Vegetarion Regions of the USSR 

List of Abbreviations 



473 


318 


474 


319 


482 


324 


488 


328 


491 


330 


493 


332 


493 


332 


494 


332 


498 


335 


498 


336 


500 


337 


500 


337 


501 


338 


505 


339 


528 


363 




365 




381 




384 



IV 



SUBJECTS AND CONTRIBUTORS 

Characteristics of the Labiatae and key for 

the genera 
Genera: Teucrium, Scutellaria, 

Sideritis, Meehania, Neustruevia Arranged by S.V. Yuzepchuk 

Genus Ajuga Arranged by V.V. Pis'yaukova 

Genus Amethystea Arranged by E.V. Volkova 

Genera: Rosmarinus, Lavandula, 

Dracocephalum Arranged by B.K. Shishkin 

Genera: Drepanocaryum, Agastache, 

Schizonepeta, Nepeta, Kudrjaschevia, 

Melittis Arranged by A.I. Poyarkova 

Genera: Thuspeinantha, Prunella Arranged by A.G. Borisova 

Genera: Marrubium, Lagopsis, 

Pseuderemostachys Arranged by O.E. Knorring 

Genus Lophanthus Arranged by E.G. Levin 

Genus Glechoma Arranged by L.A. Kupriyanova 

Genera: Lallemantia, Hymenocrater Arranged by S.G. Gorshkova 

Genus Hypogomphia Arranged by I.T. Vasil'chenko 

Addenda XIX - Diagnoses plantarum novarum in 
tomo XX Florae URSS commemoratarum. 



The plates were drawn by the following artists: N. A. Moiseeva - I, II, V-VIII, X, XI, 
XVI; Z.V. Kobyletskaya - III, IV, IX, XII, XIII, XV, XVII-XIX, XXV, XXVI; M.R. 
Gabe - XIV, XX-XXIV, XXVII. 



SYSTEMATIC INDEX OF SPECIES IN VOLUME XX* 
Family CXI. Labiatae Juss. 
Subfamily 1. Ajug^oideae Benth. 
Tribe 1. Ajugeae Benth. 
Genus 1240. Ajuga L. 

Section 1. Chamaepitys Benth. 

Russian 

page** 

11608. 1. A. chia Schreb 20 

2. A. pseudochia Shost 21 

11610. 3. A. chamaepitys (L.) Schreb 25 

4. A. salicifolia (L.) Schreb 26 

5. A. oblongata M. B 26 

6. A. chamaecistus Ging 27 

7. A. turkestanica (Rgl.) Briq 28 

8. A. Laxmannii (L.) Benth 29 

Section 2. B u g u 1 a Benth. 

9. A. reptans L 30 

10. A. shikotanensis Miyabe et Tatewaki 32 

11. A. genevensis L 32 

12. A. multiflora Bge 36 

11620. 13. A. pyramidalis L 37 

14. .A. orientalis L 38 

Genus 1241 . T e ucr i urn L. 
Section 1. Teucris Benth. 

1. T. orientale L 42 

2. T. Taylori Boiss. . „ 43 

3. T. parviflorum Schreb 44 

Section 2. Stachyobotrys Benth. 

4. T. hircanicum L. , 45 

4a. T. ussuriense Kom 528 

* [This index has been reproduced photographically from the Russian original.) 
** [Russian page numbers appear in the left-hand margin of the text.) 



vu 



Section 3. Scordium Rchb. 

5. T. scordium L 46 

6. T. scordioides Schreb 60 

7. T. botrys L 60 

Section 4. Chamaedrys Schreb. 

11630. 8. T. nuchense C. Koch 52 

9. T. multinodum (Bordz.) Juz 53 

10. T. trapezunticum (Reching. fil.) Juz 54 

11. T. chamaedrys L 55 

12. T. pulchrius Juz 57 

13. T. Fischeri Juz 57 

14. T. syspirense C. Koch 58 

15. T. krymense Juz 61 

16. T. excelsum Juz 62 

17. T. canum Fisch. et Mey 63 

Section 5. Polium Schreb. 

11640. 18. T. jailae Juz 65 

19. T. praemontanum Kick 66 

20. T. pannonicum Kern 67 

21. T. polium L 68 

Genus 1242, Amet hg ste a L. 

1. A. coerulea L 70 

Tribe 2. Rosmarineae Briq. 
Genus 1243. Rosmarinus L. 

1. R. officinalis L 71 

Subfamily 2. Scutellarioideae Briq. 
Genus 1244. Scutellaria L. 
Subgenus 1. Euscutellaria Briq. 
Section 1. G a 1 e r i c u 1 a r i a A. Hamilt. 

1. S. galericulata L 90 

2. S, Krasevii Kom. et Schischk 92 

3. S. yezoensis Kudo 95 

4. S. Regeliana Nakai 95 

11650. 5. S. Ikonnikovii Juz 96 

6. S. hastifolia L 97 

7. S. dubia Tal. et Schir 97 

8. S. dependens Maxim 98 

9. S. scordiifolia Fisch 99 

10. S. Taquetii Levi, et Vaniot 100 

11. S. tuminensis Nakai 101 

12. S. polyphylla Juz 102 

13. S. moniliorrhiza Kom 102 



VIU 



Section 2. Stachymacris A. Hamilt. 

14. S . baicalensis Georgi 103 

11660 15. S. altissima L 105 

16. S. Tournefortii Benth 106 

17. S. ussuriensis (Rgl.) Kudo 106 

18. S. pacifica Juz 107 

19. S. shikokiana Mak 108 

20. S. albida L Ill 

21. S. pallida M. B Ill 

22. S. Woronowii Juz 112 

23. S. pontica C. Koch 113 

24. S. Helenae Alb 114 

Section 3. Nevskianthe Ju7. 

11670. 25. S. colpodea Nevski 115 

26. S. striatella Gontsch 116 

27. S. cristata M. Pop 116 

Section 4. L u p u 1 i n a r i a A. Hamilt. 

28. S. navicularis Juz 118 

29. S. chenopodiifolia Juz 119 

30. S. Juzepczukii Gontsch 119 

31. S. Litwinowii Bornm. et Sint 120 

32. S. luteo-coerulea Bornm 121 

33. S. ramosissima M. Pop 122 

34. S. intermedia M. Pop 123 

11680. 35. S. glabrata Vved 124 

36. S. squarrosa Nevski 124 

37. S. araxensis Grossh 128 

38. S. taurica Juz 129 

39. S. Stevenii Juz '. 130 

40. S. Tatianae Juz 130 

41. S. Raddeana Juz 130 

42. S. leptostegia Juz 132 

43. S. grandiflora Sims 132 

44. S. tuvensis Juz 133 

11690. 45. S. turgaica Juz 133 

46. S. karkaralensis Juz 134 

47. S. nepetoides M. Pop 135 

48. S. picta Juz 136 

49. S. flabellulata Juz 136 

50. S. comosa Juz 137 

51. S. oxystegia Juz 138 

52. S. microdasys Juz 139 

53. S. Androssovii Juz 139 

54. S. mesostegia Juz 141 

11700. 55. S. Kurssanovii Pavl 141 

56. S. Titovii Juz 142 

57. S. Catharinae Juz 145 



IX 



58. S. transiliensis Juz 146 

59. S. Albert! Juz 147 

60. S. Przewalskii Juz 148 

61. S. soongorica Juz 148 

62. S. Krylovii Juz 149 

63. S. Sieversii Bge ' 150 

64. S. hypopolia Juz • . . . 151 

11710. 65. S. heterochroa Juz 152 

66. S. hirtella Juz 152 

67. S. novorossica Juz • 153 

68. S. polyodon Juz 153 

69. S. oreophila Grossh 154 

70. S. oschtenica Juz 155 

71. S. daghestanica Grossh 155 

72. S. granulosa Juz 156 

73. S. orientalis L 157 

74. S. Sedelmeyerae Juz 158 

11720. 75. S. platystegia Juz ■ 161 

76. S. Grossheimiana Juz 162 

77. S. artvinensis Grossh 162 

78. S. Karjaginii Grossh 163 

79. S. rhomboidalis Grossh 164 

80. S. Prilipkoana Grossh 164 

81. S. darriensis Grossh 165 

82. S. Sosnovskyi Takht 165 

83. S. amphichlora Juz 166 

84. S. araratica Grossh 167 

11730. 85. S. adenostegia Briq 168 

86. S. bucharica Juz 169 

87. S. Gontscharovii Juz 169 

88. S. haematochlora Juz 170 

89. S. tschimganica Juz 171 

90. S. urticifolia Juz 171 

91. S. Iskanderi Juz 172 

92. S. pycnoclada Juz 172 

93. S. angrenica Juz 175 

94. S. cordifrons Juz 175 

11740. 95. S. adsurgens M. Pop 176 

96. S. phyllostachya Juz 177 

97. S. sevanensis Sosn 178 

98. S. verna Bess 179 

99. S. creticola Juz 180 

100. S. Chitrovoi Juz 181 

101. S. cisvolgensis Juz 181 

102. S. oxyphylla Juz 182 

103. S. supina L 183 

104. S. altaica Fisch 184 

11750. 105. S. mongolica Sobolevsk. 185 

106. S. irregularis Juz 185 

107. S. Knorringiae Juz 186 

108. S. xanthosiphon Juz 186 



109. S. oligodonta Juz 187 

110. S. Paulsenii Briq 188 

111. S. filicaulis Rgl . 189 

112. S. kugarti Juz 190 

113. S. Popovii Vved 190 

114. S. talassica Juz 193 

11760. 115. S. Linczevskii Juz 194 

116. S. subcordata Juz 194 

117. S. subcaespitosa Pavl 195 

118. S. lanipes Juz 196 

119. S. toguztoraviensis Juz 196 

120. S. Alexeenkoi Juz 197 

121. S. ocellata Juz 198 

122. S. karatavica Juz 199 

Subgenus 2. Cysfaspls Juz. 

123. S. physocalyx Rgl. et Schmalh 201 

124. S. Anitae Juz 201 

11770. 125. S. pamirica Juz 202 

126. S. microphysa Juz 205 

127. S. polytricha Juz 205 

128. S. jodudiana B. Fedtsch 206 

129. S. darvasica Juz 207 

130. S. schachristanica Juz 208 

Subgenus 3. Anaspis (Reching. fil.) Jur. 

131. S. Fedtschenkoi Bornm 209 

132. S. hissarica B. Fedtsch 210 

133. S. velutina Juz. et Vved 211 

134. S. Nevskii Juz. et Vved 211 

11780. 135. S. heterotricha Juz. et Vved 212 

136. S. schugnanica B. Fedtsch ^ . . 212 

137. S. Lipskyi Juz \ 213 

138. S. villosissima Gontsch 214 

139. S. holosericea Gontsch . . . . 217 

Subgenus 4. Apeltanthus Nevski 

140. S. andrachnoides Vved 219 

141. S. baldshuanica Nevski 220 

142. S. poecilantha Nevski 221 

143. S. megalodonta Juz 221 

144. S. orbicularis Bge 222 

11790. 145. S. immaculata Nevski 223 

146. S. guttata Nevski 223 

147. S. rubromaculata Juz 224 

148. S. leptosiphon Nevski 224 

Subfamily 3. Lavanduloideae Briq. 
Genus 1245. Lavandula L. 

1. L. spica L 225 



XI 



Subfamily 4. Drepanocaryoideae Pojark. 

Genus 1246. Dr e p a no c a r y u m Pojark. 
1. D. Sewerzowii (Rgl.) Pojark 



228 



11800. 



11810. 



11820. 



Subfamily 5. Stachydoideae Briq. 
Tribe 1. Marrubieae Endl. 
Genus 1247. Tkuspeinantha Durand 
1. Th. persica (Boiss.) Briq 



232 



Genus 1248. M a r r u h i um L. 
Section 1. Marrubium Benth. 

1. M. vulgare L 235 

2. M. alternidens Rech. fil 236 

3. M. Woronowii N. Pop 239 

4. M. praecox Janka 239 

5. M. parviflorum Fisch. et May 240 

6. M. plumosum C. A. M 241 

7. M. catariifolium Desr 242 

8. M. leonuroides Desr > 242 

9. M. nanum Knorr 243 

10. M. propinquum Fisch. et May 244 

11. M. persicum C. A. M 244 

12. M. peregrinum L 245 

13. M. goktschaicum N. Pop 246 

14. M. purpuraum Bge 246 

15. M. Turkaviczii Knorr 247 



1. L. 

2. L. 

3. L. 

4. L. 



Genus 1249. Lag op sis Bge. 

marrubiastrum (Staph.) Ik. -Gal 

flava Kar. et Kir 

eriostachya (Benth.) Ik. -Gal 

supina (Steph.) Ik. -Gal 



Genus 1250. Sideritis L. 
Section 1. Empedociea (Rafin.) Benth. 
taurica Steph 



1. s 

2. S. chlorostegia Juz. . 

3. S. catillaris Juz. . . 

4. S. Marschalliana Juz 

5. S. imbrex Juz. . . . 

6. S. conferta Juz. . . 

7. S. euxina Juz. . . . 



Section 2. H e s i o d i a (Moench) Benth. 



8. S. montana L 

9. S. comosa (Roche!) Stank. 
10. S. Balansae Boiss 



248 
249 
250 

250 



256 
260 
261 
263 
264 
266 
266 



268 
271 
272 



xu 



Tribe 2. Nepeteae Benth, 
Genus 1251. Agastache Clayt. 

1. A. rugosa (Fisch. et Mey.) O. Ktze 274 

Genus 1252. Lopha nthus Adans. 

1. L. chinensis (Rafin.) Benth 276 

2. L. Krylovii Lipsky 277 

3. L. Schrenkii Levin 278 

11830. 4. L. tschimganicus Lipsky 279 

5. L. Schtsohurowskianus (Rgl.) Lipsky 279 

6. L. Lipskyanus Ik. -Gal 280 

7. L. subnivalis Lipsky 281 

8. L. elegans (Lipsky) Levin 282 

Genus 1252a. Meeha ni a Britt. 

1. M. urticifolia (Miq.) Makino 529 

Genus 1253. S chi z o ne p et a Briq. 

1. S. multifida (L.) Briq 283 

2. S. annua (PalL) B. Schischk 285 

Genus 1254. Nepeta L. 
Section 1. Glechomanthe Pojark. 

1. N. longibracteata Benth 303 

2. N. Lipskyi Kudr 305 

11840. 3. N. glutinosa Benth 306 

4. N. Knorringiana Pojark 310 

5. N. subhastata Rgl 311 

Section 2. Spicatae (Benth.) Pojark. 

6. N. podostachys Benth 312 

7. N. densiflora Kar. et Kir 314 

8. N. transiliensis Pojark 315 

9. N. kokamirica Rgl 316 

10. N. Mariae RgL 317 

11. N. pulchella Pojark 318 

12. N. tytthantha Pojark 320 

11850. 13. N. odorifera Lipsky 321 

14. N. bucharica Lipsky 322 

15. N. maussarifii Lipsky 323 

16. N. alatavica Lipsky 324 

17. N. schugnanica Lipsky 325 

18. N. supina Stev 326 

19. N. Buschii Sosn. et Mand 327 

20. N. kokanica Rgl 328 

21. N. pseudokokanica Pojark 330 

22. N. pamirensis Franch 333 



XIU 



Section 3. Capituliferae (Benth.) Pojark. 

11860. 23. N. vakhanica Pojark 335 

24. N. pseudofloccosa Pojark 336 

25. N. badachschanica Kudr 337 

26. N. ladanolens Lipskyi 338 

27. N. Olgae Rgl 339 

Section 4. Macronepeta Benth. 

28. N. manchuriensis S. Moore 341 

29. N. sibirica L 342 

30. N. formosa Kudr 344 

31. N. Gontscharovii Kudr 346 

32. N. consanguinea Pojark 347 

Section 5. Cat aria Benth. 

11870. 33. N. cataria L 349 

34. N. cyanea Stev 353 

35. N. kubanica Pojark 355 

36. N. czegemensis Pojark 356 

37. N. Biebersteiniana (Trautv.) Pojark 357 

38. N. grandiflora M. B 359 

39. N. betonicifolia C.A.M 361 

40. N. somkhetica Kapell 362 

41. N. Grossheimii Pojark 363 

42. N. strictifolia Pojark 365 

11880. 43. N. alaghezi Pojark 366 

44. N. Buhsei Pojark 367 

45. N. Mussinii Sprang 369 

46. N. Reichebachiana Fisch. et Mey 371 

47. N. hajastana Grossh 373 

48. N. transcaucasica Grossh 374 

49. N. noraschenica Grossh 378 

Section 6. Micranthae Boiss. 

50. N. amoena Stapf 380 

51. N. Troitzkii Sosn 382 

52. N. micrantha Bge 383 

11890. 53. N. Meyeri Benth 384 

54. N. pallida C. Koch 386 

55. N. saccharata Bge 388 

56. N. ispahanica Boiss 389 

57. N. santoana M. Pop 390 

Section 7. Schizocalyx Pojark. 

58. N. longituba Pojark 392 

59. N. Sosnovskyi Asker 393 

60. N. fissa C. A. M 395 

61. N. Trautvetteri Boiss. et Buhse 3% 



XIV 



62. N. velutina Pojark 397 

11900. 63. N. daghestanica Pojark 399 

64. N. iberica Pojark 400 

65. N. brevifolia C. A. M 401 

66. N. lamiifolia Willd 405 

Section 8. Orthonepeta Benth. 

67. N. pannonica L 406 

68. N. sulphurea C. Koch 409 

69. N. Sintenisii Bornm 411 

70. N. Komarovii E. Busch 412 

Section 9. Oxynepeta Benth. 

71. N. parviflora M. B 414 

72. N. ucrainica L 415 

11910. 73. N. Schischkinii Pojark 417 

74. N. kopetdaghensis Pojark 418 

75. N. erivanensis Grossh 420 

Section 10. Micronepeta Benth. 

76. N. bracteata Benth 422 

77. N. daenensis Boiss 425 

78. N. spathulifera Benth 427 

79. N. pungens (Bge.) Benth. . 429 

80. N. Fedtschenkoi Pojark 430 

81. N. microcephala Pojark 432 

82. N. satureioides Boiss 435 

Genus 1255. Glechoma L. 

11920. 1. G. hederacea L 437 

2. G. hirsuta W. at K 438 

3. G. longituba (Nakai) Kuprian 439 

Genus 1256. Dracocephnlum L. 
Subgenus 1. Eudracocephalum Briq. 
Section 1. Bu guide a Benth. 

1. D. pinnatum L 444 

2. D. palmatum Steph 445 

3. D. origanoides Steph 445 

4. D. Bungeanum Schischk. et Serg 446 

5. D. discolor Bge 447 

6. D. Paulsenii Briq 448 

7. D. botryoides Stev 448 

11930. 8. D. grandiflorum L 451 

9. D. fumosum Gontsch 452 

10. D. imberbe Bge 453 

11. D. Stellerianum Hiltebr 454 



XV 



11940. 



11950. 



12. 


D. 


13. 


D. 


14. 


D. 


15. 


D. 


16. 


D. 


17. 


D. 


18. 


D. 


19. 


D. 


20. 


D. 


21. 


D. 


22. 


D. 


23. 


D. 


24. 


D. 


25. 


D. 


26. 


D. 


27. 


D. 


28. 


D. 


29. 


D. 


30. 


D. 


31. 


D. 



fragile Turcz 454 

multicolor Kom 455 

Krylovii Lipsky 456 

fruticulosum Steph 456 

integrifolium Bge ' 457 

nutans L 458 

thymiflorum L 459 

nodulosum Rupr 460 

oblongifolium Rgl 461 

scrobiculatum Rgl 461 

foetidum Bge 462 

moldavica L 463 

peregrinum L 464 

bipinnatum Rupr 464 

heterophyllum Benth 465 

diversifolium Rupr 466 

spinulosum M. Pop 466 

Komarovii Lipsky 469 

multicaule Montbr. et Auch 470 

subcapitatum (Ktze.) Lipsky 470 



Subgenus 2. Ruyschiana (Mill.) Benth. 

32. D. argunense Fisch 

33. D. Ruyschiana L 

34. D. austriacum L 



471 
472 
472 



Subgenus 3. Fedczenkiella (Gontsch.) Schischk. 
35. D. stamineum Kar. et Kir 



473 



Genus 1257. K udr j a sc h ev i a Pojark. 



1. K. allotricha Pojark , 

2. K. Korshinskyi (Lipsky) Pojark. 
11960. 3. K. Nadinae (Lipsky) Poiark. . . 

4. K. Jacubi (Lipsky) Pojark. . . 



475 
476 
480 
481 



Genus 1258. L a 1 1 e m a nt i a Fisch. et Mey. 



1. L. peltata (L.) Fisch. et Mey. . 

2. L. canescens (L.) Fisch. et Mey. 

3. L. iberica (Stev.) Fisch. et Mev. 

4. L. Royleana Benth 

5. L. baldshuanica Gontsch 



484 
485 
485 
486 
487 



Genus 1259. H y m e no c r a t e r Fisch. et Mey. 

1. H. elegans Bge '. . . . . 

2. H. bituminosus Fisch. et Mey 



Genus 1260. Hypogomphia Bge . 

1. H. turkestana Bge 

11970. 2. H. elatior (Rgl.) Vass 



489 
490 



491 
492 



XVI 



Tribe 3. Stachydeae Briq. 
Subtribe 1. Brunelllnae Briq. 
Genus 1261. Prunella L. 

1. P. vulgaris L 495 

2. P. grandiflora (L.) Jacq 496 

3. P. laciniata L 497 

Subtribe 2. Melittinae Briq. 
Genus 1262. Mel ittis L. 
1. M. melissophyllum L 499 

Subtribe 3. Lamlineae Briq. 
Genus 1263. Pseudoeremostachys M. Pop. 
1. P. Sewertzowii (Herd.) M. Pop 500 

Genus 1264. Neustruevia Juz. 
11976. 1. N. karatavica Juz 502 



xvu 



PREFACE 

This volume deals with the first part of the Labiatae. It describes 369 species of which 
ten percent are new. The largest genera are Scutellaria (S.V. Yuzepchuk) and Nepeta 
(A.I. Poyarkova). As the latter proved to be too heterogeneous, it was considered neces- 
sary to introduce the genera Drepanocaryum and Kudrjaschevia. In addition to these 
two genera segregated by Poyarkova from Nepeta, Yuzepchuk established Neustruevia 
in the Stachydeae. 

Some genera of this family are of economic importance as they include plants yielding 
essential oil. Of these, Lavandula and Rosmarinus are grown in the southern regions 
of the USSR. 

The second part of Labiatae is presented in Volume XXL 

Editorial Board 



1 Family CXL. LABIATAE JUSS. 

Flowers always zygomorphic, with bilateral symmetry, rarely almost regular (Mentha, 
etc.), bisexual or gynodioecious, rarely polyoecious, always heteromerous and diagram- 
matically very uniform; calyx usually, corolla always 5-merous; lobes of corolla alternate 
with lobes or teeth of calyx; stamens alternate with lobes of corolla; androecium essen- 
tially 5-merous, tetra- or diandrous by reduction; gynoecium distinctly 2-merous, with 
median transverse carpels. 

Calyx variable, always with distinct tube, campanulate, tubular, infundibular, globular, 
urceolate or legeniform, 5-, rarely 4 -toothed or lobed or variously 2-lipped (more than 
5 teeth in genera with so-called commissural teeth, in Marrubium, Ballota and others 
6-10, in Molucella and others up to 13; calyx-tube straight or curved, in some cases 
one-sided (due to dorsal flattening), with squamiform or utricular dorsal projections (Scu- 
tellaria), and other features; limb of calyx likewise regular or irregular, with a range of 
variability from obsolescent lobes (Lavandula) or acute triangular teeth (Mentha, Sta- 
chys, etc.) to very large, coriaceous, colored lobes with reticulate venation (Hymeno- 
crater, etc.). There are three basic types of 2-lipped calyx: 1) formula ^/2, with 3 lobes 
in the upper lip and 2 in the lower lip (Salvia, Melissa, Satureja, Thymus, etc.), a bi- 
labiate calyx with entire lips being regarded as a modification of this type; 2) formula V4 
(some species of Teucrium, Dracocephalum, Wiedemannia, Ocimum), with transi- 

2 tion toward the ^/2 formula where the lateral teeth or lobes of the calyx are not involved 
in the formation of the lips (Plectranthus); 3) formula ^/o, with cleft lower lip (Majo- 
rana). The calyx is often more or less markedly accrescent, it is usually persistent in fruit, 
with nutlets ripening in the tube; sometimes it separates into two segments, one falling 
away, the other persistent (Scutellaria). Normally, the calyx has 5 principal nerves reach- 
ing the apex of the teeth and 5 secondary nerves terminating at the sinuses. The total 
number of nerves is often increased to 15 as a result of parallel doubhng of the secondary 
nerves (Nepeteae, Hyssopus); in other cases only the lower secondary nerves are dou- 
bled and the calyx becomes 13-nerved (it should be noted that very often the secondary 
nerves are doubled only in the middle or the lower third of the calyx-tube, and so the 
number of nerves has to be determined at the appropriate level. Corolla sympetalous, 
usually divided into tube and limb, the tube mostly cylindrical or enlarged above, erect 

or variously curved or twisted, of uniform width or expanded above into a wide throat, 
rarely campanulate; tube naked within or with a hairy ring lying below attachment of 
stamens forming a protective mechanism for nectar (nectarostegium); limb of corolla 
sometimes almost regularly 4-merous (Mentha, Lycopus, Dy sophylla), more often 
distinctly 2-lipped or 1-lipped. Three types of lip structure maybe distinguished: 1) most 
common is a corolla of the ^/a type, with the upper lip flat (Hyssopus, Satureja, 



Origanum, Thymus) or concave or convex (Galeopsis, Lamium, Stachys), and the 
middle lobe of the lower lip usually larger than the lateral and of different shape; 2) the 
corolla of most Ocimoideae is of the 7i formula, the single lobe of lower Up narrow and 
flat (Ocimum), concave (Plectranthus), navicular, etc.); 3) corolla of the "/s type, due 
to the tube being cleft above, all five lobes forming a lower lip (Teucrium); there is a 
transitional type of corolla with the lateral lobes in intermediate position, referred to the 
upper lip by some authors and to the lower Up by others. Variously shaped projections, 
called "pleuridia," frequently occur on the lobes; they are filiform on the lateral lobes in 
Lamium, rounded in Teucrium, and rounded on the lower lobe in Nepeta, Draco- 
cephalum. Salvia, Stachys, Lamium, etc. Stamens commonly 4, attached to corolla- 
tube, usually didynamous, the upper (posterior) mostly shorter or rarely longer than the 
lower (anterior) (Nepeteae), more rarely subequal (Mentha) or reduced to 2 and then the 
upper pair represented by staminodes or completely suppressed (Rosmarinus, Salvia, 
Lycopus, etc.), very rarely only the upper pair developed, the lower being absent or re- 
duced to staminodes (Orthodon). Filaments free, usually filiform, rarely connate into a 
sheath, sometimes bearing processes (apophyses) or hairy; anthers introrse, often 2-celled, 
globose, ovoid or oblong-linear, dehiscing longitudinally, mostly aUke, but deviations from 
this norm occur: thus, in Salvieae the anterior cell is obsolete or has another function, in 
Sideritis and Scutellaria the lower stamens have only one fertile cell, etc. The cells are 
normally parallel, but marked variations occur: the cells may be divergent or divaricate 
or even connivent apically and fusing into a single cell (Lavandula, Marrubium, Ocimoi- 
deae). The connective is variable: in Ocimoideae obsolete, in Satureieae strongly develop- 
ed, in Meriandreae forming a wide ledge from which anther-cells are suspended, in Salvieae 
filiform-elongate. Pollen grains usually ellipsoid, with 3-6 furrows. Gynoecium always 
2-merous, formed by 2 carpels but soon becoming seemingly 4-merous by constriction 
and false partitions. Ovary superior, sessile, rarely borne on a gynophore, 4-parted to 
base, rarely 4-lobed. A thickened, fleshy hypogynous disk is usually formed by processes 
of the receptacle; it is uniformly developed or 2-4-lobed and then its lobes as a rule alter- 
nate with those of the ovary, the lower lobe usually functioning as nectary, larger than 
others and differing in shape, the upper lobe rarely enlarged; lower part of the disk some- 
times (subfamily Scutellarioideae) forming a gynophore; in the Lavanduloideae the lobes 
of the disk are exceptionally opposite those of the ovary. The style usually develops from 
the base of ovary lobes, but it may be incompletely gynobasic or even subapical in rare 
cases (Ajugoideae, etc.); it is usually glabrous, sometimes hairy (species of Salvia), rarely 
simple, usually 2-partite at apex, the branches lobed or acute (in Sideritis etc. one branch 
lobelike, the other acute), often very unequal; ovules solitary in each ovary cell, nearly 
always anatropous, with micropyle facing downward, in some Ajugoideae inserted above 
the base of the ovary cell, semianatropous (Ajuga); in the Scutellarioideae amphitropous, 
concave or incurved. Placentation axial, mostly basiaxial; integument usually thin. Fruit 
commonly of 4 equal, glabrous, hairy or tuberculate nutlets; areola (scar of attachment) 
small, basal, rarely large, ventral (Ajugoideae), very rarely subdorsal. Seed soHtary in each 
nutlet, erect, very rarely transverse and curved, with thin testa; endosperm absent or 
scanty; embryo with flat, rarely convex or folded, somewhat fleshy cotyledons parallel 
to fruit axis, rarely transverse; radicle short, downturned, exceptionally curved, adjacent 
to cotyledons or lying on one of them (Scutellarioideae). 



Herbs (annual or perennial) or shrubs, very rarely (in tropics) trees or climbers, mostly 
aromatic, usually (not always) with epidermal glands. The main root usually persists 
throughout the hfe of the plant, very rarely (in moist habitats) it dies toward the end of 
the first vegetative season (Scutellaria, Stachys palustris, Mentha); the roots are 
sometimes thickened and serve as storage organs (species of Nepeta, Phlomis, Scutel- 
laria, etc.). Stems usually 4-angled, branches mostly opposite, sometimes whorled. Leaves 
usually opposite, rarely whorled (Dysophylla, etc.), simple, the blades entire, toothed, 
lobed, cut, etc. Spines rather rare, by transformation of upper leaves or their teeth (Cha- 
maesphacos)orbracts and bracteoles or merely theirnerves(Lagochilus). Inflorescence 
cymose, two cymes from a pair of leaves forming an apparent whorl [pseudowhorl]; brac- 
teoles usually 2, small, in monochasial cymes only one of each pair developed (Lamium, 
Satureja, etc.), more rarely both suppressed (Teucrium, Prunella, etc.). Flowers rarely 
solitary in axils of cauline leaves or bracts (Scutellaria, etc.), mostly crowded in the axils 
and forming so-called apparent whorls (verticillasters), each consisting of 2 cymose inflo- 
rescences subtended by opposite leaves; in its simpler form the verticillaster represents a 
3-flowered dichasium (Dracocephalum moldavica. Salvia pratensis, Brunella, etc.); 
mostly each cluster composed of dichasia which pass over into a pair of monochasial 
cymes; these may be few-flowered (Am ethystea, Lamium, etc.) or many-flowered 
(Nepeta, Ballota, Satureja, Lycopus, etc.); the axes of all verticillasters may be much 
reduced (Galeopsis, Lycopus, etc.) or reduction may involve all axes but one (Mentha, 
Ocimum, etc.) and then the entire cluster may be borne on a common peduncle; in some 
cases all the axes are developed; all verticillasters may be axillary or crowded at the apex 
of the stem, and then the upper leaves are usually transformed into distinctive bracts and 
the entire inflorescence becomes an apparent spike (spicastrum), apparent panicle or um- 
bellate inflorescence; when strongly contracted and congested the spike becomes a head; 
in some species all flowers are crowded toward one side of the main axis, producing a secund 
or dorsiventral inflorescence (Teucrium, Scutellaria, Nepeta, Salvia, etc.); dorsiven- 
trality is more pronounced when the bracts are opposite the flowers (e.g., Elsholtzia). 

A large family of cosmopohtan distribution, but especially abundant in the Mediterra- 
nean region (broadly conceived) and in mountainous countries of the subtropics; it does 
not reach the Arctic region nor is it typical for high mountains. Several thousand species 
in over 300 genera are known (adoption of a narrower concept of genera would produce 
many more). 

Major credit for the classification of the Labiatae is due to the English scientist Bentham. 
The two editions of his monograph (1832-1836 and 1848) were accepted as standard for 
many years. However, at the end of the last century the Swiss botanist Briquet revised this 
classification. He placed the Ajugoideae and Prostantheroideae, characterized by their 
non-gynobasic style, at the head of the family. He pointed out their affinity to Verbena- 
ceae (actually referring to them as Verbenoids) and their distinctiveness from the other 
subfamilies. It was Briquet, too, who separated the very unique subfamily Scutellarioi- 
deae. In his treatment of the Stachydoideae he radically departed from Bentham by con- 
sidering it as the nucleus of the entire family. Briquet's classification of Labiatae was in- 
corporated in Engler's system of plant families and has therefore been adopted in the 
present work. The imperfections of this classification are obvious; one of them, for 
example, would be the high rank given to the Satureieae and, in particular, to the Menthi- 
nae, though the simple structure of their flowers must be considered primitive. 



In the generic key, we have retained Briquet's classification as far as possible, so as to 
present it to the users of the "Flora." Yet, to facilitate the use of the key, we had to make 
significant modifications in many instances. More particularly, the generic breakdown as 
presented in the "Flora" is more far-reaching than in Briquet's classification; also quite a 
few new genera have since been described. 

Economic importance. The economic importance of the Labiatae is largely due to their 
specialized glands which excrete essential oils. Many species yield essential oil used in per- 
fumery, cookery and medicine. Some produce edible tuberlike rhizomes; many are culti- 
vated as ornamentals. For further details, see notes to genera and species. 



Key to Genera 

1 . Style inserted above base of lobes of ovary; nutlets with lateroventral attachment, 
the large areola (scar of attachment) extending over half the height of the nutlet; 
corolla 1 -lipped or apparently so owing to underdeveloped upper lip, rarely 2-hpped 
(Amethystea and Rosmarinus) (subfamily Ajugoideae) 2. 

+ Style inserted at base of lobes of ovary, i.e. basal or gynobasic; nutlets with basal, 
very rarely basidorsal or ventral attachment, the areola usually small, very rarely 
large; corolla not 1-lipped 5. 

2. Corolla 1-lipped or apparently so, rarely 2-lipped and then the upper lip not concave; 
stamens 4, rarely 2; anthers 2-ceUed; nutlets rugose (tribe Ajugeae) 3. 

+ Corolla distinctly 2-lipped, the upper lip concave; stamens 2; anthers 1 -celled; nut- 
lets smooth (tribe Rosmarineae) 1243. Rosmarinus L. 

3. All 4 stamens well developed; corolla 1 -lipped or apparently so 4. 

+ Only 2 lower stamens developed, the upper 2 reduced to staminodes; corolla 

2-lipped 1242. Amethystea L. 

4. Corolla essentially 2-hpped but appearing 1-hpped because of the pronounced dispar- 
ity of the lips: the upper lip very short, 2-partite or 2-lobed, the lower large, with 
highly developed middle lobe 1240. Ajuga L. 

+ Corolla genuinely 1-lipped, all 5 lobes dechned, lying decidedly below the stamens 
and style, exserted from the 2 upper lobes and forming a single lower lip ... . 
. . 1241. Teucrium L. 

5. Calyx 2-Upped, the entire short Ups of 2 parts or segments, the upper segment usual- 
ly with rounded, concave dorsal scale, rarely with a bladderlike appendage or unap- 
pendaged, usually deciduous, the lower segment frequently persistent; upper lip of 
corolla apparently 3-lobed, its lateral lobes adnate to the middle lobe, rarely free 
and then it may also be included in the lower lip (subfamily Scutellarioideae). . . 
1244. Scutellaria L. 

+ Calyx different; upper lip of corolla (in our genera) usually not 3-lobed. ... 6. 

6. Lobes of disk oblong-rounded, opposite lobes of ovary and covering their lower part; 
nutlets with basidorsal areola. Perennial plants, cultivated (in USSR); leaves linear- 
lanceolate, with revolute margins; calyx 2-lipped, the upper lip with 1 tooth, the 
lower with 4 teeth (subfamily Lavanduloideae) 1245. Lavandula L. 



+ Lobes of disk (if distinct) alternate with those of ovary (the large, crescent-shaped 
appendages enveloping the nutlet in Drepanocaryum (described at the next stage) 
should not be mistaken for disk lobes) 7. 

7. Disk with 4 large crescent-shaped appendages alternating with disk lobes and envel- 
oping the base of ovary cells (and nutlets) obsolescent in fruit; areola large, basi- 
ventral. Annuals, with small flowers; calyx pellucid, scarious, with 18-20 nerves, 
obhquely inflated at base; 4 fertile anthers (subfamily Drepanocaryoideae) . . . 
1246. Drepanocaryum Poj ark. 

-I- Disk without appendages; areola small, usually basal; calyx different 8. 

8. Stamens included in corolla-tube or exserted, ascending or projecting forward (sub- 
family Stachydoideae) 9. 

+ Stamens exserted, descending, appearing to lie on slightly concave, 1-lobed entire 
lower lip of corolla; upper lip rounded, with 4 shallow even lobes (subfamily 
Ocimoideae) 68. 

9. Stamens (and pistil) included in corolla-tube, this usually enclosed in the calyx, very 
rarely stamens exserted 10. 

-I- All stamens, or at least the 2 longer ones, more or less exserted; corolla-tube mostly 
(though not in all genera) not enclosed in calyx 15. 

10. Corolla-tube narrow, exserted from calyx, the upper lip narrow, nearly flat; nutlets 

narrow, oblong (nearly linear). Annuals (Central Asia) 

1247. Tuspeinantha Durand. 

-I- Corolla-tube enclosed in calyx; nutlets ovoid or ellipsoid. Perennials, rarely annuals 
11. 

11. Corolla very small, not exserted from calyx-tube; corolla-tube without a ring of 
hairs; verticillasters 4 or 5, very distant, 2-flowered. Densely tomentose perennials, 
with oblong entire leaves 1283. Phlomidoschema (Benth.) Vved. 

-1- Limb of corolla exserted from calyx-tube; verticillasters many-flowered. . . .12. 

12. Anthers with divaricate cells; calyx 5-10-toothed; verticillasters axillary, in axils of 
ordinary leaves. Plants usually perennial (very rarely annual) 13. 

+ Anthers of upper stamens with divergent cells; calyx 5-toothed; verticillasters in 
false spike, subtended by differentiated bracts, or axillary and then plants annual . 
1250. SideritisL. 

13. Calyx 5-10-nerved, with 5-10 equal or unequal teeth; corolla-tube with a faint and 
uneven ring of hairs or glabrous inside; upper Up of corolla nearly flat or concave; 
filaments glabrous; base of leaves not sagittate 14. 

+ Calyx 10-nerved, with 5 equal teeth; corolla-tube with well developed ring of hairs; 
upper lip of corolla slightly inflated; filaments villous-hairy; base of leaves sagittate 
1280. Metastachys Knorr. 

14. Calyx 5-10-toothed; corolla-tube with a faint and uneven rmg of hairs; upper lip 
notched or 2-fid; leaves orbicular or ovate, usually crenate. . 1248. Marrubium L. 

+ Calyx 5-toothed; corolla-tube without ring of hairs; upper lip entire. Leaves round- 
ed, palmatifid 1249. Lagopsis Bge. 

15. Anther-cells spreading, divergent or parallel, oblong or ovoid, not confluent or indis- 
tinctly confluent at apex, not disposed in one plane after dehiscence 16. 

+ Anther-cells spreading, globular, confluent at apex, disposed in one plane after 

dehiscence (tribe Pogostemoneae) 67. 



16. Connective of 2 fertile lower stamens linear-filiform, loosely or firmly jointed with 
filament, its posterior limb bearing a normally developed linear cell, the anterior 
limb naked or bearing a modified barren cell (tribe Salvieae) 17. 

+ Stamens different 18. 

17. Calyx green or violet (in our species); lips of fruiting calyx unchanged or slightly 
modified, the upper lip distinctly 3-toothed. 1285. Salvia L. 

. + Calyx pink or yellow (in our species); fruiting calyx with strongly accrescent, coria- 
ceous, netted-veined lips, the upper lip entire 1286. Schraderia Moench. 

18. Corolla distinctly 2-lipped, with lips of unequal length, the upper generally concave 
or convex (flat only inChamaesphacos, in which the leaves are spine-tipped . 19. 

+ Corolla distinctly 2-Upped, with nearly flat upper Up, lips faintly differentiated, or 
nearly actinomorpliic 51, 

19. Each of the 2 lower fertile stamens bearing 2 parallel, oblong-linear, slightly curved 
cells, these approximate and hanging from a slightly dilated connective. Central 
Asian herbs or subshrubs, with heavily pubescent 2-lipped calyx and deeply toothed 
or cut leaves (tribe Meriandreae) 1287. Perovskia Karel. 

+ Fertile stamens usually 4, or 2 and then connective not dilated and anther-cells ovoid 
or ellipsoid, not pendulous 20. 

20. Upper stamens longer than the lower, the latter sometimes obsolete (tribe Nepeteae) 
21. 

+ Upper stamens shorter than the lower (tribe Stachydeae) 30. 

2 1 . Only upper stamens or all stamens fertile, in any case the lower with very small 

10 anthers, mostly concealed in corolla-tube 22. 

+ All 4 stamens fertile, with normally developed, equal or nearly equal anthers (in 
some genera there are staminate individuals with 4 staminodes included in the 
corolla-tube) 23. 

22. Only upper stamens fertile, the lower filiform, apically clavate staminodes; upper 
hp of corolla narrow, much longer than the lower; calyx campanulate, with nearly 
straight throat. Annuals 1260. Hypogomphia Bge. 

+ Lower stamens short, mostly included in corolla-tube, rarely exserted, not stamino- 
dial, or upper stamens reduced to sterile staminodes and then plants annual; upper 

lip of corolla broad, shorter than the lower; calyx 2-lipped 

1257. Kudrjaschevia Pojark. 

23. Fruiting calyx unchanged or only shghtly modified 24. 

+ Fruiting calyx enlarged at apex into a large coriaceous reticulate 5-lobed fringe . . 

1259. Hymenocrater Fisch. et Mey. 

24. Flowers seemingly inverted by contortion of the corolla-tube; upper lip of corolla, 
normally overlying stamens and style, occupies the position of the lower lip and 
vice-versa 1252. Lophanthus Adans. 

+ Flowers not inverted 25. 

25. Pairs of stamens not parallel 26. 

+ Pairs of stamens parallel, arched under the upper lip of corolla 27. 

26. Upper stamens declinate, the lower ascending; lobes of disk hardly discernible; mid- 
dle lobe of lower lip of corolla not unguiculate 1251. Agastache Clayt. 

+ Upper stamens ascending, the lower directed forward; lobes of disk well developed; 
middle lobe of lower lip of corolla unguiculate .... 1253. Schizonepeta Briq. 



27. Upper lip of corolla with protruding fold inside; pedicels flattened 

1258. Lallemantia Fisch. et Mey. 

+ Upper lip of corolla smooth inside; pedicels not flattened 28. 

28. Calyx usually tubular, 5-toothed, rarely 2-lipped, always without protuberance in 
sinuses 29. 

+ Calyx distinctly tubular (formula V4 or ^/j), with a protuberance in some or all 

sinuses 1256. Draco cephalum L. 

29. Anther-cells diverging at an angle of 180°. Plant without creeping stems .... 
1254. NepetaL. 

+ Anther-cells diverging at a right angle. Plant with creeping stems and obtuse leaves 

1255. Glechoma L. 

++ Anther-cells parallel. Plant with creeping stems and acute leaves 

1252a. Meehania Britton. 

30. Calyx 2-lipped, the lower lip turned toward the upper after flowering and covering 
the throat (subtribe Prunellinae) 1261. Prunella L. 

+ Calyx different 31. 

31. Calyx broadly campanulate, membranous, with 3-5 broad unequal teeth or lobes 
(subtribe Melittinae) 1262. Melittis L. 

+ Calyx tubular, campanulate or rotate, 5-10-toothed, rarely 2-lipped (subtribe 

Lamiinae) 32. 

32. Upper lip of corolla concave or hooded, rarely convex, usually very hairy . . .33. 
+ Upper lip of corolla flat, glabrous or very sparsely hairy 

1284. Chamaesphacos Schrenk. 

33. Style-branches very unequal, the upper much shorter than the lower, rarely nearly 
equal (in many species of Eremostachys) and then all or only upper stamens with 
basal appendages 34. 

+ Style-branches equal or nearly so, filaments without basal appendages 36. 

34. Lower stamens hardly exserted from corolla-tube, the upper not exserted; filaments 
unappendaged; nutlets with tufts of hairs at apex; bracts Unear-subulate, not con- 
nate; leaves ovate, entire 1263. Pseuderemostachys M. Pop. 

+ Anthers of all stamens much exserted from corolla-tube; filaments generally with, 
rarely without appendages 35. 

35. Appendages of upper filaments spurlike, smooth; calyx-teeth rather long and narrow, 
setaceous from obtuse apex; nutlets pubescent or glabrous; leaves always entire . 
1266. PhlomisL. 

+ Appendages of all or only of the upper filaments more or less deeply or pectinately 
cleft, very rarely spur-shaped, and then tuberculate-dentate; calyx-teeth very short 
and broad, abruptly mucronate from truncate apex, the mucro mostly short; nutlets 

very hairy at apex; leaves commonly pinnatisect, rarely entire 

1265. Eremostachys Bge. 

36. Upper lip of corolla short, half as long as the lower, nearly square, shghtly convex; 
anther-cells divaricate, apically confluent with a common slit; verticillasters few, 
very dense and compact; bracts linear-lanceolate, falcate, rigid, prickly, connate at 
base; leaves pinnatisect 1264. Neustruevia Juz. 

+ Flowers and other parts different 37. 



37. Lower lip of corolla with 2 hollow hornlike protuberances; anther-cells forming a 
right angle at anthesis 1268. Galeopsis L. 

+ Lower lip without protuberances; anther-cells parallel or divergent 38. 

38. Nutlets (lobes of ovary less distinctly so) acutely trigonous, obtuse 39. 

+ Nutlets (and lobes of ovary) ovoid, more or less rounded at apex 48. 

39. Calyx tubular or campanulate, and if infundibuliform then filaments joined by thick 
pubescence; anther-cells with ciUate slit 40. 

+ Calyx clearly infundibuliform, with 5 short arched mucronate lobes, filaments not 
pubescent; anther-cells glabrous 1277. Molucella L. 

40. Corolla-tube usually elongate, markedly exserted from calyx; calyx-teeth rather soft, 
not prickly or spinous 41. 

+ Corolla-tube included in calyx or slightly exserted; calyx-teeth more or less prickly 
or spinous, rarely mucronate 44. 

41. Calyx-teeth equal or nearly so 42. 

+ Calyx 2-Hpped, the upper lip 1-lobed, the lower composed of 4 teeth united at base 

1272. Wiedemannia Fisch. et Mey. 

42. Lateral lobes of lower Up of corolla relatively large, ovate, obtuse, entire or emargi- 
nate; leaves approximate, broadly rhombic or cuneate-flabelliform, densely 

velutinous-tomentose. Central Asian high-mountain plant 

1271. Erianthera Benth. 

+ Lateral lobes of lower lip of corolla and other parts different 43. 

43. Lateral lobes of lower hp of corolla relatively well developed, lanceolate, acute, en- 
13 tire; flowers yellow 1270. Galeobdolon Adans. 

+ Lateral lobes of lower lip of corolla reduced to small appendages with one to several 
acute marginal teeth; flowers not yellow. . 1269. Lamium L. 

44. Upper Up of corolla 2-fid or emarginate. Central Asian subshrub, with spines at base 

of verticillasters (modified bracts) and frequently also in leaf axils 

1276. Lagochilus Bge. 

+ Upper Hp of corolla entire; no spines at base of verticillasters or in leaf axils . . 45. 

45. Anther-cells strongly divergent ; leaves ovate, entire 46. 

+ Anther-ceUs parallel; leaves of different shape, deeply divided 47. 

46. Flowers 5-7 mm long; corolla scarcely longer than calyx-teeth; calyx-tube without 

a ring of hairs; stamens slightly exserted; nutlets puberulent at apex 

1273. Chaiturus Willd. 

+ Flowers over 1 cm, commonly 1 .5-2 cm long; corolla much longer than calyx, its 

tube with a ring of hairs; stamens long-exserted; nutlets glabrous 

1267. Stachyopsis M. Pop. et Vved. 

47 Calyx usually small, not more than 9 mm long, funnel-shaped, 5-nerved, with nar- 
rowly triangular teeth; corolla pink, enlarged (inflated), with a ring of hairs; upper 
lip of corolla slightly concave, mostly tapering at base; lower Hp usually de flexed . 
1274. Leonurus L. 

+ Calyx much larger (13-1 8 mm), tubular-campanulate, 10-nerved (5 nerves less dis- 
tinct than the rest), with broadly triangular teeth; coroUa yeUow, its tube narrow, 
without ring of hairs; upper lip hooded, not tapering at base; lower lip straight, 
only the lobules of middle lobe deflexed 1275. Panzeria Moench. 



10 



48. Calyx enlarged above to an oblique fringe, the upper lip short, entire or 3-toothed, 
the lower large, angled or 4 -toothed 1278. Otostegia Benth. 

+ Calyx different 49. 

49. Calyx tubular-infundibular, with 5-10 (or more) teeth enlarged at base or united 
into a round toothed fringe; stamens straight after flowering . . .1279. Ballota L. 

+ Calyx tubular, with 5 triangular equal acute teeth or upper teeth longer than the 
lower, rarely calyx 2-hpped (formula ^/2); after flowering stamens recurved against 
sides of upper lip 50. 

50. Spikelike inflorescences usually dense, short-cylindrical or subcapituliform; corolla- 
tube exserted, without ring of hairs; anther-cells parallel . . . 1282. Betonica L. 

+ Spikelike inflorescences different, flowers sometimes axillary; corolla-tube usually in- 
cluded in calyx, with ring of hairs within; anther-cells divergent . 1281. Stachys L. 

51. Corolla 2-lipped 52. 

+ Corolla nearly actinomorphic 64. 

52. Only lower stamens fertile, the upper reduced to staminodes or absent .... 53. 
+ All 4 stamens fertile (tribe Saturejeae) 54. 

53. Calyx narrowly cylindrical, the limb much shorter than the tube; upper Up of corol- 
la entire, the lower 3-lobed. Perennials or annuals, with flowers crowded in capitate, 

approximate, rarely remote verticillasters (genus of tribe Saturejeae) 

1288. Ziziphora L. 

+ Calyx campanulate, the limb as long as the tube; upper lip of corolla shortly 3-lobed, 
the lower faintly emarginate. Annuals, with elongate, racemiform inflorescences 
(subtribe Perillinae) 1304. Orthodon Benth. et Oliver. 

54. Calyx regular or nearly so, 5-toothed 55. 

+ Calyx irregular, at least after flowering distinctly 2-lipped, sometimes 1 -lipped . 57. 

55. Calyx 15-nerved, stamens long-exserted from corolla (subtribe Hyssopinae) . . . 
1296. Hyssopus L. 

+ Calyx 10-13-nerved, or 15-nerved and then stamens shorter than upper lip of corolla 
56. 

56. Flowers subsessile, distinctly bracteolate; calyx 13-nerved, the nerves distinct only 
at base; calyx-teeth incurved 1298. Origanum L. 

+ Flowers on long pedicels, with poorly developed bracteoles; calyx distinctly 13-15- 
nerved, the teeth straight 1291. Micromeria Benth. 

57. Bracts large, equaling or exceeding the calyx, rounded or rounded-ovate. . . .58. 
+ Bracts small, shorter than the calyx, rarely longer and then of different shape. . 59. 

58. Calyx cylindrical, the upper lip entire or 3-toothed, the lower small, 2-toothed; spic- 
ules small, drooping, with colored bracts 1295. Amaracus Gleditsch. 

+ Calyx campanulate, with obUque throat, the upper lip subentire, the lower entire or 
absent; spicules larger, erect; bracts pale green .... 1297. Majorana Moench. 

59. Calyx 10-nerved 60. 

+ Calyx 13-1 8-nerved 62. 

60. Corolla exserted, ascending below middle and deflexed .... 1289. Melissa L. 
+ Corolla included in calyx or exserted, straight or slightly curved 61. 

61. Calyx 2-lipped; stamens straight, directed forward, distant, protruding at sides of 
lower lip 1299. Thymus L. 



11 



16 



+ Calyx with 5 subequal teeth, stamens curved, ascending, approximate, lying under 
upper lip 1290. Satureja L. 

62. Bracts long, setiform, long-hairy; calyx with hairy throat . 1293. Clinopodium L. 
+ Bracts different; calyx with ring of hairs in throat 63. 

63. Calyx tubular or tubular-campanulate, not gibbous at base; the triangular teeth not 
acuminate; pedicels terete 1292. Calamintha Lam. 

+ Calyx curved-urceolate, constricted at throat and gibbous at base, the teeth acumi- 
nate; pedicels flattened 1294. Acinos Moench. 

64. Calyx usually regular even in fruit, rarely irregular, with 5 equal, rarely unequal 
teeth, not drooping in fruit (subtribe Menthinae) 65. 

+ Fruiting calyx 2-lipped, drooping (subtribe Perillinae) 1303. Perilla L. 

65. All 4 stamens fertile, well developed 66. 

+ Only the 2 lower stamens fertile, the upper reduced to filiform staminodes. . . . 

1300. Lycopus L. 

66. Calyx 10-nerved; corolla-tube not exserted from calyx; anther-cells parallel. Herbs, 
with toothed, non-coriaceous leaves 1301. Mentha L. 

+ Calyx 15-nerved; corolla-tube exserted from calyx; anther-cells divergent. Shrubs, 
with entire, coriaceous leaves 1302. Gontscharovia Boriss. 

67. Corolla slightly 2-lipped. Annuals, with opposite leaves and one-sided, compact, 
spicate inflorescences, the large, orbicular imbricated ciliate bracts facing the flowers 
1305. Elsholtzia WiUd. 

+ Corolla with nearly equal lobes. Perennials, with whorled leaves and non-secund cy- 
lindrical inflorescences, the small bracts imbricated and not facing the flowers . . 
1306. Dysophylla Blume. 

68. Lower lip of corolla long, concave, or navicular; corolla-tube exserted 

1307. Plectranthus L'Herit. 

+ Lower lip of corolla flat or slightly concave, barely exceeding the lobes of upper 
lip, corolla-tube included in the calyx 1308. Ocimum L. 



Subfamily 1 . AJUGOIDEAE Benth. Lab. gen. -t sp. (1835) 651 ; id. in Benth. et Hook, 
fil. Gen. 111.(1876) 1171, emend.; Briq. in Pflanzenfam. IV, 3a (1895) 206. - Style not 
gynobasic, inserted above the base of ovary lobes. Nutlets with lateroventral attachment, 
the area of attachment usually large, often more than half the height of the nutlet. 

Note. Ajugoideae and the purely Australian Prostantheroideae occupy an isolated po- 
sition among the Labiatae because of the unique attachment of the style and ovules. Bri- 
quet even went so far as to maintain that these two subfamilies were phylogenetically un- 
related to the Labiatae proper and proposed their inclusion in the Verbenaceae. 



Tribe 1. AJUGEAE Benth. in DC. Prodr. XII (1848) 971 ; Briq. in Pflanzenfam. IV, 3a 
(1895) 206 et 208. - Calyx 10-nerved, with 5 equal teeth according to formula V4, very 
rarely ^/2, or with 2 entire lips; corolla 2-lipped of the V3 type, often with obsolete upper 
lip or 1-Hpped (^/s); stamens 4, rarely 2; anthers parallel or divergent; ovary shortly 4- 
celled or divided for V3 of its height. Nutlets obovoid, with large ventral areola. Embryo 
straight, with short straight radicle. 

12 



Genus 1240* Ajuga** L. 

L. Sp. pi. (1753) 561; Benth. Lab. gen. et sp. (1835) 690. - Moscharia Forsk. Fl. aegypt.-arab. 
(1775)158.- Cham aepitys Link, Handb. 2(1829)453.- Phleboanthe Tausch in Flora, XI 
(1828) 322. - Rosenbachia Rgl. in Tr. Bot. sada, IX, 2 (1886) 613. 

Flowers ebracteate, in 2- to many-flowered axillary verticillasters, sometimes aggregated 
at ends of shoots to form dense spiciform inflorescences; calyx campanulate or obovoid, 
5-toothed or 5-fid, regular except for the sometimes shorter upper tooth; corolla marces- 
cent, 2-hpped, the upper lip very short, bipartite or retuse, the lower long, 3-lobed, the 
large obcordate or reniform middle lobe more or less deeply cleft; corolla-tube with ring 
of hairs in lower part, straight, sometimes spirally twisted so that the lower lip is turned 
upward; stamens 4, exserted or included in corolla-tube, the upper pair longer than the 
lower; anthers 2-ceUed at first, the divergent cells at length confluent and dehiscent by 
common slit; nutlets reticulate or transversely rugose, with large areola, sometimes envel- 
oped by an appendage; ants which feed on these appendages contribute to seed dispersal. 
Annual, biennial or perennial plants or subshrubs; stems erect or ascending, sometimes 
stoloniferous; leaves entire or the cauline 3-lobed or 3 -partite; bracts usually resembling 
cauline leaves, rarely differentiated and tinged reddish or blue. 

Ajuga has about 50 species widely distributed in the Old World. 

Note. Since the publication of De Candolle's "Prodromus" (1848), Ajuga has been 
divided into three sections; of these Champaepitys Benth. and Bugula Benth. are re- 
presented in the Soviet Union. Later authors were mainly engaged in splitting the sections 
into subsections or series, until Boissier (Flora orientalis, IV, 1 879, 804) set up the section 
Phleboanthe, consisting of a single species (A. laxmannii) and characterized by its trun- 
cate corolla-tube. In his very useful treatment of section Bugula, Maksimovich (Diagn. 
plantar, nov. asiatic, V, 1 883) established a number of series composed of morphological- 
ly similar species. His classification of the section Bugula was fully adopted by Briquet 
(Engl. - Pr. Pflanzenfam. IV, 3a, 1897). The section Chamaepitys Benth. was divided 
by Briquet into two subsections — Ivae and Phleboanthe, thus incorporating the section 
Phleboanthe Boissier. Beside A. laxmannii, Briquet included in Phleboanthe all the 
integrifoliate species - A. salicifolia, A. oblongata, A. turkestanica, A. chamaecis- 
tus. We find it impossible to combine these Irano-Anatolian species, all characterized by 
closed corolla-tube, with A. laxmannii, and we place them in a separate subsection. A. 
laxmannii, with its open corolla-tube, appears to be a transition to Teucrium. 

Economic importance. The species Ajuga chia, A. pseudochia, A. reptans and 
A. genevensis are widely known for their medicinal properties. The leaves of A. rep- 
tans contain provitamin A (527). Currently, of special interest are A. chia, A. pseudo- 
chia and A. reptans, because of their hemostatic and wound-healing action. 

1. Flowers soHtary or rarely paired in the axil of each bract, thus verticillasters 

2-flowered, rarely 4-flowered 2. 

+ Verticillasters 6- to many-flowered 9. 

* Treatment by V.V. Pis'yaukova. 

** Ajuga may be derived from aguia, and this in turn from the Greek agyios, meaning joints, referring 
to the use of the plant in the treatment of gout. Alternately, ajuga is a distortion of abiga, Pliny's 
name (Hist. Nat., XXIV) for Chamaepitys. Later Latin authors write Ajuga instead of Abiga. 



13 



2. Leaves 3-lobed or 3 -partite, only radical leaves entire or toothed; flowers yellow 3. 
+ All leaves entire 5. 

3. Flowers much shorter than the bracts; calyx about equaling the corolla-tube; nut- 
lets oval, reticulate-rugose 3. A. chamaepitys (L.) Schreb. 

+ Flowers about equaling or exceeding the bracts; calyx about half as long as the 

corolla-tube; nutlets oblong, transversely rugose, reticulate-rugose only at apex 4. 

4. Stems and leaves densely covered with spreading hairs; calyx uniformly hairy . . 
1 . A. chia Schreb. 

+ Stems with short appressed hairs on alternate ribs and sparse spreading hairs else- ~ 
where; calyx hairy under the teeth, glabrous below. . . . 2. A. pseudochia Shost. 

5. Flowers yellow, with purple stripes along the nerves of corolla-tube 6. 

-I- Flowers bright purple 8. 

6. Corolla-tube cleft from upper lip to nearly the middle; calyx-teeth broad, ovate- 
lanceolate, obtusish; stems simple, thick, villous-hairy, densely leafy, the sessile 
leaves to 50 mm long 8. A. laxmannii (L.) Benth. 

19 + Corolla-tube entire; calyx-teeth lanceolate, acute; leaves 20-35 mm long; stems 

branching; plants with spreading or appressed hairs 7. 

7. Leaves lanceolate-elliptic or broadly eUiptic, short-petiole d, acutish, remote; plants 
appressed-hairy, with many slender stems; calyx 1/3 as long as corolla, hairy all 
over 4. A. salicifolia (L.) Schreb. 

+ Leaves mostly lanceolate, sessile, obtuse, densely covering stem; plants patent- 
villous; stems with declined branches; calyx 1/4 to 1/5 as long as corolla, glabrous 
below 5. A. oblongata M.B. 

8. Stems 3-5 mm in diameter, leafy, glaucescent, very finely soft-pubescent mainly in 
upper part; leaves 30-60 mm long, 12- 18 mm broad; corolla-tube nearly twice as 
long as calyx; calyx-teeth accrescent in fruit ... 7. A. turkestanica (Rgl.) Briq. 

+ Stems 1 .5-3 mm in diameter, white, profusely short -branched, hispid as are the 
leaves; sterile shoots often leafless, becoming woody, spinous; leaves 18-30 mm 

long, 4-10 mm broad; corolla-tube shorter; calyx-teeth not accrescent 

6. A. chamaecistus Ging. 

9. Plants with creeping stolons 10. 

+ Plants non-stoloniferous 11. 

10. Radical leaves spatulate, rounded at apex, entire or obscurely and broadly emargi- 

nate; lateral lobes of lower lip truncate; calyx glabrous below 

10. A. shikotanensis Miyabe et Tatewaki. 

+ Radical leaves broadly ovate, sinuate-dentate; lateral lobes oflower lip round-tipped; 
calyx commonly hairy all over 9. A. reptans L. 

1 1 . Stamens and style included; corolla-tube twisted and thus the lower lip turned up- 
ward; low, lanate-villous plants 14. A. orientalis L. 

+ Corolla-tube straight; stamens and style exceeding the corolla-tube 12. 

12. Inflorescence compact, 4-angled-pyramidal, starting close to soil level; bracts broad- 
ly ovate or oblong-ovate, entire, rarely obscurely toothed . .13. A. pyramidalis L. 

+ Inflorescence not 4-angled, starting higher up, at least lower verticillasters remote . 
13. 



14 



13. Plants 8-25 cm high, densely white-villous; bracts entire or sparsely toothed at apex; 
verticillasters with 8 or more flowers; calyx covered all over with long hairs, the nar- 
row acute teeth longer than the tube 12. A. multiflora Bge. 

+ Plants 10-40 (50) cm tall, villous-hairy mainly in upper part; bracts 3-lobed or 
coarsely 3-toothed at apex; verticillasters 4-6-flowered; calyx with woolly hairs 
mainly in upper part, the triangular-lanceolate teeth shorter than the tube .... 
11. A. genevensis L. 



Section 1. Chamaepitys Benth. Lab. gen. et sp. (1835) 691. — Verticillasters 2-flow- 
ered, rarely 4-flowered; corolla yellow or purple, with very short upper lip. 



Subsection 1. Ivae Briq. in Pflanzenfam. IV, 3a (1897) 210. - Cauline leaves lobate 
or 3-partite, with very narrow or linear lobes. Upper lip of corolla weakly developed, with 
obsolete lobes. 

1. A. chia Schreb. PI. vert, unilab. (1774) 25; Benth. Lab. gen. et sp. 699; in DC. 
Prodr. XII, 601 , p. p.; C. Koch in Linnaea XXI, 703, p. p.; Ldb. Fl. Ross. Ill, 449, p. p. 
(quoad pi. taur. Steveni); Boiss. Fl. or. IV, 802, p. p.; Shmal'g. Fl. II, 346, p. p.; N. Pop. 
in Mat. fl. Kavk. IV, 3, 11, p. p.; Vizn. rosl. URSR (1950) 407. - A. intermedia Boiss. 
1. c. pro synon. A. chiae. — A. chamaepitys Guss. Fl. Sic. Prodr. II (1828) 66; M. B. 
Fl. taur.-cauc. II, 34 p. p. — A. chamaepitys v. grandiflora Stev. in Bull. Soc. Nat. 
Mosc. 30(1857)366. - Teucrium chium Poir. Encycl. Meth. Suppl. II (1812) 772.- 
Ic: Sibth. et Sm. Fl. Graec. VI, tab. 524; N. Pop. in Tr. Bot. sada Yur'evsk. univ. XIV, 
24, Fig. 2 - fruit. 

Perennial, sometimes subshrub, canescent; stems often reddish, erect or ascending, 
usually evenly covered all round with long, white, sometimes rather coarse hairs, and if 
hairy on two faces only then the hairs long, spreading; lower leaves entire or apicaUy 3- 
toothed or 3-lobed, spatulate or oblong-lanceolate, tapering to a long petiole, the upper 
3-partite with linear entire lobes, all leaves villous-pilose; flowers axillary, solitary (verti- 
cillasters 2-flowered), subsessile, exceeding or about equaling the bracts, usually forming 
a rather compact, apical spikelike inflorescence; calyx broadly campanulate, long-hairy 
all over, 4-6 mm long, slightly ribbed, the 5 teeth broadly triangular at base, abruptly 
acuminate, nearly subulate, the upper teeth shorter than the lower; corolla 19-25 mm 
long, 4-5 times the length of calyx, yellow, with purple spots and stripes on lower lip and 
along the nerves of tube, densely covered outside with long white hairs as well as short 
clavate glandular hairs, the latter more abundant on inner surface; corolla-tube greatly 
exceeding the calyx, sometimes twice as long, the upper Up slightly 2-lobed, the large 
middle lobe of lower lip long-unguiculate and broadly cleft, the lateral lobes obovate, 
obtuse; filaments and style densely covered with long hairs, much exserted; nutlets 
oblong, 3-4 mm long, reticulate at apex, transversely rugose beneath, rarely with few 
oblique anastomoses between wrinkles, the areola 2/3 the length of the nutlets. May- 
September. (Plate I, Figure 2.) 



15 



Dry steppe slopes. — European part: U. Dns.. BL, Crim.; Caucasus: Cisc. (Mozdok), 
W. Transc. Gen.distr.: Med., Bal.-As. Min. Described from Chios. Type in Paris. 

Note. A very polymorphous species, originating from the eastern Mediterranean area 
and Asia Minor. In the USSR it is confined to ancient Pontus. One specimen in the her- 
barium similar to A. chia but subglabrous was collected by A. Shelkovnikov (17 VII 
1928) near Lake Sevan; it comes very close to A. glabra Presl (Fl. Sic. 36) which grows 
outside the USSR, e.g. near Lake Van. 

2. A. pseudochia Shost. in Hot. mat. gerb. Bot. inst. ANSSSR(1940) 147; Vizn. rosl 
URSR,407. - A. chia Benth. in DC. Prodr. XII (1848) 601, p.p.; Ldb. Fl. Ross. Ill, 
449, p.p.; Boiss. Fl. or. IV, 802, p.p.; Shmal'g. FL II, 346, p.p.; Fedch. Rast. Turk. 671; 
N. Pop. in Mat. Fl. Kavk. IV, 3, 1 1, p.p. et auct. Fl. Ross. - A. chia var. pseudocha- 
maepitys N. Pop. I.e.; Grossg. Fl. Kavk. Ill, 281. - A. chamaepitys M. B. Fl. taur.- 
cauc. II (1808) 34, p. p.; Guss. Fl. Sic. Prodr. II, 66, p. p.; Ldb. 1. c. p. p.; Shmal'g., op. 
cit. p.p.; Boiss. I.e. p.p.; C. Koch in Linnaea XXI, 703; Bge. Lab. Pers. 84, non L. - 
A. chamaepitys var. grandiflora Lindem. Fl. Elisabethgr. (1868) 120. - A. chamae- 
pitys p. hirta Freyn in Oesterr. Bot. Zeitschr. XXVI (1876) 405. - Teucrium chamae- 
pitys Falk. Beitr. Kenntn. Russ. Reichs II (1786) 204, non L. - Ic: N. Pop. in Tr. Bot. 
sada Yur'evsk. univ. XIV, 24 - fruit. 

Perennial, sometimes subshrub; stems 8-30 (37) cm long, branching at base, erect or 
ascending, alternately appressed-hairy on two faces, sometimes (mostly at nodes) with 
sparse spreading hairs and thus appearing less hairy than the preceding species; cauline 
leaves lanate, 3-lobed, the lobes 1 5-30 (35) mm long, 1 .5-3 mm wide; bracts as long as 
tlie flowers, rarely slightly longer; flowers solitary in leaf axils, often forming rather 
dense spikelike terminal inflorescences; pedicels to 2 mm long; calyx campanulate, 4- 
5 mm long, hairy only in upper part, usually glabrous below, with prominent nerves and 
acutely triangular teeth to 2-2.5 mm long, shorter than corolla-tube by 2-3 mm; corolla 
16-20 mm long, lemon-yellow with purple stripes along nerves and spots on lower Up; 
lower lip broadly obcordate, sometimes subreniform, broadly emarginate; lateral lobes 
broad, obtuse; corolla filaments and style less hairy than in preceding species; nutlets 
oblong, 2.5-3 mm long, reticulate at apex, transversely rugose below, the often oblique 
anastomoses between the wrinkles relatively numerous. Of frequent occurrence are an- 
dromonoecious specimens with bisexual flowers at ends of branches and sterile male 
flowers below. May— September. (Plate 1. Figure 1.) 

Steppes, steppe and stony slopes and taluses, chalky outcrops, wastelands, fields, road- 
sides. - European part: U. Dnp.. M. Dnp.. V.-Don, Transv., U. Dns., Bes., Bl., Crim., L. 
Don,L.V.; Caucasus: universal; Centr. Asia: Mtn. Turkm. Gen.distr.: Centr. Eur., 
Med., Bal.-As. Min.. Arm.-Kurd., Iran. Described from near Izyum. Type at Kharkov 
University. 

Note. Very close to A. chia; constantly confused with either A. chia or A. cha- 
maepitys because of its intermediate features. Yet, the distribution area and the aggre- 
gate of morphological characters quaUfy it as a separate species. A. chia and A. pseu- 
dochia occur together in southern Italy and on the Aegean and Black Sea coasts. 
A. pseudochia has apparently spread from there to eastern and northeastern Europe, 
which is today its principal distribution area. 



16 



23 




PLATE I, 1 - Ajuga pseudochia Schost., general aspect, flower, nutlet; 2,2a - A. chia Schreb.^ 
general aspect, flower, nutlet; 3,3a- A. chamaepitys (L.) Schreb., general aspect, flower, nutlet. 



17 



Visiani (Flora dalmatica, II, 1847, 222) described a form from Dalmatia resembling 
A. chia and A. chamaepitys, and called it A. chamaepitys var. grandiflora. Judg- 
ing from his description, this plant is in no way distinguished from A. pseudochia. 
Specimens from Sicily determined as A. chamaepitys (L.) Schreb. var. grandiflora Vis. 
have also proved to be A. pseudochia. 

25 3. A. chamaepitys (L.) Schreb. PI. vert, unilab. (1774) 24; Benth. Lab. gen. et sp. 699, 
p.p.; DC.Prodr. XII,601,p.p.; Ldb. Fl. Ross. Ill, 449, p.p.; Shmal'g. Fl. II, 346, p. p. 
- Teucrium chamaepitys L. Sp. pi. (1753) 562. - Bugula chamaepitys Scop. Fl. 
earn. (1772) 417. - Chamaepitys trifida Dumort. Florul. Belg. (1827) 42. - Ch. vul- 
garis Link, Handb. (1829) 453. - Ic: Rchb. Ic. Fl. Germ. XVIII, tab. 1235; N. Pop. in 
Tr. Bot. sada Yur'evsk. univ. XIV, 24 - fruit. 

Annual or biennial; stems often numerous, 5-20 cm high, procumbent or ascending, 
leaves, except for the radical, deeply 3-partite, with linear entire lobes, the middle one 
often broader than the others; stems and leaves covered with soft white hairs; bracts 
longer than flowers, resembling cauline leaves; flowers soUtary in leaf axils, yellow with 
purple spots and stripes on lower lip and corolla-tube; pedicels 1-1 .5 mm long; calyx 
campanulate, 5-6 mm long, equaling the corolla-tube or not more than 1 mm shorter, 
covered with long white hairs, the acutely triangular teeth to 3 mm long; corolla 10- 
16 mm long, only 2-3 times the length of calyx, pubescent outside; middle lobe of lower 
Up obcordate, gradually tapering to claw, slightly 2-fid, with a broad apical slit; lateral 
lobes broadly lanceolate, ca. 2 mm long, obtuse: filaments and style with scattered long 
spreading hairs; nutlets ovoid, rarely somewhat oblong-ovoid, to 3 mm long, the entire 
surface reticulate-rugose. June. (Plate I, Figure 3.) 

Dry, often calcareous slopes; mostly wastelands, fields and meadows. - European 
part: U. Dnp. (western part), U. Dns., Bes. Gen. distr.: Centr. and Atl. Eur., Med. 
(western part), Bal.-As. Min. (Bal.). Described from Portugal. Type in London. 

Note. A. chamaepitys has a typically West European distribution. It extends east- 
ward only to the upper parts of the Dniester and Pripet. A single specimen from the vici- 
nity of Zemlyansk in the Voronezh region, preserved in the herbarium of the Botanical 
Institute of the Academy of Sciences of the USSR, was determined by B.M. Kozo-Polyan- 
skii as A. chamaepitys. In fact it comes closest to A. chamaepitys except for the 
nutlets which are more coarsely alveolate compared with the type. More collections are 
needed to determine the eastern boundary of A. chamaepitys. 



Subsection 2. Pseudophleboanthe Pissjauk. - Sect. Phleboanthe Briq. in Pflan- 
zenfam. IV, 3 (1897) 210, p.p. - Leaves entire; corolla-tube closed, upper lip broadly 
and slightly 2-lobed; corolla covered with jointed hairs. 

26 4. A. salicifolia (L.) Schreb. PL vert, unilab. (1774) 26; M. B. Fl. taur.-cauc. II, 34; 
III, 388; Benth. Lab. gen. et sp. 699; DC. Prodr. XII, 599; Ldb. Fl. Ross. Ill, 449; Boiss. 
Fl. or. IV, 800. - N. Pop. in Mat. Fl. Kavk. IV, 3, 19. - A. oblongata Schmalh. Fl. II 
(1896) 346, p. p. quoad pi. taur. (non M. B. 1. c. III. 388, quae est pi. transcaucas.). - 



18 



27 



A. rhodopea Velen in Sitzber. bohm. Gesellsch. Wiss. (1892) 19. - Teucrium salici- 
folium L. Mantissa (1767) 80. - Ic: Schreb. Ic. et descr. I, 17, tab. 9; Sibth. et Sm. Fl. 
Graec. VI,tab. 526. 

Perennial, strongly rhizomatous, to 20-30 cm high; stems numerous, often woody at 
base, simple or rarely branching at base, ascending or spreading, in lower part slender, 
somewhat nodose, subterete, glabrous, in upper part slightly 4-angled, covered with short 
crisp hairs, sometimes almost tomentose; leaves distant, short-petioled, broadly lanceolate 
or lance-elliptic, 25-30 mm long, 10-12 mm wide, acute, entire, rarely 2-3-toothed in up- 
per part, with short stiff hairs above and beneath, sometimes almost strigose; bracts resem- 
bling the leaves, exceeding the flowers; flowers to 25 mm long, yellow with purple stripes 
on corolla-tube and on lateral lobes of lower lip; pedicels to 3 mm; calyx campanulate, 
8- 10 mm long, not more than half the length of the corolla-tube, glaucous all over with 
short appressed hairs, the teeth lanceolate, obtuse, upright, equaling or exceeding the tube, 
the upper tooth often smaller than the others; corolla nearly twice as long as calyx, also 
finely appressed-hairy, the upper lip with 2 obtuse short lobes, the lower with long lance- 
olate lateral lobes and narrowly unguiculate deeply cut middle lobe; corolla-tube broadly 
campanulate at apex; filaments and style pubescent; nutlets oblong, 3-3.5 (4) mm long, 
reticulate-rugose. May-July. (Plate II, Figure 2.) 

Steppes, stony slopes. — European part: Crim. Gen. distr.: Bal.-As. Min., Arm.-Kurd. 
Described from Southwest Asia. Type in London. 

5. A. oblongata M. B. Fl. taur.-cauc. Ill (1819) 388; Benth. Lab. gen. et sp. 698; in 
DC. Prodr. XII, 599; Ldb. Fl. Ross. Ill, 499; Boiss. Fl. or. IV, 801 ; Shmal'g. Fl. II, 346, 
p.p. (quoad pi. transcaucas.); N. Pop. in Mat. fl. Kavk. IV, 3, 18. — A. salicifolia Stev. 
in Mem. Soc. Nat. Mosc. Ill (1812) 265, non Schreb. 

Perennial, 10-30 (50) cm high, patent -villous; stems stout, often with sturdy branches; 
leaves densely covering the stem, narrowly lanceolate or oblong-elliptic, obtuse, entire, 
2-4 cm long, 5-10 mm wide, tapering to base, subsessile, the terminal leaves generally 
shorter than flowers; flowers yellow, on pedicels to 1 mm long, in a rather dense spikelike 
inflorescence; calyx rounded-ovoid or campanulate, 5-6 mm long, 1/2 to 1/3 the length 
of corolla-tube, with sparse long hairs confined to upper part, the teeth ca. 3 mm, narrow- 
ly lanceolate, acute, the uppermost shorter than the others; corolla 20-25 mm long, yel- 
lowish,, without purple spots or stripes, thin (the tube almost scarious), with long white 
hairs and short glands on the outside; corolla-tube infundibular; upper lip slightly 2-lobed; 
lower lip large, with lanceolate obtuse lateral lobes and reniform retuse middle lobe; fila- 
ments and style with scattered hairs; nutlets oblong, reticulate-rugose. May —July. 
(Plate II, Figure 4.) 

Dry steppes. — Caucasus: E. Transc. Gen. distr.: Med. (east), Iraq. Described from 
the estuary of the Kura River. Type in Leningrad. 

6. A. chamaecistus Ging. ex Benth. Lab. gen. et sp. (1835) 698; in DC. Prodr. XII, 
600; Briq. in Pflanzenfam. IV, 3a, 210; Bge. Lab. Pers. 84; Boiss. Fl. or. IV, 801; 
Grossg. Fl. Kavk. Ill, 281; Bornmull. in Bull. Herb. Boiss. 8, 2, 121. 

Low subshrub or shrub, with large root and dense divaricate or ascending branches, 
loosely cespitose; stems and leaves covered with coarse spreading, sometimes bristlelike 



19 



hairs; stems slender, 1.5-2 (3) mm in diameter, white, sturdy, nodose; sterile shoots often 
leafless, woody, nearly spinous; leaves variable: on sterile shoots lanceolate or lanceolate- 
linear, on the fertile oblong-elliptic or ovate-elliptic, obtuse, entire, sessile, always attenuate 
at base, sUghtly amplexicaul, 20-35 mm long, 4-10 mm wide, with thin revolute margins, 
often canescent, remote in lower part of stem; bracts resembling cauline leaves, about 
equaling the flowers; flowers solitary, bright purple, 18-25 (not more than 30) mm long; 
pedicels 2-3 mm long; calyx tubular-campanulate, densely covered with stiff appressed 
hairs, almost lanate, gray, to 8 mm long; teeth triangular-lanceolate, as long as the tube, 
not accrescent, the uppermost slightly shorter than the others; corolla (except for lower 
lip) densely hairy, the thin subscarious tube purple-striped; upper lip with scarcely visible 
obtuse lobes; lower lip large, the lateral lobes broad, rounded, spreading at an obtuse 

28 angle, the middle lobe 10-13 mm wide, short-glandular deeply cut, broadly unguiculate 
with rounded, slightly incised-dentate lobules; filaments sparsely short-glandular, with a 
tuft of long hairs below the anthers; style glabrous; nutlets 5-6 mm long, 2 mm wide, 
longitudinally reticulate-rugose, with very large areola. May -June. (Plate II, Figure 1.) 

Slopes of hills, stony slopes of mountains up to 2000 m. — Caucasus: S. Transc. Gen. 
distr.- '^""", -Kurd., Iran. Described from Iran. Type in Paris? 

7. A. turkestanica (Rgl.) Briq. in Bull. Herb. Boiss. 2 (1894) 71 1 et in Pflanzenfam. IV, 
3a, 210; Lipsk. in Tr. Bot. sada, XVIII, 1, 107. — Rosenbachia turkestanica Rgl. in 
Tr. Bot. sada, IX (1886) 613. - A. chamaecistus B. Fedtsch. Rast. Turk. (1915) 671, 
non Benth. - Ic: Tr. Bot. sada, IX (1886) tab. X, fig. 21, a-f, flowers and fruit. 

Subshrub with a robust root; stems 40-50 (60) cm long, stout, 3-5 mm in diameter, 
pale brown, sometimes reddish, rarely more or less whitish below, glabrous, grayish, in 
upper part finely and softly appressed-hairy like the leaves; leafless, woody and spinous 
branches usually absent; sterile shoots leafy above; leaves larger than in the preceding 
species, (30) 45-60 mm long, (12) 14-18 mm wide, oblong-elliptic or obovate, sometimes 
rhombic, only on short sterile branches occasionally oblong-lanceolate, entire or rarely 
with 2-4 apical teeth, tapering at base; lower leaves short-petioled, the upper sessile, 
amplexicaul; pubescence of leaves and other parts sparser than in the preceding species; 
flowers 25-40 mm long, purple; pedicels 3-4 mm long at flowering, later 6 mm; calyx 
campanulate or narrowly so, (8) 10-12 mm long at flowering, 13-16 mm in fruit, covered 
all over with fine short appressed hairs, glaucous, the teeth narrowly lanceolate, nearly as 
long as calyx-tube, accrescent in fruit to nearly twice their length and becoming broadly 
lanceolate, foliaceous; corolla-tube long, broadly infundibular, finely appressed-hairy, 
l-V-h times as long as calyx; upper lip slightly developed, with 2 obtuse, hardly discern- 
ible lobes; lower lip very large, slightly glandular, the lateral lobes broadly lanceolate or 
ovate, diverging at an acute angle, deeply bipartite, 12-15 mm wide, the middle lobe long- 
unguiculate, the lobules rounded, faintly undulate-dentate; filaments ribbon-shaped in 
lower part, glabrous like the style; nutlets oblong-ovoid, 7-9 mm long, 3 mm wide, thinly 

29 longitudinally reticulate-rugose, with very large areola. April— July. (Plate II, Figure 3.) 

Stony and gravelly mountain slopes, rocks and taluses, to 2500m. — Centr. Asia: Pam.-Al. 
(west and southwest). Endemic. Described from Tadzhikistan. Type in Leningrad. 

Note. This species is the more northern, mesophyllous race of A. chamaecistus, to 
which it is allied. Inspite of Borimiiiller's objections to the segregation of A. turkestanica 



20 



(Bull. Herb. Boiss. 2 ser., 8, 1908, 121), this species is very clearly distinguished from 
A. chamaecistus, even from the "luxuriantly developed specimens" from Northern and 
Western Iran, to which it is referred. We had at our disposal a specimen of A. cham ae- 
cistus collected by Bornmiiller on 12 V 1902 between Resht and Kazvin (spurs of Elburz 
Range), with critical notes attached, as well as a specimen collected on 1 1 V 1904 near 
Kermanshah and determined by Bornmiiller. Study of these plants showed that in respect 
of vesture, structure of flowers (especially calyx) and character of branches, they belonged 
to A. chamaecistus and were readily distinguishable from the Pamir-Alai forms. More 
collections are needed from Southern Transcaucasia, Talysh and the western and south- 
western regions of Pamir-Alai. 



Subsection 3. Phleboanthe (Boiss.) Briq. in Pflanzenfam. IV, 3a (1897) 210, p. p. 
(quoad typ.); Boiss. Fl. or. IV (1879) 804 (pro sect.). — Corolla-tube closed from upper 
lip to nearly middle, otherwise as in preceding subsection. 

8. A. laxmannii (L.) Benth Lab. gen. et sp. (1835) 697; in DC. Prodr. XII, 599; Ldb. 
Fl. Ross. Ill, 448; Shmal'g. Fl. II, 346. - Teucrium laxmanni L. Syst. veg. (1784) 
526; Georgi, Beschr. Russ. R. Ill, 5, 1077; M. B. Fl. taur.-cauc. II, 35; III, 389. - T. iva 
Georgi, 1. c. 1076 (non L.). — T. pseudiva Giildenst. Reis. Russl. et Cauc. (1787) 140. — 
Phleboanthe laxmanni Tausch in Bot. Zeitschr. I (1828) 323. — Ic: Rchb. Ic. Fl. 
Germ. XVIII, tab. 1236 (sub Phleboanthe laxmanni). 

Perennial, with woody branching root; stems stout, to 8mm in diameter, simple, 
numerous, densely leafy, erect or ascending, to 20-50 cm long, patent-villous, more 
copiously so at nodes; leaves oblong- or oval-elliptic, obtuse, 35-45 (55) mm long, 10-12 
(30) mm wide, sessile, sUghtly amplexicaul, entire, on sterile shoots lanceolate, long-attenu- 
ate at base, at apex rarely crenate-serrate, grayish with appressed-lanate hairs on both sides; 
bracts leaflike, exceeding the flowers; verticillasters two-flowered, remote or crowded in 
30 spikelike terminal inflorescence; flowers large, to 30-35 mm long, yellow, with dense net- 
work of purple nerves; pedicels 3-4 mm; calyx campanulate, 10-15 mm long, wooUy- 
canescent with clavate hairs; calyx-teeth lanceolate or oblong, obtuse, rarely subacuminate, 
nearly the length of the tube (ca. 5 mm long), the uppermost tooth sometimes slightly 
smaller than the others; corolla covered with clavate hairs and on lower Up densely short- 
glandular; tube open; lobes of upper lip small, lanceolate; lower lip very large, reclinate, 
the lateral lobes broadly oval, the middle lobe long-clawed, 2-fid, with broad oval lobules; 
filaments and style extending to the tip of lateral lobes of lower lip; style glabrous, with 
thickish stigma lobes; filaments ribbonlike at base, covered with short clavate hairs; nut- 
lets obovoid, light brown, 4-4.5 mm long, 1.5-2 mm wide, reticulate-rugose, with large 
areola. May -July. (Plate II, Figure 5.) 

Steppes, steppe and chalky slopes, coppices, margins of broadleaved forests in forest 
steppe and steppe regions. — European part: U. Dnp., M. Dnp., V.-Don, U. Dns., Bes., Bl., 
Crim., L. Don; Caucasus: Cisc, E. Transc. (?). Gen. distr.: Centr. Eur., Bal.-As. Min. 
Described from cultivated specimens. Type in London. 



21 



Section 2. Bugula Benth. Lab. gen. et sp. (1835) 692; in DC. Prodr. XII, 525; Briq. 
in Pflanzenfam. IV, 3a, 210; Maxim, in Mel. biol. XI, 808. — Verticillasters 6- to many- 
flowered; corolla blue, rarely pink, purple or white, not yellow. 



Subsection 1. Genevenses Maxim, in Mel. biol. XI (1883) 809 (pro serie). - Leaves 
tapering at base; corolla-tube straight; stamens and style exserted, 

9. A. reptans L. Sp. pi. (1753) 561; Benth. Lab. gen. et sp. 694; in DC. Prodr. XII, 
595; Ldb. Fl. Ross. Ill, 446; Shmal'g. Fl. II, 345; Syreishch. Fl. Mosk. gub. Ill, 115. - 
N. Pop. in Mat. Fl. Kavk. IV, 3, 27; Fl. Yugo-vost. VI, 134; Kryl. Fl. Zap. Sib. IX, 2992. 
- A. repens Host, Fl. Austr. II (1831) 119; Giildenst. ex Ldb. 1. c. 447. - A.pyramida- 
lis Sobol. ex Rupr. Beitr. Pflanzenk. Russ. R. IV (1847) 68. - A. alpina Fries, Mant. 3 
(1842) 54, non Linn. - A. barelieri Ten. Fl. Nap. (1842) tab. 240, fig. 2. - A. vulgaris 
ssp. reptans Rouy, Consp. Fl. Franc. (1927) 206, - Bugula reptans Scop. Fl. Carniol. 
(1760) 447. - B. densiflora Ten. 1. c. tab. 239, fig. 2. - Teucrium reptans Crantz, 
31 Stirp. Austr. ed. I (1769) 25 1. - Ic: Rchb. Ic. Fl. Germ. XVIII, tab. 1234, Fedch. and 
Fl. Fl. Evrop. Ross. 800; Syreishch., op. cit. Ill, 1 14; Maevsk. Fl. 608. 

Perennial, with rooting stolons; stems simple, 15-35 cm long, covered on two faces with 
fine, appressed hairs, these sometimes interspersed with erect articulate hairs; radical 
leaves long-petioled, spatulate or spatulate-obovate, broadly sinuate-dentate; lower cauhne 
leaves few, oblong or ovate, sessile, gradually passing into bracts; bracts entire, ovate, 
sometimes slightly crenate, the lower longer than the flowers, the upper shorter; flowers 
azure, blue, sometimes white or pink; verticillasters 6-8-flowered, distant below, forming 
above a rather dense spikelike inflorescence; calyx glandular hairy, almost villous at apex, 
the lanceolate teeth longer than, rarely as long as the tube; corolla densely glandular hairy 
outside, the tube not more than half as long again as the calyx, the upper lip deeply 2- 
lobed, the middle lobe of lower lip rounded, slightly emarginate, the lateral lobes broadly 
ovate; stamens and style exserted; filaments densely covered with long articulate and 
short clavate hairs; nutlets globose, pale brown, ca. 2.5 mm long, finely reticulate-rugose. 
May -July (August). 

Wet meadows, forests, forest glades, coppices, wooded slopes. Ascending into high 
mountains; rare in southern regions. - European part: Kar.-Lap. (south), Lad.-Ilm., 
Bah., Dv.-Pech., V.-Kama, U. Dns., U. Dnp., L. Don, Transv., L.V. (delta of Volga), Crim.; 
Caucasus: Cisc, Dag., W. and E. Transc, Tal. Gen.distr.: Scand., Atl. and Centr. Eur., 
Med., Bal.-As. Min., Arm.-Kurd., Iran (northern part). Cultivated in N. Am. Described 
from Europe. Type in London. 

Note. This widespread species is by no means homogeneous. The following fornis 
(apparently ecological) have been described for the USSR: 1) var. alpina N. Pop. in Mat. 
Fl. Kavk. IV, 3 (1916) 30 (A. alpina Vill. non Fries) - high-mountain form characterized 
by a reduced stem with a rosette of radical leaves and a short, dense inflorescence; 2) var, 
simplex Kauffm. Mosk. Fl. (1866) 389 - with very short stolons (sometimes obsolescent), 
densely hairy stem and large radical leaves. Both forms grow on dry slopes and in dry scrub. 

There are also colored fomis: var. albitlora Tin. (Fl. Luxemb. 1836, 291) and var. 
rosciflora Wildem. et Durand (Prodr. Fl. Belg. Ill, 1899, 640). For hybrids, see Note to 
A, genevensis. 

22 



32 10. A. shikotanensis Miyabe et Tatewaki in Transact. Sapporo Nat. Hist. Soc. XIV 

(1935-1936)7. -Ic: I.e. 7, fig. 3. 

Perennial, stoloniferous; stems ascending, 15-20 cm long, with patent white hairs on 
two faces, the hairs usually of two kinds: tangled subappressed-articulate and erect- 
glandular; radical leaves petiolate, spatulate, round-tipped, subentire, rarely broadly 
incised with scattered coarse hairs above, subglabrous beneath, with petioles 5-8 cm long; 
lower cauline leaves petiolate, tapering above, upper leaves and bracts sessile, oblong or 
oblong-ovate, subentire, round-tipped, with scattered woolly hairs above, glabrate beneath, 
sUghtly colored; verticillasters 5-7-flowered; calyx campanulate, with 5 subequal, triangu- 
lar-lanceolate hairy teeth, the tube subglabrous; corolla ca. 13 mm long, with scattered 
long hairs and short glands, the tube straight, the upper lip with 2 ovate-triangular lobes, 
the lower large, rarely recurved, with obcordate middle lobe and oblong truncate lateral 
lobes; stamens and style slightly exserted; stamens pubescent; style glabrous, the stigma 
with sUghtly unequal lobes. 

Grass patches along seacoasts and at foot of mountains. — Far East: Sakh. (Kurile 
Islands). Described from Shikotan Island. Type in University Herbarium, Hokkaido. 

11, A.genevensisL. Sp. pi. (1753) 561; Benth. Lab. gen. et sp. 694; inDC.Prodr. 
XII, 596; Ldb. Fl. Ross. Ill, 448; Shmal'g. Fl. II, 345; Syreishch. Fl. Mosk. gub. Ill, 1 15; 
Fedch. and Fl. Fl. Evrop. Ross. Ill, 800; N. Pop. in Mat. Fl. Kavk. IV, 3, 22. - A. alpina 
L. Mantissa (1767) 80; Falk, Beitr. Kenntn. Russ. R. 204; M. B. Fl. taur.-cauc. II, 33; III, 
388. - A. pyramidalis M. B. 1. c. II (1808) 32 et auct. plur. florae Ross. med. austr. et 
cauc. (non L.); Boiss. Fl. or. IV, 799, p. p. (quoad pi. Caucas.). — A. alpestris Dumort. 
Florula Belg. (1827) 42. - A. folio sa Tratt. Arch. Gew. I (1829) 25. - A. latifolia et 
rugosa Host, Fl. Austr. II (1831) 1 19. - A. montana Dill, ex Rchb. Fl. Germ. exc. 
(1830-1832) 314. - A. cryptostylon Lagreze-Fossat ex Nym. Consp. Fl. Eur. (1881) 
567. — A. vulgaris ssp. foliosa, alpina, genevensis Rouy, Consp. Fl. Franc. (1927) 
206. - Bugula genevensis Mill. Gard. Diet. ed. VIII, No. 4. — B. decumbens Mill. 
35 1. c. No. 2. - B. tomentosa Gilib.PI. rar. et comm. Lithuan. (1785) 17 et Fl. Lithuan. 
in Delect. Opusc. Bot. II (1793) 323. - B. alpina All. Fl. Pedem. I (1785) 45. - Teu- 
crium genevense Crantz, Stirp. Austr. ed. I (1769) 253. - Ic: Rchb. Ic. Fl. Germ. 
XVIII, tab. 1234; Syreishch. op. cit., 1 15. 

Perennial, non-stoloniferous with bundles of strongly branching lateral roots; stems 
10-40 (50) cm, rarely branching at base, villous-hairy (only on two faces in var. alpes- 
tris (Dum.) Beck.), sometimes subglabrous at base; radical leaves distant, oblong-spatulate 
or obovate, strongly crenate-dentate, sometimes almost entire, 4-12 cm long, 2-5 cm wide, 
tapering to petiole mostly marcescent; lower cauline leaves oblong, in upper part remotely 
crenate-dentate, cuneate, sessile, gradually passing into bracts; bracts coarsely 3-toothed, 
often 3-lobed, ovate or broadly ovate, the upper usually shorter than the flowers (longer 
in var. longibracteata Bizzoz.), oftei colored; all leaves covered on both sides with 
more or less articulate hairs and sometimes also with short bristles; verticillasters 4-6 (8)- 
flowered, crowded in a spikelike terminal inflorescence, remote below; flowers 12-18 mm 
long, sessile; calyx lanate in upper part, shghtly glandular, subglabrous at base, 5-6 (7)mm 
long, 1/2 to 2/5 as long as corolla-tube, the subequal triangular-lanceolate teeth shorter 
than the tube; corolla blue, rarely pink or white, sparingly pubescent outside (especially 



23 



(33) 




PLATE II. 1 - Ajuga chamaecistus Ging., la) flower; 2 - A. salicifolia (L.) Schreb., part of 
flowering branch; 2a) flower; 3 - A. turkestanica (Rgl.) Briq., part of flowering branch; 3a) flower; 
3b) fruiting calyx; 4 - A. oblongata M. B., part of flowering branch; 4a) flower; 5 - A. laxman- 
nii (L.) Benth., flower (one of the calyx teeth bent). 



24 



the tube); lobes of upper lip small, truncate; middle lobe of lower lip broad, recurved; 
stamens and style much exserted; filaments pubescent or subglabrous; nutlets to 3 mm 
long, globose, dark brown, diffusely patent-hairy, prominently netted-rugose. April— July 
(up to September). 

Meadows, scrub, forests, small woods, river valleys. — European part: Bait., Lad.-Ilm., 
U.V., U. Dnp., V.-Kama, M. Dnp., V.-Don, Transv., U. Dns., Bes., Bl., Crim., L. Don; 
Caucasus: universal. Gen. distr.: Europe, except for the Arctic, Med., Bal. -As. Min., 
Arm.-Kurd. Described from Switzerland. Type in London. 

Note. This species is remarkable for its variabiHty and heterogeneity within its distri- 
bution area (see synonymy). Further studies are required to determine the extent of sea- 
sonal variability of its individuals and the ecological adaptations of the various forms. 
Greatest variabihty is to be found in size of radical leaves and bracts, vesture of the stem, 
and color of corolla. In the typical form, the stem is hairy all round, often villous, the 
radical leaves exceed the lower stem leaves, and the bracts are about as long as the flowers. 
Other known forms are: (1) var. excelsa Lindem. Florula Elisabethgr. (1868) 120 (A. 
f oliosa Tratt.), with higher stems (to 40-50 cm) and coarsely toothed leaves which are 
2-3 times as long as in the typical form; the lower 2-6 pairs of bracts are also large and 
greatly exceed the flowers; mostly in forests, forest margins and coppices; (2) var. ela- 
tior (Fries) Briq. in Hegi, III; Fl. V, 2544; Syreishch. Fl. Mosk. gub. Ill, 115; Fl. Yugo- 
vost. VI, 134, close to the type, characterized by small radical leaves (the size of lower 
stem leaves), acutely 3-lobed lower and middle bracts that exceed the flowers and diffuse 
pubescence of the stem; (3) var. serotina Kosh. et Zing. Fl. Tul'sk. gub. (1880) 91, a 
late-flowering form (end of July-August), with very large, long-petioled radical and lower 
stem leaves. 

Forms representing color variations occur throughout the region: f. roseiflora Cho- 
roschk. (in Tr. Bot. sada Yur'evsk. univ. VIII, 1908, 17) and f. albiflora Syreischt. (Fl. 
Mosk. gub. Ill, 1910, 115). 

There are records of hybrids A. genevensis L. X A. reptans L., in varying degree 
resembling one or the other parent species: X A. genevensis var. stolonifera Zing, (in 
ByuU. Mosk. obshch. ispyt. prir. 1881, No. 2, 329), with short stolons, and X A. hybrida 
Kern, (in Oesterr. Bot. Zeitschr. XXIV, 1874, 382), similar to A. reptans but non- 
stoloniferous, with dentate radical leaves persistent in flowering. 

12. A. multiflora Bge. Enum. pi. Chin. bor. (1831) 51 ; Kom. Fl. Man'chzh. Ill, 334; 
Kom. and Alls. Opred. rast. Dal'nevost. kraya, II, 892. - A. genevensis auct.: Maxim. 
Prim. fl. Amur. (1859) 221 et in Mel. Biol. (1883) 815; Korzh. in Tr. Bot. sada, XII, 
380, non L.; Forb. et Hemsl. Ind. Fl. Sin. II, 315, p.p. - A. amurica Freyn in Oesterr. 
Bot. Zeitschr. (1902) 408. - Ic: Kom. and Alls., op. cit. 892, Plate 20. 

Perennial, non-stoloniferous, with a short main root and bundles of large lateral roots; 
stems erect, robust, solitary or 2-3 developing from one root, simple, 8-25 cm long, white- 
villous with long articulate hairs; radical leaves with petiole to 6 cm, oblong or oblong- 
elliptic, 5-10(12) cm long, 2-3 (4) cm wide, obtuse, entire or obscurely incised, glabrous 
or with remote short hairs; cauline leaves sessile, broadly lanceolate or acuminate-ovate, 
2-4 cm long, 1-3 cm wide, obscurely dentate-crenate in upper half, appressed-lanate (es- 
pecially at base), longer than the internodes, gradually smaller toward summit; upper 



25 



37 bracts white-villous, ovate or oblong-ovate, entire or sparsely toothed at apex, longer or 
(more often) shorter than the flowers; flowers subsessile, crowded in verticels of 8-10, 
bright blue; calyx 5-6 mm long, densely long-hairy, the 5 equal narrow, triangular-pointed 
teeth longer than the tube; corolla soft-hairy outside, 12-15 mm long, the tube twice as 
long as the calyx; lower lip large, 8-9 mm long, the obcordate middle lobe to 7 mm wide; 
upper lip faintly 2-lobed; stamens and style exserted; filaments pubescent; nutlets 
2.5-3.5 mm long, pale brown, prominently netted-rugose, with short scattered hairs. 
April— June. 

Meadows, meadow slopes, clearings, scrub. East Siberia: Dau.; Far East: Ze.-Bu.,Uss. 
Gen. distr.: China (Manchuria). Vicariate races in S. China and Japan. Described from 
near Peking. Type in Paris. 

13. A. pyramidalis L. Sp. pi. (1753) 561; Benth. Lab. gen. et sp. 693; in DC. Prodr. 
XII, 596; Ldb. Fl. Ross. Ill, 447, p. p. (quoad pi. Ross, septentrion.); Shmal'g. Fl. II, 
345; Fedch. and Fl. Fl. Evrop. Ross. Ill, 800. - A. nana Gilib. PI. rar. et comm. Lithuan. 
(1785) 17 et Fl. Lithuan. in Delect. Opusc. Bot. II (1793) 323. - A. alpina Sut. Fl. Helv. 
II (1802) 2 and II (1822) 2, non L. nee Vill. (?). - A. vulgaris ssp. pyramidalis Rouy, 
Consp. Fl. Franc. (1927) 206. - Bugula pyramidalis Crantz, Inst. I (1766) 518. - 
Teucrium pyramidale Crantz, Stirp. Austr. ed. II (1769) 252. - Ic: Fl. Dan. (1765), 
tab. 185; Rchb. Ic. Fl. Germ. XVIII, tab. 1234. 

Perennial, non-stoloniferous, with short main root and a bundle of strong lateral roots; 
stems simple, erect, 10-30 cm long, villous all round with stiff, patent hairs, rarely glabrate 
below; leaves in radical rosette numerous, short-petioled, large, obovate or broadly spatu- 
late, entire or crenulate at apex, rough with articulate hairs; bracts broadly ovate, the ter- 
minal oblong-ovate, entire, rarely obscurely toothed, sessile, often violet, always exceeding 
the flowers; verticillasters mostly 6-flowered, borne from base of stem and forming a dense, 
4-angled inflorescence; flowers small, 10-13 mm long, sessile, pale blue; calyx subglabrous 
below, about half as long as corolla-tube, the teeth nearly as long as the tube; corolla vil- 
lous, the upper lip 2-lobed, the lower lip with small middle lobe; filaments and style slight- 
ly exceeding the lobes of upper Up; nutlets globose, ca. 1 .5 mm long, distinctly netted- 
rugose, with scattered patent hairs. May— June. 

38 Meadows in northern regions. — European part: Bait., Kar.-Lap. Gen, distr.: Scand., 
Centr. and Atl. Eur., W. Med. (mountains), Bal.-As. Min. (Balkans). Plant of northern 
meadows or Alps. Described from Sweden. Type in London. 

Note. A typical, arctic-alpine species erroneously reported from many parts of the 
middle belt and the Caucasus; in lowlands apparently confused with some forms of A. 
genevensis (e.g. with its fall forms) and in the Caucasus mostly with A. orientalis. 
Hybrids of A. genevensis L. X A. pyramidalis and A. pyramidalis X A. reptans 
occur in regions where the parent species grow together (mainly in mountains). 



Subsection 2. Orientales Maxim, in Mel. biol. XI (1883) 824 (pro serie). - Corolla- 
tube contorted; stamens and style included in corolla-tube. 

14. A. orientalis L. Sp. pi. (1753) 561; Benth. Lab. gen. et sp. 693; in DC. Prodr. 
XII, 596; M.B. Fl. taur.-cauc. II, 32; Ldb. Fl. Ross. Ill, 447; Boiss. Fl. or. IV, 800; C. 

26 



Koch in Linnaea, XVII, 301 et in Linnaea, XXI, 702; Shmarg. Fl. II, 344; N. Pop. in 
Mat. FI. Kavk. IV, 3, 30. - Bugula obliqua Moench, Meth. (1794) 382. - B. orien- 
talis Mill. Card. Diet. ed. VIII, No.5. - Ic: Kniphof, Herb. Viv. I tab. 21; Fiori et Paol. 
Ic. Fl. Ital. 362. 

Perennial, with slightly oblique creeping rhizome; stems erect or slightly ascending, 
simple, 10-20 (30) cm long, like the leaves white- or yellowish-lanate-villous, sometimes 
glabrescent in lower part; radical leaves persistent in flowering, oblong-elliptic or broadly 
ovate, tapering to petiole, dentate-crenate or crenate, sometimes with large apical teeth; 
bracts usually sessile, eUiptic, broadly elliptic or obovate, amplexicaul, often rugose, large- 
toothed or 3-5-fid, longer than, equal to or shorter than flowers (var. condensata Boiss.), 
the terminal ones often violet (var. aenesia Heldr.); verticillasters many -flowered, more 
or less remote below, crowded above in compact spikelike inflorescence, sometimes (var. 
condensata Boiss.) crowded from base of stem; flowers subse^sile; calyx 5-6 mm long, 
campanulate, lanate-villous (sometimes glabrate at base), the teeth narrow, acute, profusely 
hairy, more than twice as long as the tube; corolla 13-16 (18) mm long, with blue hmb 
(often faded in herbarium); tube almost white, glabrous, contorted so that the lower hp 
assumes dorsal position; upper lip with 2 large ovate lobes, lower lip with small slightly 
emarginate middle lobe and closely adjacent lateral lobes; limb of corolla sparingly lanate- 
villous outside, the tube slightly exceeding the calyx; nutlets globose-ovoid, ca. 2 mm long, 
reticulate-rugose, sometimes with scattered hairs. April— June. 

Exposed stony slopes of dry hills, extending to subalpine and alpine belt, in meadows, 
clearings and scrub. - European part: Bes., Crim.; Caucasus; Cisc, Dag., W., E. and S. 
Transc. Gen. distr.: Med., Bal.-As. Min., Arm.-Kurd., Iran. Described from Southwest 
Asia. Type in London. 



Genus 1241.* Teucrium** L. 

L. Sp. pi. (1753) 562. - Scorodonia Adans. Fam. II (1763) 188. - Scordium Gilib. Fl. Lithuan. I 
(1781) 95. - Chamaedrys S. F. Gray, Nat. Arr. Brit. pi. II (1821) 369. - Monochilon Dulac, H. 
Hautes-Pyren. (1867)405. - Botrys Fourr. in Ann. Soc. Linn. Lyon, n.s. XVII (1869) 138. 

Flowers distinctly pediceled, sometimes sessile, in 1-3 (lO)-flowered cymose inflores- 
cences, borne in the axils of undifferentiated or slightly differentiated bracts, and forming 
a terminal inflorescence, this mostly secund, racemiform or spiciform, rarely capitate or 
umbellate; calyx tubular or campanulate, often curved, 10-nerved, with 5 subequal trian- 
gular teeth or with an enlarged upper tooth and thus 2-Hpped; corolla mostly deciduous, 
the tube short, included, rarely slightly exserted, without a ring of hairs inside, often hairy 
outside; upper hp seemingly absent, being cleft in two with the two halves shifted to the 
lower lip which thus appears 5-lobed; middle lobe large, rounded or oblong, entire or 
finely toothed, mostly concave, generally recurved; lateral lobes smaller, oblong, often 
also recurved or straight, divaricate; stamens 4, ascending, often long-exserted, didynamous, 

* Treatment by S.Y. Yuzepchuk. 

** From the Greek teukrion, name given by Theophrastus and Dioscorides to certain species of this 
genus; from the legendary Teucer (Greek Teukros, Latin Teucrus or Teucer), credited with discovery 
of the plant's healing properties. 



27 



the small divergent anther-ceUs confluent at apex; style with 2 short subequal branches; 
nutlets obovoid, tapering at base, reticulate or rugose. Perennial, very rarely annual herbs, 
subshrubs or shrubs, often with an aromatic, rarely disagreeable, odor. Leaves usually 
short-petioled, entire, crenate-dentate or pinnatilobate or parted. 

A genus of cosmopolitan distribution, but characteristically Mediterranean. It com- 
prises ten weU differentiated sections. 

40 1. Verticillasters forming racemiform or spiciform inflorescences; flowers reddish, 

purple or blue (white only in albinos) 2. 

+ Flowers crowded in dense umbellate or capitate inflorescences, whitish or 

yellowish 18. 

2. Leaves deeply pinnatipartite 3. 

+ Leaves entire, crenate-dentate or rather deeply incised-crenate (nearly lobate). . 6. 

3. Perennials, with eglandulose hairs 4. 

+ Glandular-hairy annuals 7. T. botrys L. 

4. Stamens long-exserted 5. 

+ Stamens not exserted 3. T. parviflorum Schreb. 

5. Plant covered with short, rarely rather long, patent hairs .... 1. T. orientale L. 
+ Plant appressed-tomentose 2. T. taylori Boiss. 

6. Upper teeth of calyx broader than the lower pair (calyx 2-lipped); bracts linear- 
subulate; inflorescence narrowly cyUndric, spicate 4. T. hircanicum L. 

+ Upper teeth of calyx broader than the lower pair; bracts lanceolate; inflorescence 

compound, rather broad, racemose 4a. T. ussuriense Kom. (page 364). 

++ Calyx-teeth subequal 7. 

7. Stems herbaceous; all flowers shorter than bracts (plant smelling of garlic). . . 8. 
+ Stems more or less woody at base; all flowers, or at least the upper, longer than the 

bracts or scarcely shorter (aromatic plants) 9. 

8. Leaves 2-3 times as long as broad, not or slightly cordate at base, acutely crenate- 
dentate, short-hairy 5. T. scordium L. 

.+ Leaves to twice as long as broad, cordate at base, obtusely crenate, tomentose-lanate 
6. T. scordioides Schreb. 

9. Plants non-stoloniferous; stems and lower side of leaves canescent with very short 
tomentum 17. T. canum Fisch. et Mey. 

+ Plants often St oloniferous; stems and lower side of leaves with patent hairs . . 10. 

10. Inflorescence of moderate length or abbreviated; verticillasters more or less approx- 
imate; bracts gradually smaller toward summit, usually all more or less toothed, the 
upper concave 11. 

41 + Inflorescence markedly elongated; verticillasters very distant; bracts abruptly di- 

minishing in size, at least the upper entire, all flat 16. 

11. Calyx-teeth with long, commonly sparse hairs on margins, short hairs wanting or 
few 12. 

+ Calyx-teeth covered with very thick short hairs, interspersed with long ones on 

margins 14. 

12. Leaves v/ith comparatively narrow cuneate base, not very densely hairy beneath. . 
8. T. nuchense C. Koch. 



28 



+ At least lower leaves not narrowly cuneate, all leaves rather densely hairy beneath, 
' often nearly velutinous 13. 

13. Calyx-teeth narrow, long, acuminate; lower leaves cordate or truncate at base . . 
9. T. multinodum (Bordz.) Juz. 

+ Calyx-teeth short and broad, acute; lower leaves with broadly cuneate or notched 
base 10. T. trapezunticum (Reching. fil.) Juz. 

14. Leaves finely crenate-dentate, densely pubescent beneath, nearly velutinous; flow- 
ers pale purple 13. T. fischeri Juz. 

+ Leaves largely and irregularly crenate-dentate, not very densely hairy beneath, not 
velutinous; flowers darker 15. 

15. Inflorescence shorter than the leafy part of stem; bracts often green, rarely colored; 
flowers of medium size, purple 11. T. chamaedrys L. 

+ Inflorescence nearly as long as the leafy part of stem; bracts nearly black-purple; 
flowers larger, very dark purple 12. T. pulchrius Juz. 

16. Leaves finely and obtusely dentate; most bracts entire 17. 

+ Leaves largely and acutely dentate; only uppermost bracts entire 

16. T. excelsum Juz. 

17. Stems densely covered with long patent hairs; leaves densely velutinous-hairy be- 
neath; upper bracts entire, oblong-ovate; corolla very dark purple 

15. T. krymense Juz. 

+ Stems with comparatively short hairs; leaves not velutinous beneath; upper bracts 
entire lanceolate; flowers purple 14. T. syspirense C. Koch. 

18. Flowers rather large, 1-1.5 cm long; all leaves entire 19. 

42 + Flowers small, 5-8 mm long; leaves crenate-dentate 21. T. polium L. 

19. Stems appressed-tomentose 20. 

+ Stems patent-hairy 20. T. pannonicum Kern. 

20. Leaves densely tomentose, gray-green above; calyx densely or very densely tomen- 

tose outside, the teeth not ciUate or obscurely ciliate along margin 

18. T.jailaeJuz. 

+ Leaves dark green, with short scattered hairs above; calyx with appressed short 

hairs outside, the teeth sparsely short-ciliate along margin 

19. T. praemontanum Klok. 



Secfion 1. Teucris Benth. Lab. gen. et sp. (1835) 661. -Shrubs or (in USSR) sub- 
shrubs, with opposite, axillary, 1-3-flowered cymes forming a racemose or paniculate 
inflorescence; calyx campanulate, erect, with 5 subequal teeth; leaves in our species 
(subsection Incisae Briq.) deeply parted. 



Series 1. Orientalia Juz. - Subshrubs with pinnatisect leaves. Flower in loose pani- 
culate inflorescence; filaments long-exserted. 

1. T. orientaleL.Sp.pl. (1753)562; Ldb. Fl. Ross. Ill, 1, 442; Shmal'g. Fl. II, 347; 
N. Pop. in Mat. Fl. Kavk. IV, 3, 64. - Ic: Bot. Mag. XXXI, tab. 1279. - Exs.; GRF, 
No. 1129; Fl. cauc. exs. No. 167. 

29 



Perennial, with woody, strongly branching root; rhizome short, branching, usually 
woody; stems few or many, 10-50 cm long, slightly ascending at base or erect, straight 
or flexuous, distinctly 4-angled, rather densely clothed with short, rarely (especially in 
inflorescence) long hairs, sometimes thickly covered all over with long multicellular sUght- 
ly crisp hairs; branches opposite; leaves rather distant, 1-3 cm long, 1.2-4 cm wide, broad- 
ly rhombic, broadly ovate or suborbicular, deeply thrice pinnatisect, the lobes rather long, 
narrowly linear, sHghtly revolute, green or canescent, with short patent or appressed hairs 
above, rather densely covered beneath with short hairs, mainly along prominent nerves, 
sometimes both sides rather densely covered with stiff crisp hairs; petioles 0.1-1 cm long; 
inflorescence lax, paniculate or paniculate-corymbiform, its branches usually more or less 

43 elongate, often with short secondary branches; peduncles 1-3-flowered, longer than the 
small dissected bract and the calyx; calyx 3-5 mm long, short-campanulate, with both 
short and long scattered hairs, the teeth spreading, triangular or lanceolate, acute, carinate, 
often subglabrous, nearly as long as the tube or slightly shorter; corolla 1-1.7 cm long, 3-4 
times as long as the calyx, blue, the middle lobe elongate, concave, acute, short-hairy 
beneath, the upper 4 lobes ovate or lanceolate, acute; filaments to 1 .7 cm long, long- 
exserted, with short crisp hairs below; nutlets glabrous, rugose, with bloom of small, 
pellucid, granular glands. June -August. (Plate III, Figures 1 and 2.) 

Gravelly-stony or clayey mountain slopes with xerophytic vegetation, mountain pine 
forests, mountain meadows and alpine pasturelands, clayey and stony banks of rivers, 
sandy hills. - Caucasus: Cisc, Dag., E. and S. Transc. Gen. distr.: Bal.-As. Min., Iran. 
Described from the "East" (probably from Armenia). 

N ote. Forms with stems and leaves covered with thick and long, spreading hairs 
resemble T. orientale /3 villosum Benth. (in DC. Prodr. XII, 1848, 577). They are very 
rare in the USSR, where they occur mainly in Southern Transcaucasia; they have also 
been found in other areas (Dagestan and even Ciscaucasia), but are more widespread in 
Asia Minor and Iran. Even there they grow alongside the typical T. orientale; thus they 
have no separate distribution area and are not otherwise sufficiently distinct from the 
typical form (c.f. similar forms in: Celakovsky, Ueber einige Arten der Gattung Teucrium, 
Bot. Centralbl. XIV, 1883, 223). 

Economic importance. Occasionally cultivated as an ornamental. 

2. T. taylori Boiss. Diagn. Ser. I, 7 (1846) 61 ; id. in Fl. or. IV, 809; Grossg. Opred. 
rast. Kavk. 325 - T. pruinosum Grossh. Fl. Kavk. Ill (1932) 282, non Boiss. 

Perennial, rhizomatous, cespitose, densely gray-tomentose with very short appressed 
hairs (vesture resembling that of the preceding species); stems many, 8-30 cm long, erect 
from ascending base, shortly paniculate-branched in upper part; leaves 0.6-3 cm long and 
as wide, rounded-ovate or orbicular, gray on both sides, bipinnatisect nearly to base, the 
segments entire, very short, Unear, obtuse, revolute-margined, prominently veined beneath; 
peduncles loosely branching, 1-3-flowered; pedicels 2-3 times as long as calyx and bracts, 

44 the latter grayish like the pedicels; calyx 4-5 mm long, short-campanulate, 5-partite to 
middle, the teeth long-subulate from triangular base; corolla ca. 1 cm long, twice as long 
as calyx, blue; middle lobe mitriform, obtuse, short-hairy beneath; filaments long-exsert- 
ed as in T. orientale but (at least in forms with normal vesture) in lower half more dense- 
ly hairy and glandular than in the (otherwise similar) preceding species. June. 



30 



Stony places and slopes. — Caucasus: S. Transc. (near Erevan). Gen. distr.: Iran. 
Described from the foot of "Sabst-Buschom" mountain near Shiraz. Type in Geneva. 

Notes. 1) We have not seen the type of T. taylori Boiss. or any other authenticated 
material; the identification of the Erevan plant with this species is Grossgeim's. Some 
features of this plant, in particular the character of the vesture and the narrow teeth of 
the calyx, also occur in the next species. The plant is possibly of hybrid origin. 

2) In addition to T. orientale L. and T. taylori Boiss. the series Orientalia contains 
a wide array of distinct races, all of which, however, are to be found outside the USSR 
(Southwest Asia). One of them is the Iranian T. pruinosum Boiss., erroneously reported 
by Grossgeim for Armenia; it was confused with T. taylori Boiss. although it differs 
markedly from that species in habit and vesture. 



Series 2. Parviflora Juz. — Like the preceding series, but filaments scarcely exserted 
from corolla. 



3. T. parviflorum Schreb. PI. Vert. Unilab. gen. et sp. (1774) 31 ; Boiss. Fl. or. IV, 810; 
N. Pop. Mat. for Fl. Kavk. IV, 3, 6,9; Grossg. Fl. Kavk. Ill, 282. - Ic: Schreb. 1. c. tab. 
p. XXXI, No. 18. 

Perennial, rhizomatous; stems many, erect, herbaceous, 0.4-1 mlong, stout, obtusely 
4-angled, divaricately long-branched from middle upward, the entire plant, including ca- 
lyces, gray with dense, very fine and rather slender, slightly crisp hairs; leaves somewhat 
distant, 3 to 6 cm long, 2-8 cm wide, broadly ovate or (the upper) lanceolate from cordate 
base, 3-partite nearly to base, the lobes pinnatisect into 2-4 pairs of segments, these nar- 
rowly linear, entire, divergent, slightly revolute-margined, mucronate, with a prominent 
midrib beneath, leaves on side-shoots also 3-partite but the middle lobe 3 -sect, the lateral 
lobes entire; inflorescence broadly and shortly pyramidal-paniculate; bracts small, 3-part- 
ite into Hnear lobules; peduncles 1 -flowered, remote or spreading, filiform, as long as or 
longer than the bracts, nearly half as long as the calyx; flowers small; calyx 5-7 mm long, 
campanulate, ribbed, like pedicels short-tomentose, the narrowly lanceolate acute teeth 
nearly as long as tube; corolla 6-8 mm long, barely exceeding to at most half as long again 
as the calyx, blue, with obtuse lobes, tomentulose outside; filaments short, scarcely ex- 
serted; nutlets with short hairs. June. 

Dry mountain slopes, arable and weed-infested land. - Caucasus: S. Transc. (Mount 
Aragats, Kipchak). Gen. distr.: Bal.-As. Min., Iran. Described from Armenia. Type 
possibly in Leipzig. 

Note. As regards occurrence in the Russian flora, we are only aware of Radde's old 
specimens, from the localities indicated. Apart from T. parviflorum, the series Parvi- 
flora, which corresponds to Micrandrae Briq., also contains T. macrum Boiss. et Hausskn. 
from S. Iran. 



Section 2. Stachyobotrys Benth. Lab. gen. et sp. (1835) 662. — Perennials; flowers 
soUtary or in irregular verticillasters, forming a simple, more or less one-sided spike; calyx 
cam.panulate, 2-lipped, with 3 larger, broadly ovate upper teeth. 

31 




PLATE III. 1 - Teucrium orientale L., general aspect, flower, calyx; 2 -T. orientale var. villo- 
sum Benth., part of stem with leaves; 3 - T. hircanicum L., summit of stem with inflorescence, 
flower, calyx; 4 - T. scordium L., general aspect, flower, calyx; 5 - T. scordioides Schreb., part 
of stem with leaves. 



32 



4. T. hircanicum L. Syst. X (1759) 1096; Sp. pi. (1763) 789. - T. hyrcanicum 
Ldb.Fl. Ross. Ill, 1,442; N. Pop. in Mat. Fl. Kavk. IV, 3, 61. - Scorodonia spicata 
Moench, Meth. (1794) 385. - Ic: Bot. Mag. XLV, tab. 2013. - Exs.: Herb. Fl. Cauc. 
No. 345. 

Perennial, with short, ascending, woody rhizome; stems 20-70 cm long, herbaceous, 
few-branched or simple, densely covered with soft, short or rather long, spreading hairs, 
often reddish, especially below; leaves 2.5-8 cm long, 1-5 cm broad, ovate-oblong, gradu- 
ally tapering from deeply cordate or truncate base, obtuse, incised-crenate, with many 
(10-30 on each margin) large obtuse or subacute unequal teeth, rarely bicrenate, green 
above, with scattered or rather dense hairs above, grayish beneath with very long somewhat 
crisp hairs, densely hairy on very prominent veins; calyx 0.3-2 cm long, with vesture as on 
stems; lateral branches, if present, shortish, ascending; inflorescence showy, dense, spicate, 
6-25 cm long, cylindrical; lateral inflorescences, if present, much smaller; bracts marked- 
46 ly differentiated from upper cauline leaves, 5-8 mm long, linear-subulate, villous-hairy, 
equaling or slightly shorter than the calyx, concealed among flowers; pedicels straight, 
spreading, shorter than the calyx; calyx 3-5 mm long, sUghtly accrescent, campanulate, 
drooping, densely patent-viUous, 2-lipped, the upper teeth broader than the others, broad- 
ly ovate, mucronate, membranous, sparingly pubescent, netted-nerved, the lateral teeth 
short, nearly semiorbicular, obtuse, the lower teeth ascending, lanceolate, acute; corolla 
purple, short-villous-hairy outside, half as long again as calyx; filaments long-exserted; 
nutlets subglobose, ca. 1 mm long, granulate, glandular. June-August. (Plate 111, 
Figure 3.) 

Forests, wood margins, coppices, clearings, meadows. — Caucasus: Dag., W. and E. 
Transc, Tal. Gen. distr.: Iran. Described from Girkan province. Type in London. 

Economic importance. Sometimes grown as ornamental. 



Section 3. Scordium Rchb. Fl. germ. exc. (1831) 314; Benth. Lab. gen. et sp. 662.- 
Perennials, with remote, 2-6-flowered, axillary verticillasters; calyx with obUque, gibbous 
base, with subequal teeth. 



Series 1. Scordia Juz. — Eglandular perennials, with entire leaves. 

5. T. scordium L. Sp. pi. (1753) 565; Ldb. Fl. Ross. Ill, 1, 443; Shmal'g. Fl. II, 347; 
N. Pop. in Mat. Fl. Kavk. IV, 3, 59; Kryl. Fl. Zap. Sib. IX, 2293. - T. palustre Lam. Fl. 
Fr. 11(1795)411.-1. arenarium S. G. Gmel. Reise, I (1770) 149. - T. scordium ssp. 
palustre Gams in Hegi, III. Fl. V, 4 (1927) 2531. — T. scordium var. borysthenicum 
Schost. in Bot. mat. gerb. Bot. inst. AN SSSR, VIII, 9 (1940) 148. - Chamaedrys scor- 
dium Moench, Meth. (1794) 384. - Ic: Gmel. 1. c. tab. XXVI (T. arenarium Gmel); 
Rchb. Ic. Fl. Germ. XVIII (1856) tab. 38, fig. II, 2-4. - Exs.: Fl. pol. exs. No. 226; Eesti 
Taimed, No.83,83-a. 

Perennial, with creeping rhizome and freely rooting stoljons; stems ascending, then 
erect, 10-25 (40) cm long, simple or branching, subterete, softly villous-hairy all over, 
49 often — like leaves — violet-tinged; leaves closely approximate, 15-30 (50) mm long, 



33 



4-12 (20) mm wide, thin, oblong-elliptic, 2-3 times as long as broad, only the lower slight- 
ly cordate or rounded at base, these on branches tapering to sessile base, crenate with 4-6 
large acutish teeth on each margin, pinnately nerved with short appressed hairs above and 
spreading hairs beneath, especially on veins, otherwise glabrate or with short spreading 
hairs; middle leaves largest; bracts only slightly smaller than leaves, much longer than the 
flowers; inflorescence slender; verticillasters about half as long as the bracts, usually 2-5- 
flowered; flowers 6-10 mm long, short-pediceled; calyx tubular-campanulate, gibbous at 
base, glandular-villous, green or more or less violet-tinged, the teeth lanceolate, acute, half 
as long to nearly as long as the tube; corolla twice as long as the calyx, pale crimson-red, 
rarely white, the lower lip sparingly pubescent beneath, the rounded middle lobe and the 
lateral lobes recurved; stamens sliglitly exserted; nutlets ca. 1 mm long, reticulate-pitted. 
Plant smeUing of garlic. July-August. (Plate III, Figure 4.) 

Damp meadows, ditches, banks of rivers and lakes (especially sandy banks). — European 
part: Bait., V.-Kama, U. Dnp., M. Dnp., V.-Don, Transv., Bl., L. Don, L.V.; West Siberia: 
U. Tob., Irt.; Centr. Asia: Ar.-Casp. Gen. distr.: Scand., Centr. and Atl. Eur., Med. 
(Spain, Italy). Described from W. Europe. Type in London. 

Note. This species is not sharply differentiated from the next. Transitional forms occur 
where the distribution areas overlap, e.g. with leaf base and serration as in T. scordium 
but vesture excessive for this species, or, conversely, with the vesture of T. scordium but 
exceptionally obtuse-crenate leaves, etc. The variety estabhshed by Shostenko apparently 
represents one of these transitional forms. 

Close study of some of the southern Russian forms of T. scordium (in particular the 
Don and S. Kazakhstan specimens) may lead to their separation from the genuine T. scor- 
dium; they seem to represent a poorly differentiated race gravitating toward T. scordio- 
ides (stems more strongly branching, leaves with more obtuse teeth, vesture slightly more 
profuse, flowers smaller) that may qualify for the name T. arenarium S.G. Gmel. 

Economic importance. The plant contains essential oils, bitter principles and tannin, 
50 and has long been known for its medicinal properties and was highly valued in antiquity. 
It has been used as antiseptic, tonic and anthelmintic, for treatment of bubonic plague, 
lung diseases, etc. 

6. T. scordioidesSchreb. Fl. Vert. Unilab. (1774) 37; Ldb. Fl. Ross. Ill, I, 444; 
Shmal'g. Fl. II, 347; N. Pop. in Mat. Fl. Kavk. IV, 3, 60. - T. scordium var. scordio- 
ides Arcang. Fl. Ital. (1882) 559. - T. scordium subsp. scordioides Rouy, Fl. Fr. XI 
(1909) 237. - T.lanuginosum Hoffmgg. et Link, Fl. Portug. I (1809) 80. - Ic: Rchb. 
Ic. Fl. Germ. XVIII, tab. 38, fig. Ill; Hoffmgg. et Link, 1. c. tab. 1. - Exs.: Orph. Fl. 
Graec. No. 540. 

Perennial, with creeping rliizome and rooting stolons; stems usually erect, 12-60 cm 
long, mostly profusely branching, 4-angled, uninterruptedly, densely and softly lanate- 
tomentose; leaves closely approximate, 0.5-4.5 cm long, 0.25-2.5 cm wide, oblong-ovate 
or short-eUiptic, usually not more than twice as long as broad, rounded at base (especially 
on side-shoots) or mostly cordate, semiamplexicaul, sessile, obtusely crenate, with 3-12 
large crenations on each margin, thin, more or less densely tomentose-villous on both 
sides with rather long hairs; bracts usually not more than twice the length of flowers; 
main inflorescence long and slender; axillary inflorescences up to half the length of the 



34 



bracts, 2-5-flowered; flowers 5-8 mm long; pedicels rather long, especially in fruit; calyx 
campanulate, gibbous at base, densely tomentose-villous, usually green, the teeth lanceo- 
late, acute, half as long as the tube; corolla twice as long as the calyx, purple, sometimes 
white, with slightly exserted stamens. In all other respects similar to T. scordium L. 
July -September. (Plate III, Figure 5.) 

Wet, sometimes solonetzic meadows, marshes, sandy and stony banks of rivers, brooks 
and lakes, fluvial plains, seacoasts, rarely steppes or mountain slopes. - European part: 
Bes., Crim.; Caucasus: universal (becoming rare in S. Transc); Centr. Asia: Balkh., Mtn. 
Turkm., Syr D.,Pam.-Al., T. Sh. Gen. distr.: Atl. Eur., Med., Bal.-As. Min., Iran. De- 
scribed from Crete. 

Economic importance. Presumably as in preceding species; according to Grossgeim, 
used as a condiment in the Caucasus. 



Series 2. Bo try s Juz. — Annuals, with strongly developed glandular vesture and pin- 
natipartite leaves. 

7. T. botrysL.Sp.pl. (1753) 562; Ldb. Fl. Ross. Ill, I, 444; Shmal'g. Fl. II, 348; 
Klok. in Vizn. rosl. 408. — Chamaedrys botrys Moench, Meth. (1794) 383. — Scoro- 
donia botrys Ser. Bull. Bot. (1830) 317. - Botrys chamaedryoides Fourr. in Ann. 
51 Soc. Linn. Lyon. 17 (1869) 138. - Ic: Rchb. Ic. Fl. Germ. XVIII, tab. 38, fig. II, 1; 
Hegi,III. Fl. V,4, fig. 3368. 

Annual, rarely perennial, with oblique, frequently curved root; stems mostly much 
branched from low down, rarely simple; branches (5) 10-39 (40) cm long, arcuately 
ascending or straight at base, then dechned or nearly upright, obtusely 4-angled, green or 
reddish, entire length covered with horizontally spreading short and long glandular hairs 
interspersed with fewer long simple ones; leaves 1-2.5 cm long and as wide, ovate to 
broadly ovate, usually deeply 1-2-3-pinnatifid, the lobes short, linear or spatulate, obtuse, 
1 -nerved, dark green with short scattered hairs above, grayish with sessile and stipitate 
glands beneath, the veins and margins with recurved hairs; petioles 0.5-1.5 cm, covered 
with hairs of the same kind as on stems; verticillasters 2-4-flowered, secund, borne in 
axils of undifferentiated upper leaves, slightly shorter than the latter; flowers 10-15 mm 
long; pedicels 3-7 mm long, erect, spreading or declined with hairs of the same kind as 
those on stem and calyx; calyx campanulate, reticulate-nerved, curved, strongly gibbous, 
densely covered with stipitate glandular hairs, the teeth triangular, erect or sHghtly con- 
verging, the two lower smaller than the rest; corolla purple, the short tube included in 
calyx, the lower lobe of lip pale, large, concave, sparingly hairy outside, the lateral lobes 
acute, 3-fid, the upper lobes small, acute; stamens and style long-exserted, curved; nutlets 
subglobose, 1 .5-2 mm long, scabrous-pitted, with deep furrow at areola. The whole plant 
has a disagreeable odor. July — August. 

Dry hills, calcareous rocks, shrubs. — European part: U. Dns. Gen. distr.: Atl. and 
Centr. Eur., Med. Described from W. Europe. Type in London. 



Section 4. Chamaedrys Schreb. Verticill. gen. (1774) 19; Benth. Lab. gen. et sp. 
(1835) 662. — Subshrubs, with 2-6-flowered verticillasters forming a loose, rarely compact. 



35 



racemose inflorescence; calyx tubular, attenuate at base and slightly gibbous, with 5 lan- 
ceolate or triangular subequal teeth. 

Note. Until recently, T. canum Fisch. et Mey. was the only species in this group 
segregated by Russian botanists from T. chamaedrys L., while all other forms were 
combined under the latter name. They constitute, however, a very complex cycle with 
52 many variable forms. K. H. Rechinger (Fil. Bot. Arch. 42, 1941, 335-420) quite aptly 
refers to this cycle as "confused but very attractive." The present treatment of the section 
is far from definitive. We have endeavored to arrange the known forms of the cycle of 
T. chamaedrys s.l. in three series though some of these may crossbreed and produce 
numerous "transitional" forms, which may obliterate the boundaries between series. This, 
of course, seriously complicates the observed relationships within the group and the classi- 
fication of the forms included. 



Series 1. Nuchensia Juz. - Stems and lower side of leaves covered with simple spread- 
ing or recurved hairs (not tomentose). Margins of calyx-teeth with scattered or rather nu- 
merous long hairs. Inflorescence often abbreviated; bracts gradually diminishing in size, 
all more or less dentate, the upper more or less concave, scaphoid. A predominantly Cau- 
casian group. 

8. T. nuchense C. Koch in Linnaea, XXI (1848) 703; Bornm. in Mitt. Thiir. Bot. Ver. 
N. F. 37, 6. - T. chamaedrys Ldb. Fl. Ross. Ill, I (1849) 444, p. p.; Shmal'g. Fl. II, 
348, p.p.; N. Pop. in Mat. Fl. Kavk. IV, 3, 49, p. max. p.; Grossg. Fl. Kavk. Ill, 284, p. 
max. p. et auct. plur. Fl. Cauc, non L. — T. chamaedrys ssp. nuchense Reching. fil. 
in Bot. Archiv, 42 (1941) 370. - Ic: Reching. fil. I.e. fig. 6 (p. 371) et tab. I, fig. 10, 
tab. Ill, fig. 47. 

Perennial; plant woody at base only; flowering stems 10-30 (40) cm, with geniculately 
curved base, mostly stoutish or stout, upright, simple or short-branching above, on 2 or 4 
faces diffusely hairy, the hairs somewhat crisp, reclinate, shorter than diameter of stem; 
leaves 1-2.5 cm long, 0.5-1 .4 cm wide, short-elliptic from cuneate base, rounded at apex, 
VA-2 times as long as wide, rather evenly crenate or incised-crenate with 5-12 short obtuse 
teeth, furcately 3-6 (mostly 5)-veined on each side, thickish, green on both sides, covered 
very sparsely above, more densely beneath, with short patent hairs; petioles to 3 mm long, 
often scarcely deHmited from blade, well developed only in lower leaves, the petiole and 
base of blade rather sparsely and minutely ciliolate; inflorescence mostly very short, con- 
53 tracted, rarely lax, at ends of stems and branches; bracts cuneate or somewhat rounded at 
base, entire from base to middle, with ciHate margin, the lowermost longer than flowers, 
the upper navicular, shortly toothed at apex, the uppermost entire, often distally 2-fid; 
verticillasters many-flowered, more or less crowded; pedicels often much shorter than 
calyx; calyx campanulate, sparsely covered on the nerves, rarely all over, with short re- 
clinate hairs, the teeth triangular, scarcely longer than wide, approximately half the length 
of the tube, short-acuminate, with scattered long stiff cilia along margin, short cilia absent 
or very sparse; corolla (including lower lobe of Up) to 2 cm long, purple, copiously villous 
outside. June— August. (Plate IV, Figure 1.) 



36 



Bare or grassy dry mountain slopes, rocks, meadows, wood margins, pine forests. — 
Caucasus: all regions. Gen. distr.: Bal.-As.Min. Described from "Nuchi." Type in Berlin. 

Note. As observed by Rechinger (1. c. 1941, 372-373), the more or less typical forms 
of this rather polymorphous member of section Chamaedrys, which is very widespread in 
the Caucasus, grow together with forms gravitating toward T. chamaedrys L. It is very 
likely that the latter represent hybrids of these species. Today, the typical T. chamae- 
drys L. appears to be rare in the Caucasus (see Note to T. chamaedrys L. below). 

9. T. multinodum (Bordz.) Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 
21. — T. chamaedrys var. multinodum Bordz. apud N. Pop. in Mat. Fl. Kavk. IV, 3 
(1916)57. 

Perennial; stems rather few, woody at base, mostly upright or scarcely ascending at 
base, 1 5-30 cm long, very stout, robust, straight or slightly flexuous, densely covered with 
short recurved hairs, frequently dark purple; leaves many, commonly 10-15 pairs, 2.5 cm 
apart, medium size, 1-2.2 cm long, 0.5-1.7 cm wide, equaling or shorter than the inter- 
nodes, broadly ovate or the upper ovate, approximately IVi times as long as broad, the 
lower subcordate, the middle and upper truncate or broadly cuneate at base, all leaves 
abruptly cuneate at base, rounded or obtuse at apex, rather evenly and shallowly incised- 
54 crenate, very often with double or even treble crenations on each margin reaching 1/5-1/4 
of half-blade width; blade rather firm, flat, with scattered or rather dense, long, nearly 
straight hairs above, denser and shorter recurved hairs beneath, the basal ciUa very distinct; 
lateral veins 4-6 on each side of midrib, usually branching, impressed above, very promi- 
nent beneath; petioles very short, to 3 mm long (sometimes leaves subsessile), with vesture 
Uke that of stems; inflorescence 2-9 cm long, dense, sometimes subcapitate, with only low- 
er nodes frequently remote, the internodes to 1 cm long; axis of inflorescence with hairs 
like those on stem but denser; lowermost bracts undifferentiated, the others more or less 
concave, eUiptic, acute, with entire convex margins in lower half, finely and obtusely 
toothed in upper half, very short-petioled or sessile; verticillasters 2-6-flowered; flowers 
1-1 .5 cm long; pedicels to half the length of the calyx; calyx campanulate, rounded at 
base, more or less purple-violet, covered with longish, stiff, patent or reclinate hairs, the 
teeth slightly spreading, broadly short-triangular, mucronate, very sparingly hairy, with a few 
longish cilia on the margin, corolla probably pale, with dark purple lower lip. June— July. 

Grassy habitats, coppices and forest margins in lower mountain zone. — Caucasus: 
W. Transc. Endemic. Described from Akhali-Afoni. Type in Leningrad. 

Note. The typical form is known only from Abkhazia; specimens from the more 
northern locations are markedly anomalous, with less hairy leaves, strongly branching 
stems and very loose inflorescences (T. circassicum Juz. ined.). Toward the south 
T. multinodum seems to vicariate with the next species. 

10. T. trapezunticum (Reching. fil.) Juz. comb. nova. — T. chamaedrys ssp. trapez- 
unticum Reching. fil. in Bot. Arch. 42 (1941) 369. - Ic: Reching. fil. 1. c. tab. I, fig. 9 
et tab. 3, fig. 46. 

Perennial, similar to T. multinodum (Bordz.) Juz. but stems with denser, softer and 
longer hairs; leaves broadly notched-cuneate at base, ovate-spatulate, ca. 2 cm long,. 1.2- 
1.5 cm wide, incised-crenate or nearly lobate with incisions 1/3 of half-blade width, 



37 



apically more or less short-crenate or incised, diffusely covered with long semi-appressed 
55 hairs above, almost tomentose beneath; petioles short, frequently obsolescent; calyx with 
dense, long, spreading hairs, the teeth short and broad, approximately Wi times as long as 
wide, long-acuminate, with longscattered marginal ciUa; otherwise like T. multinodum. 
June— July. 

Exposed stony and rocky sites. -Caucasus: W.Transc. (Adzhar). Gen.distr.: Bal.-As. 
Min. (NE Anatolia, Trebizond). Described from vicinity of Trebizond. Type in Vienna. 

Note. We did not see the type of this inadequately investigated form but, judging by 
the author's description, we have tentatively included in it a specimen collected by Litvi- 
nov somewhere between Adzharis-Tskali and Keda in Adzhar. This specimen is easily 
distinguishable from all T. multinodum (Bordz.) Juz. plants we have seen by the 
somewhat different vesture of stems and leaves and the characteristic leaf base. However, 
both species are closely related to T. nuchense, and may merely represent local races. 



Series 2. Eu-chamaedrys Juz. Like the preceding series but the teeth of the leaves 
with both long and short dense marginal hairs. Inflorescence longer and looser; bracts 
narrower, more or less concave, only the uppermost sometimes entire. A predominantly 
European group. 

11. T. chamaedrys L. Sp. pi. (1753) 565,s. str.; Ldb. Fl. Ross. 1, 1,444, p. p.; Shmal'g. 
Fl. II, 348, p.p. - T. chamaedrys ssp. eu-chamaedrys Reching. fil. in Bot. Arch. 42 
(1941)344. -T. chamaedrys |3. lucidum Celak. Bot. Centralbl. 14(1883) 220. - T. 
chamaedrys var. glanduliferum Hausskn. in Mitt. Thiir. Bot. Ver. N. F. 11 (1897) 181. 

- ? T. pseudochamaedrys Wender in Flora, IX, 1 (1826) 358. - T. officinale Lam. 
Fl. Fr. II (1779) 414, p. p. - Chamaedrys officinalis Moench, Meth. (1794) 383, p. p. 

- Ic: Reching. fil. 1. c. tab. 1, fig. 1-4. - Exs.: GRF, No. 1493. 

Perennial, with woody base; flowering stems 10-45 cm long, slender to fairly stout, 
ascending at base or upright, usually curved or tlexuous, rarely almost straight, simple or 
commonly slightly branching, covered on two or all four faces with short, sparse, soft, 
recurved hairs, pale green or reddish; leaves rather many, more or less remote, 0.7-4 cm 
long, 0.5-2.5 cm wide, usually ovate, rarely elliptic or oblong, \V2-2]6. times as long as 
broad, cuneately tapering from broadened base, gradually passing into petiole from which 
56 they are hardly differentiated (only the lowermost well delimited from petiole), rounded 
or acutish at apex, largely crenate-dentate or with broad rounded or pointed lobes, entire 
or more or less crenatc or incised; teeth or lobes 4-9 on each margin; lateral veins 4-5 on 
each side of midrib, scarcely impressed above, scarcely protruding beneath, mostly rather 
tliin. softly herbaceous in live state, mostly chartaceous when dry, bright green above, just 
a shade paler (but not glaucous) beneath; hairs on upper surface scattered, appressed, on 
lower surface more profuse or else confined to veins, curved and mostly pointing down- 
ward or sideways, the base of blade and petiole with mostly shortish cilia; petioles l-7mm 
long; inflorescence short or elongate but usually much less than half the length of stem; 
bracts fairly large, nearly always exceeding the calyx, more or less concave (especially the 
upper), entire at cuneate base (lower third and more), otherwise crenate-dentate, generally 
green, rarely the upper lilac-tinged; verticillasters 2-5-fiowered, usually somewhat distant; 



38 



pedicels shorter than or sometimes equaling the calyx; flowers large, purple; calyx 5-6 mm 
long, rather broadly campanulate, the short, usually curved hairs scattered or confined to 
nerves, the triangular short-acuminate teeth to P/4 times as long as broad, very densely be- 
set along the margin with short cilia and short-stipitate glands forming a kind of fringe in 
addition to scattered or fairly dense long cilia; corolla (including lower hp) 1-1.3 cm long; 
nutlets ca. 1 .5 mm long, broadly ellipsoid, brown, smooth or scarcely rugose. June— August. 
(Plate IV, Figure 2.) 

Steppes and steppe slopes, deciduous and pine forests, forest margins, coppices, rocks 
and outcrops, stony mountain slopes and taluses, sands. — European part: M. Dnp., BL, 
L. Don, Bes., U. Dns., Crim.; Caucasus: Cisc, Dag., S. Transc. (also reported for many 
parts of the Caucasus, but most of these reports clearly refer to T. nuchense). Gen.dist.: 
Centr. Eur. (southern part), Med. (Apennines), Bal.-As. Min. (Balkans, AnatoUa).. Describ- 
ed from Germany, Switzerland, France. Type in London. 

Note. A typical feature in the distribution within the USSR of this most eury topic 
member of the section is its penetration from the north and south into the Caucasus 
where it encounters and partly coexists with forms of T. nuchense C. Koch, that are very 
common and unquestionably autochthonous in that region. 
57 Economic importance. The plant contains essential oil, tannin and other substances. It 
has long been known as a medicinal plant and is still valued in popular practice. The aerial 
part of the flowering plant is dried and boiled as an infusion. It has been used as excitant, 
stomachic, tonic, diuretic, antifebrile, antidote, etc. Known in horticulture as a border 
plant. Tanning agent. 

12. T. pulchrius Juz. in Bot. mat. gerb. Bot. Inst. AN SSSR, XIV (1951) 22. - T. cha- 
maedrys auct. Fl. taur. p. p. non L. s. str. 

Perennial, 8-20 cm high; stems short, hairy, ascending at base; leaves in relatively few 
approximate pairs, rather heavily pubescent but bright green, the teeth mostly shallow and 
broad, simple or double; flowers appearing early so that inflorescence usually accounts for 
nearly half the overall length of stem; inflorescence frequently branching in lower part, 
contracted, with very crowded verticillasters, the lower bracts foliaceous, the upper dark 
lilac, mostly subscaphoid, even the uppermost dentate at apex; flowers very large; calyx 
ca. 7 mm long; corolla (including lower lip) ca. 16 mm long, dark purple. Otherwise Hke 
T. chamaedrys L. June — August. 

Grassy mountain slopes, meadows. — European part: Crim. Endemic. Described 
from Avind (Nikitskaya Yaila). Type in Leningrad. 

Note. A rather weakly expressed Crimean high-mountain race of the preceding species 
that appears to be a Crimean analogue of the Balkan T. olympicum Reching. fil. (pro 
subsp. T. chamaedrys). 

13. T. fischeri Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 23. - T. tauri- 
cum Fisch. in herb, ex Juz. 1. c. - T. cham aedry s auct. fl. taur. p. p. non L. s. str. - T. 
chamaedrys var. hirsutum Celak. in Bot.Centralbl. 14(1883) 220. -T. chamaedrys 
var. pubescens N. Pop. in Mat. Fl. Kavk. IV, 3 (1916) 57, p. p. - T. syspirense var. 
hirsutum Reching. fil. in Bot. Arch. 42 (1941) 375, p.p. 

Perennial, to 30 cm high; stems slightly ascending at base or nearly upright, covered 
with thick, horizontally spreading or often distinctly reclinate, rather long white hairs; 

39 



leaves in many subremote pairs, ovate-oblong, cuneately tapering to petiole, with many 
fine subequal simple or double obtuse teeth, gray -green with profuse patent hairs on both 
sides, subvelutinous beneath; inflorescence at most 1/4-1/3 the overall length of the stem, 
58 simple, rarely long-branched in lower part, contracted, mostly with rather crowded verti- 
cillasters; bracts gradually smaller toward summit, the lower resembling the stem-leaves, 
the middle entire laterally, the uppermost often entire throughout, lanceolate, nearly flat 
or mostly navicular, green, or the upper sometimes dull violet; flower medium size; calyx 
4-6 mm long; corolla (including lower lip) 9-12 mm long, pale pink, the lower Up purple. 
Otherwise like T. chamaedrys L. June— August. 

Forests (mainly coniferous) and stony mountain slopes. — European part: Crim. 
(southern shore); Caucasus: W. Transc. (Novorossiisk area). Endemic. Described from 
Cape Mart'yan. Type in Leningrad. 

Note. We have included this plant in the series Euchamaedrys principally on account 
of the navicular upper bracts. In such characters as leaf dentation and vesture, and the 
often completely entire bracts, it closely resembles some forms of T. syspirense affiUa- 
tion, with which it was apparently confused by Rechinger. It might well be suspected of 
being a hybrid, possibly a cross between T. chamaedrys L. and T. krymense Juz., were 
it not for some of its particular features, notably the paleness of the corolla, not associated 
with either of these species. 



Series 3. Syspirensia Juz. - Like the preceding series, but inflorescence markedly 
elongated; verticillasters very distant; bracts abruptly diminishing in size, all flat, the 
upper entire. Essentially Anatohan and Iranian groups. 



14. T. syspirense C. Koch in Linnaea, XXI (1 848) 704; Bornm. in Mitt. Thiir. Bot. 
Ver. N. F. 37, 59-60, p. p. - T. chamaedrys var. canum Boiss. Fl. or. IV (1879) 816, 
p.p. — T. chamaedrys var. pubescens N. Pop. in Mat. Fl. Kavk. IV, 3 (1916) 57, p. p. 
— T. chamaedrys ssp. syspirense Reching. fil. in Ann. Nat. Mus. Wien. 51 (1941)427; 
id. in Bot. Arch. 42 (1941) 373. - Ic: Reching. fil. in Bot. Arch. 42, 372, fig. 7; tab. I, 
fig. 11; tab. 3, fig. 48. 

Perennial, with woody taproot and woody stem base; stems ascending or upright, 15- 
25 cm long, slender, flexuous, covered with patent and reclinate hairs, these shorter than 
61 diameter of stem; leaves 0.7-1 .5 cm long, 0.4-0.8 cm wide, ovate or oblong-ovate, shortly 
and broadly incised-cuneate at base, rounded or obtuse at apex, Wi times as long as wide, 
rather regularly and finely crenate-dentate, with 4-8 entire, rarely cleft teeth on each mar- 
gin, dark green with sparse appressed hairs above, canescent with hairs beneath; petioles 
clearly delimited, about 1/3 as long as the blade, obscurely ciliate; inflorescence often 
markedly elongated; bracts abruptly diminishing in size, shorter than calyx, cuneate- 
obovate to lanceolate, acute, dentate only at apex, the upper lanceolate, entire, all flat; 
verticillasters 4-6-flowered, distant; lower pedicels nearly as long as the calyx, the upper 
shorter; calyx 5-6 mm long, tubular-campanula te, with short spreading hairs, very often 
purple-violet; teeth triangular, rather long-acuminate, the margins covered with very pro- 
fuse short hairs and scattered long bristles; corolla half as long again as the calyx, purple, 
the middle lobe dark purple. June — July. 

40 




PLATE IV. 1 - Teucrium nuchense C. Koch, summit of stem with inflorescence, cauhne leaf, bract, 
flower, calyx, calyx-tooth; 2-T. chamaedrys L., leaf, bract, flower, calyx, calyx-tooth; 3 - T. kry- 
mense Juz., summit of stem and inflorescence, leaf, bract, flower, calyx; 4 - T. jailae Juz., general 
aspect, leaf, bract, flower, calyx; 5 - T. polium L., general aspect, leaf, bract, flower. 



41 



Dry stony mountain slopes. -Caucasus: W. and S. Transc? Gen. distr.: Bal.-As.Min., 
Iran. Described from Lesser Armenia (Sber and Ispir). Type in Berlin. 

Note. We have interpreted this species in its original and more restricted sense. The 
very wide scope assigned to it by Bornmiiller and Rechinger hardly provides a practical 
approach. The typical T. syspirense C. Koch is unknown in the USSR, but it was col- 
lected by Nesterov in Oltyn province and by Turkevich in Artvin province, and may occur 
in the USSR. For this reason we have included it in "Flora." 



15. T. krymense Juz. in Bot. mat. gerb. Bot. inst. ANSSSR,XIV(1951) 19. - T. cha- 
maedrys var. pubescens N. Pop. in Mat. Fl. Kav. IV, 3 (1916) 57, p. p. — T. chamae- 
drys ssp. syspirense Reching. fil. in Bot. Arch. 42 (1941) 373, p. p. non T. syspirense 
C. Koch. -T. syspirense Zef. Gubotsv. Kryma (1951) 5, non C. Koch. 

Perennial, woody at base, branching; flowering stems many, ascending or nearly up- 
right, 20-40 cm long, slender or fairly robust, somewhat flexuous, densely covered with 
soft grayish, slightly declined hairs and nearly vellutinous like the leaves and their petioles; 
leaves small or medium size, 10-20 mm long, 6-12 mm wide, ovate, approximately twice 
as long as wide, the lower broadly cuneate at base, the upper more narrowly cuneate, 
62 rounded at apex, rather evenly and finely crenate-dentate, with about 7 often double teeth 
at each side more than 1/3 the width of half the blade, the blade rather firm, flat; lateral 
veins about 5, often branching, impressed above, slightly protruding beneath but very dis- 
tinct against the lighter background of hairs; upper surface loosely subappressed-hairy, the 
lower less velutinous-tomentose with white hairs, basal cilia indistinct; petioles approxi- 
mately 1/3 as long as blade, clearly dehmited from blade in lower leaves; bracts flat, short- 
petioled; the lowermost often resembling stem-leaves, others smaller, oblong or oblong- 
ovate, cuneate at base, acute, entire laterally, with few small acute teeth only at apex, the 
uppermost broadly lanceolate, entire, vesture of inflorescence axis like that of the stem; 
verticillasters 3-6-flowered, all or only the lower and the middle ones remote, forming a 
loose, often very long simple raceme; flowers 10-12 mm long, on pedicels 2/3 as long as 
calyx; calyx tubular-campanulate, very dark purple -violet (almost black, Uke stems in up- 
per part), rather densely covered with patent or reclinate hairs of medium length, the 
teeth triangular, long-acuminate, IM times as long as wide, with very profuse short and 
scattered long cilia; corolla very dark purple (nearly black). June— August. (Plate IV, 
Figure 3 .) 

Grassy mountain slopes. —European part: Crim. Endemic. Described from the vicinity 
of Old Crimea, Golyi Agarmysh Mountain. Type in Leningrad. 

Note. The distribution area of this species is completely separated from that of the true 
T. syspirense C. Koch which has not so far been found anywhere in the Caucasus and for 
this reason we feel justified in treating the Crimean form as a separate taxonomic unit, not- 
withstanding its relatively limited distinctiveness. 



16. T. excelsum Juz. sp. nov. in Addenda, XIX, 505. — T. chamaedrys ssp. syspi- 
rense Reching. fil. in Bot. Arch. 42 (1941) 377, p. p. non T. syspirense C. Koch. — Exs.: 
Sintenis, It. transcasp.-pers. 1900-1901, No. 1054. 



42 



Perennial, woody and branching at base; flowering stems usually many, upright or near- 
ly so, 12-45 cm long, moderately sturdy, straight or slightly flexuous, rather densely cover- 
ed with white, spreading and slightly recUnate hairs; leaves 1 .2-3 cm long, 0.8-2 cm wide, 
elhptic or broadly ovate, approximately IVi times as long as wide, broadly cuneate at base 
or (the upper) narrowly cuneate, obtuse or acute at apex, irregularly and largely incised- 
crenate-dentate, sparsely covered above with whitish curved hairs, more densely so beneath, 
but not velutinous; teeth 4-7 on each margin elongate, obliquely antrorse, often double, 
nearly half of half-blade width, blade flat; lateral veins 4-6, usually branching, scarcely 
impressed above, slightly protruding beneath; petioles very short, not more than 1/4 the 
length of blade, sometimes in lowermost leaves clearly delimited from blade; lower bracts 
resembling upper cauhne leaves, gradually smaller toward summit, oblong-rhombic with 
narrowly cuneate base, acute, sharp-toothed in upper half; upper bracts oblong-eUiptic, 
with few acute denticles only at apex, subsessile, flattish, the uppermost frequently entire; 
vesture of inflorescence axis like that of the stem; verticillasters 2-5-flowered, the lower 
distant, the others closely approximate; inflorescence racemose, or distally rather dense, 
sometimes slightly branching at base; flowers 1.3 cm long, on pedicels exceeding half the 
length of calyx; calyx obconical or subcampanulate, green or mostly purple-violet, densely 
covered with moderately long patent recurved hairs; teeth triangular, long-acuminate, IV^ 
times as long as wide, hairy on the margins and sometimes on the midnerve; corolla purple. 
End of May — July. 

Gravelly mountain slopes with steppe vegetation, pebble-beds along banks of mountain 
streams. — Centr. Asia: Mtn. Turkm. Gen. distr.: Iran. Described from Ai-Dere gorge. 
Type in Leningrad. 

Note. In referring T. excelsum to the series Syspirensia, we have adopted the approach 
of Rechinger who does not distinguish this species from T. syspirense C. Koch. It should 
be noted, however, that in some of its features, notably the size and shape of the leaf teeth, 
and the bracts that are not quite typical for this group, T. excelsum comes very close to 
the species of the preceding series (Eu-chamaedrys Juz.). In such "intermediate" nature 
of its characters it is comparable to T. fischeri Juz. In general, its taxonomic position 
needs further clarification. 



Series 4. Cana Juz. — Stems and lower side of leaves appressed-white-tomentose; 
stems and leaves nearly devoid of simple spreading hairs. 



17. T. canum Fisch. et Mey. Ind. sem. hort. Petrop. 1 (1835) 40; Ldb. Fl. Ross. Ill, 
445; Reching. fil. in Bot. Arch. 42, 405. - T. chamaedrys var. canum Boiss. Fl. or. 
IV (1879) 813, p. p. - Ic: Reching. fil. 1. c. f. 32 and 69. 

Perennial, woody at base, non-stoloniferous; flowering stems elongate, virgate, 15- 
45 cm long, arched-ascending or upright, canescent all over with very short appressed to- 
mentum; leaves distant, 0.8-2.5 cm long, 2-3 mm wide, generally shorter than internodes, 
erect, subcoriaceous, the lower narrowly ovate or obovate, the upper ovate-lanceolate or 
oblanceolate, convex, strongly revolute-margined, long-cuneate at base, gradually tapering 
to petiole, obtuse or acute at apex; teeth 5-8 on each margin, fine, short, rounded or acute; 



43 



lateral veins 4-5, irregularly branching; upper surface sparsely covered with fine short 
curved hairs; lower surface with vesture like that on the stem; petioles lacking cilia, in 
lower and middle leaves to 1/3 the length of blade, in uppermost leaves very short; in- 
florescence axis with hairs like those on the stem, interspersed above with longer patent 
hairs; lowermost bracts resembling stem-leaves, others entire, sessile, strongly revolute- 
margined, ovate-lanceolate, acuminate, devoid of cilia, much shorter than calyx; verticil- 
lasters many, very distant, the lower many-flowered (6-flowered on the average); flowers 
1 .2-1 .5 cm long, purple; pedicels about the length of calyx or shorter; calyx 5-7 mm long, 
campanulate, erect or sHghtly declined, covered with profuse short and fewer long patent 
hairs sometimes interspersed with very few glandular hairs, often slightly suffused with 
purple; teeth subequal, triangular, abruptly short-acuminate, scarcely longer than wide, 
half the length of the tube or slightly longer, the margin with profuse short and scattered 
long cilia; corolla nearly twice as long as the calyx, densely villous-hairy outside. June — 
August. 

Dry hills, clayey slopes, rocks, stony places, mountain steppes, dry scrub. —Caucasus: 
Dag., E. Transc. Endemic. Described from Somkhetia. Type in Leningrad. 

Note. The characters observed in Syspyrensia appear to be much more pronounced in 
the series Cana. Other species of this series grow in Lesser Armenia (T. Ixodes Bornm.) 
and Cappadocia (T. leucophyllum Montbr. et Aucher). 

A plant which is undoubtedly a hybrid T. canum X T. chamaedrys L. or more likely 
T. canum X T. nuchense C. Koch, which we have named T. X alexeenkoanum Juz. 
(in Addenda XIX, 342), was collected by Alekseenko near "Khaltanskaya Kazma" along 
the Ata-Chai River in the district of Kuba as No. 1 1037. The stems of this hybrid are 
moderately covered with short, crisp hairs; as compared with T. canum, the leaves, green- 
ish beneath, have a broader base, they are more sparingly hairy, their teeth are cut more 
65 deeply, and the bracts are shorter and broader; verticillaster containing up to 7 flowers; 
calyx deeply purple -violet. Popov included this plant in his T. chamaedrys var. pubes- 
cens N. Pop. This "variety" is an exceedingly composite concept which, among other 
forms, also includes T. syspirense C. Koch. Alekseenko's plant undoubtedly displays 
superficial resemblance to the latter but differs markedly in the character of its vesture. 



Section 5. Polium Schreb. Verticill. gen. (1774) 20; Senth. Lab. gen. et sp. (1835) 
663. - Perennials; verticillasters crowded in terminal ovoid or globular heads; calyx tubu- 
lar-campanulate, the teeth subequal or the lower teeth slightly narrower and more acute 
than the upper. 



Series 1. Montana Juz. — Flowers 1-1.5 cm long. Leaves entire. 



18. T.jailae Juz. inSpisokrast.gerb. fl. SSSR, XI (1949) 144. -T.montanum auct. 
fl. taur. non L.; Ldb. Fl. Ross. Ill, 445, p. p.; non L.; N. Pop. in Mat. Fl. Kav. IV, 3, 40. 
-Exs.: Gerb.fl. SSSR, No. 3467. 

Subshrub, with robust woody root to 1 cm in diameter, producing procumbent tufts of 



44 



varying size; stems low, much-branched, prostrate or ascending at the ends, densely soft- 
tomentose, rather slender, terete, whitish or with some reddish color showing from under 
the tomentum, the reduced internodes many times shorter than the leaves; leaves 4-18 mm 
long, 0.3-4 mm wide, crowded, lanceolate or sublinear, gradually tapering at base to very 
short petiole, obtuse at apex, usually markedly revo lute-margined, densely tomentose and 
gray-green above, very densely white-tomentose beneath, with loose tomentum consisting, 
as on stems, of rather long but slightly crisp hairs; midrib strongly impressed above, pro- 
minent beneath; lateral veins indistinct,completely concealed by tomentum; flower-heads 
mostly solitary, usually 3-7 -flowered, occasionally containing up to 20 and more flowers, 
dense, obconical or subglobular; bracts resembUng stem-leaves but shoiter; flowers 1.5 cm 
long, on short or very short pedicels; calyx tubular, gradually tapering at base, more or 
less densely tomentose, faintly anastomosing, pale green or faintly purple; teeth 2/5-1/3 
as long as the tube, somewhat unequal, narrowly triangular, gradually passing into short 
awn, erect or the upper at length moderately recurved; corolla sparingly pubescent out- 
side, the tube not exceeding the calyx; lower lip much enlarged, the middle lobe nearly 
square, with a few lobules, finely denticulate, the lateral lobes much smaller, oblong-elHptic, 
nearly flat, rounded at apex, recurved Hke the middle lobe, without marginal ciUa, the up- 
per lobes similar to the lower but slightly larger, ciliate at margin, with greenish stripes on 
back; anthers greenish; nutlets ca. 2 mm long, ellipsoid, black, slightly rugose, with large 
pale areola. June -August. (Plate IV, Figure 4.) 

Stony places (meidows and pastures), rocks, taluses. — European part: Crim. Endemic. 
Described from Ai Petrinskaya Yaila. Type in Leningrad. 



19. T. praemontanum Klok. sp. nov. in Addenda XIX, 342. 

Subshrub producing rather dense tufts 10-40cm long; current year's flowering branches 
2-10 cm long, densely canescent with short (to 0.25 mm) appressed or subappressed hairs; 
leaves oblong-obovate to sublinear, 5-20 mm long, 1 .25-4 mm wide, entire, cuneately ta- 
pering at base to short or indistinct petiole, more or less revolute-margined, obtuse or 
rounded at apex, whitish beneath with fine short tomentum, dark green above with scat- 
tered hairs, these more distinct (to 0.5 mm long) in lower part; bracts linear-lanceolate, 
with petiolar attenuation at base, 5-12 mm long, 0.75-2 mm wide, the lower longer than 
calyx (but shorter than whole flower), the upper shorter; inflorescence terminal, capitate, 
rather dense, 5-15-flowered; pedicels 1-2 mm long (the lower sometimes to 3.5mmlong), 
covered with semi-appressed short hairs; calyx subtubular, 8-9 mm long, the tube covered 
outside with short appressed hairs; teeth sparsely long-ciliate at margin, the lower longer 
and narrower, lanceolate, 2.25-3 mm long, to 1 mm wide, terminating in short point, the 
upper shorter and wider, acutely triangular, unequal, the middle 1 .75-2 mm long and as 
wide, the lateral 1.25-2rrmi long, 1.25-1.5 mm wide, acuminate or tapering to a point 
0.25-0.5 mm long, straight or often markedly recurved; corolla nearly twice as long as 
the calyx, whitish-yellowish, the tube broadening above, nearly erect, 5-6 mm long, the 
Up 7-10 mm long, the terminal lobe rounded, entire, concave, the lateral lobes oblong- 
elliptic, tapering to base, much smaller, the upper lobes oblong-ligulate, longer than the 
lateral; nutlets ovoid, ca. 1.75 mm long, 1 mm wide, rugose-pitted, blackish. June-July. 

Limestone and chalk outcrops. — European part: U. Dns. (Ternopol' Region, Ukrainian 



45 



SSR, Berezhan district, Trostyanets and vicinity of Kremenets). Endemic. Described from 
Trostyanets. Type in Kiev. 

Note. According to M.V. Klokov, who described this species, many specimens had 
been collected on a number of occasions by different people. Judging by these collections, 
the species occurs in the absence of T. pannonicum Kern, to which it does not appear to 
be closely related. 



20. T. pannonicum Kern, in Oesterr. Bot. Zeitschr. XIII (1863) 384. - T. montanum 
Ldb. Fl. Ross. 111,445, p.p.; Shmal'g. Fl. II, 348, p.p. non L. - T. montanum var. pan- 
nonicum Briq. Pflanzenfam. IV, 3a(1897) 821. -T. montanum (i. hirsutum Boiss. 
Fl. or. IV(1879)819.-Exs.: Fl. exs. Austro-Hung. No. 2643. 

Subshrub or small shrub, branching from woody base, with woody root; stems several 
to many, 8-25 cm long, prostrate or ascending, radially spreading, rarely suberect, loosely 
cespitose, stoutish, terete, villous with profuse white, slightly crisp hairs; internodes rather 
well developed, half as long as to equaling the leaves; leaves 0.8-2 cm long, 1-5 mm wide, 
sessile or very short-petioled, oblong, lanceolate or linear, tapering at base, subacute or 
obtuse at apex, entire, revolute-margined, villous-tomentose on both sides, sparsely so and 
hence dark green above, almost snow-white beneath; midrib moderately impressed above, 
somewhat prominent beneath, the lateral veins indistinct; verticillasters crowded in a ter- 
minal, subglobular, many-flowered head; bracts much smaller than the leaves; flowers 1- 
1 .5 cm long, very short-pediceled; calyx 5-8 mm long, tubular-campanulate, mostly sparse- 
ly gray -hairy, pale green; teeth 1/5-1/4 as long as the tube, terminating in a short subulate 
point, cihate on the margin, 5-nerved; corolla whitish-sulfureous, the 5-lobed lip sparingly 
pubescent outside, the upper lobes with network of pale purple nerves; nutlets ovoid, 
dark brown, impressed-punctate. June— August. 

Stony and rocky places (limestone), needlegrass-dominated areas. — European part: M. 
Dnp. (western part), U. Dns., Bes. Gen.distr.: Centr. Eur. (southeastern part). Described 
from Hungary and Transylvania. Type in Vienna. 

Note. This species differs much more sharply from the genuine T. montanum L. 
68 (not encountered in the USSR) and from the two preceding species than do all these three 
species from each other. Possibly, T. pannonicum will have to be separated into a series 
which, we presume, would also include the Balkan T. scropili Velen. 



Series 2. Polia Juz. — Flowers 5-8 mm long; leaves toothed. 



21. T. poUumL.Sp.pl. (1753)566; Ldb. Fl. Ross. Ill, 445; Shmafg. Fl. 11, 348; N. 
Pop. in Mat. Fl. Kavk. IV, 3, 42. - T. polium 7. vulgare Benth. Lab. gen. et sp. (1835) 
685. - T. angustifolium Benth. 1. c. 686. - Polium album Mill. Card, Diet. ed. VIII 
(1768) N0.4. - Ic: Hayne Arzn.-Gew. 8, tab. 5; Pflanzenfam. IV, 3a, 211, fig. 73 C; 
Fedch. and Fl. Fl. Evrop. Ross. Ill, fig. 708 (malum). — Exs.: Lang et Szovits, Herb. ruth. 
No. 92; Rehm. et Wol. Fl. pol. No. 227; GRF,No.581; Novopokr.Gerb.donsk.fi. 
No. 147; Sintenis, It. transcasp.-pers. 1900-1901, No. 26. 



46 



Subshrub with woody root, strongly branching from woody base; stems many, ascend- 
ing at base and often strongly flexuous, rarely suberect, 5-40 cm long, terete, tough, dense- 
ly short-appressed-tomentose, grayish, gray or sometimes snow-white, rarely yellowish or 
reddish (mostly in lower part), short-paniculate or often distally corymbiform, the inter- 
nodes varying greatly in length; leaves half to twice the length of internodes, 0.5-3.5 cm 
long, 1 .8 mm wide, linear, oblanceolate or oblong, cuneate at base, obtuse at apex, shallow- 
ly crenate-dentate or in lower part (1/3-2/3) entire, very often revolute-margined, frequent- 
ly somewhat wrinkled, sessile, with vesture as on stems (denser beneath), green or grayish 
above, grayish or gray beneath; inflorescences many at ends of stems and branches, short- 
peduncled, comprising few very crowded verticillasters, densely capitate, globular or ovoid, 
0.8-2 cm across; bracts small, equaling or mostly shorter than the flowers, 3-5 mm long, 
linear or linear-spatulate, entire, revolute-margined, tomentose; flowers subsessile, 5-8 mm 
long, whitish or yellowish; calyx shortly tubular-campanulate, densely whitish- or white- 
tomentose, sulcate, the teeth subequal, short-triangular, obtuse, much shorter than the 
tube, concealed by tomentum; corolla only slightly exceeding the calyx, tomentose-villous 
outside; stamens slightly exserted. June— September. (Plate IV, Figures.) 

Dry hills, clayey and stony slopes and bluffs, rocks, taluses, chalk and limestone out- 
crops, steppes, coastal sands. - European part: M. Dnp., U. Dns., Bes., BL, Crim., L. Don, 
69 L.V.; Caucasus: universal; Centr. Asia: Ar.-Casp. (Mangyshlak), Mtn. Turkm. Gen. 
distr.: Med., Bal.-As. Min., Iran. Described from Italy and other West Mediterranean 
countries. Type in London. 

Note. The plant varies markedly in habit, vesture, shape and dentation of leaves. Its 
race composition has not been thoroughly investigated in the Soviet Union, but there is 
no doubt that widely different forms occur within the regions. This would suggest that 
the sporadically observed variability is due to differences in habitat conditions, or possibly 
we may have here some evidence of the occurrence of genuine "Jordanean" species. It 
should be mentioned that a whole series of "elementary species" of this cycle has been 
established for parts of the distribution area of T. polium s.l. outside the USSR. Many 
of them were discovered long ago by Miller. Jordan himself also had a part in the analysis 
of this cycle. Very many, if not the great majority, of the West European "species" of the 
cycle are cited by Popov (op. cit., 42-43) as synonyms of T. polium L. Popov, however, 
went to the other extreme in declaring that T. polium "isconstant in every respect except 
for shght variations in the shape of the leaf blade" (p. 49). 

Economic importance. Used in Western Europe as far back as the 16th century in the 
treatment of various ailments and has been cultivated in gardens. 



Genus 1242.* Amethystea** L. 

L. Sp.pl. (1753)21 

Flowers small, 3-5 mm long, blue, in peduncled cymes forming a loose paniculate in- 
florescence; calyx subglobose-campanulate, regular, cut to middle into 5 equal subulate- 
lanceolate acute teeth, usually blue, densely covered with short glandular hairs; corolla 

* Treatment by E.V. Volkov. 
** Amethyst - a bluish-violet precious stone; referring to the color of the flowers. 



47 



70 



azure, ciliate, without a ring of hairs inside, 5-lobed; middle lobe of lower lip larger than 
the others, obovate, broader at front; other lobes equal, acuminate; stamens 2, inserted 
near base of lower lip, the upper sterile stamens reduced to short slender filaments without 
anthers; nutlets obovoid, convex, rugose, with curved lower margin, brown, 2 mm long. 
Annual herbs, with erect branching stem; leaves 3-5-partite, with lanceolate lobes; upper 
leaves very small. 



1. A.coeruleaL. Sp.pl. 1(1753) 21; Bge. in Ldb. Fl. Alt. I, 19; Ldb. Fl. Ross. Ill, 
441 ; Turcz. Fl. baic.-dahur. II, 435; Kom. Fl. Man'chzh. Ill, 336; Kom. and Alls. Opred. 
rast. Dal'nevost. kraya, II, 895; Kryl. Fl. Zap. Sib. IX, 2294. 

Annual; stems 10-20 cm long, usually bluish, erect, branching, 4-angled, covered with 
very short glandular hairs, more profusely so at internodes and in uppermost part; leaves 
2-6 cm long, 1-2 (3) cm wide, cuneately tapering to short petiole, deeply 3-5-sect into 
lanceolate, dentate lobes, the terminal larger than the lateral, the upper side smooth or 
with sparse and very short hairs, the lower side paler; flowers sparingly pubescent, in 
cymes forming a small paniculate inflorescence; peduncles as long as or longer than leaves; 
calyx 2-3 mm long, 1 .5 mm wide, campanulate, usually bluish, cut to the middle into 5 
subulate-lanceolate acute teeth, rather densely punctate with short-stipitate glands; corolla 
longer than calyx, 3-3.5 mm long, blue, 5-lobed, ciliate at margins; stamens 2, exserted; 
nutlets brown, rugose, 2 mm long. July August. 

A weed, ubiquitous in inhabited areas, also pastures, river banks, stony slopes and 
taluses. - Centr. Asia: Dzu.-Tarb.,T. Sh.; West Siberia: Alt.; East Siberia: Ang.-Say., 
Dau.; Far East: Uss. Gen. distr.: Iran., Mong., China (Manchuria). Described from 
Siberia. Type in London. 



Tribe 2. ROSMARINEAE Briq. in Pflanzenfam. IV, 3a (1895) 207 et 216. - Calyx 
12-nerved, 2-hpped, according to ^/j formula; corolla 2-lipped, with concave upper lip 
and 3-parted lower lip; stamens 2 (the lower), attached under upper Up, filaments with 
a tooth at base; antlrers 1-celled; nutlets obovoid, with ventral surface of attachment. 



Genus 1243.* Rosmarinus** L. 

L. Sp.pl. (1753)23. 

Calyx ovoid-campanulate, 8-r2-nerved, 2-lipped, the upper lip with 3 short teeth, the 
lower 2-toothed; corolla blue-violet (rarely wliite); upper lip emarginate or 2-fid; lower 
lip 3-lobed, the large middle lobe strongly concave and dechned; stamens 2, with 1-celled 
anthers; nutlets smooth. Shrub, with evergreen entire leaves. 

* Treatment by B.K. Shishkin. 

** The derivation of the name Rosmarinus is unclear. Some authors attribute it to Greek rhops, low 
shrub, and myrinos, balsam shrub. Others suggest a less likely derivation from the Liitin ros, dew, 
and marinus, of the sea, marine. 



48 



71 The genus contains four or five species native to the Mediterranean region. It is consid- 

ered as monotypic by many taxonomists. 

1. R. officinalis L. Sp. pi. 0753) 23. - R. angustifolius Mill. Card. Diet. ed. VIII 
(1758)No. l.-R. latifolius Mill. I.e. (1758) No.2. -R. officinalis vulgaris Alefel, 
Landwirt. Fl.(1860) 186. - R. officinalis var. genuina Turill in Kew Bull. (1920) 105. 
- Salvia rosmarinus Schleid. Handb. Med.-Pharm. Bot. I (1852) 265. ~ Ic: Barlikh. 
Russk. lekarstv. rast. IV, Plate 98 (1900); Komarov, Sbor, sushka i razved. lekarstv. rast. 
izd. 3-e, Plate 64 (1917). 

Shrub, 50-200 cm high, strongly branching; branches ascending or erect, covered with 
gray, easily peeling bark; young branches obtusely 4-angled, pubescent, with axillary fas- 
cicles; leaves on very short petioles, evergreen, linear, obtuse, revolute-margined, rather 
thick, dark gre^n above, subglabrous (with solitary minute stellate hairs), white-tomentose 
beneath, 1 .5-3.5 cm long, 1 .5-3 mm wide; flowers subsessile, in 5-10-flowered racemiform 
inflorescences at the end of short shoots; calyx gray-tomentose, with glabrous throat; co- 
rolla blue-violet, sparingly pubescent outside, the tube slightly exceeding the calyx; upper 
lip deeply emarginate; lower lip slightly longer, the large middle lobe toothed at margins; 
stamens exserted; nutlets globose-ovoid, 1.5-2 mm long, smooth, brownisu. /.pril-May. 

The plant is distributed in the Mediterranean region from southern France and North 
Africa (Algeria, Tunisia) to Asia Minor. It is well known in England, France, Italy and 
other countries for its volatile oil. 

In the USSR it is cultivated in the Crimea and Caucasus. It is native to the West Medi- 
terranean floristic region where it is often a component of maquis vegetation. 

Economic importance. Oil of rosemary, esteemed by the ancient Greeks and Romans, 
is obtained from the leaves and flowering shoots which are clipped up to three times a 
year. The yield of oil varies from 0.19 to 1 .7% (up to 2% from some leaves) depending 
on origin and management. The composition of the oil is complex, the main component 
being pinene (up to 80%). Oil of rosemary is used in many brands of eau de Cologne; it 
is also known as moth-repellent. 



Subfamily 2. SCUTELLARIOIDEAE Briq. in Pflanzenfam. IV, 3a (1895) 207 et 224. 
— Calyx 2-lipped, both lips entire; corolla 2-Upped, the upper galeate; stamens 4, ascending 
under the upper lip, the lower longer than the upper; ovary 4-parted; nutlets depressed- 
globose, with dry exocarp, basally attached to receptacle (gynophore); surface of attach- 
72 ment small, orbicular; ovules amphitropous; seeds transverse; embryo with a bent radicle 
lying on one cotyledon. Herbs, seldom shrubs. 

Note. The characteristic position of the ovules and shape of the embryo, which do not 
occur in any other members of the family, prompted segregation of this group as a family 
(Caruel. Sur la nouvelle famille des Scutellariacees. Bull, de la Soc. Bot. de France, tab. 
XXXIII, 1886, 266-268). In the Soviet literature, a similar approach was adopted by 
Nevskii. However, considering that this group does not otherwise differ from other Labia- 
tae (the other features of Scutellarioideae are to be found in various genuine Labiatae), we 
treat it, in agreement with Briquet, as a subfamily, all the more so since its extent has not 



49 



been finalized to this day. Camel included in this group the genera Scutellaria L., Sala- 
zaria Torr., Perilomia H.B.K. and (provisionally) Catopheria Benth.; Briquet admitted 
only the first two. In his classification, Briquet assigns to Perilomia the status of a tribe, 
Perilomieae, and attaches it to the subfamily Stachydoideae, but he himself expresses doubt 
as regards the taxonomic position of one of the sections of Perilomia Kunth., namely 
Aprosphyla Briq. (species of the latter have been included in the genus Scutellaria by 
some authors). 



Genus 1244.* Scutellaria** L. 

L. Sp.pl. (1753)598. 

Flowers soUtary, in axils of undifferentiated or more or less differentiated bracts, in the 
latter case forming racemose or spicate inflorescences; calyx campanulate, dorsoventrally 
flattened, 2-lipped, with entire broadly rounded lips that are closed in fruit, usually split- 
ting into two unequal flapUke parts, the lower persistent and the upper falling away (some- 
times both persistent or both deciduous); upper part of calyx frequently bearing a rounded 
and usually concave scalelike appendage (fold), a so-called scutellum, as long as or longer 
than the remaining part of the upper "flap," rarely of different shape or obsolete; coroUa- 
73 tube long, mostly exserted, curved upward, dilated above; Umb of corolla 2-lipped; upper 
lip ascending to upright, strongly inflated, galeate, entire or emarginate, the lateral lobes 
freeor adnate to the upper rather than the lower lip; lower hp broad, flat, shorter or longer 
than the upper; stamens 4, didynamous, ascending, the ciliate anthers approximate in pairs; 
the lower longer, 1 -celled by partial abortion, the upper with 2 divergent cells; disk elon- 
gating into columelhform or turbinate, erect or curved gynophore, this bearing 4 nutlets 
and an unevenly 2-cleft style with very short upper lobe; nutlets depressed-globose or 
ovoid, mostly tubercled, often pubescent, rarely smooth; embryo with curved radicle. 
Perennial or very rarely annual herbs, subshrubs, seldom shrubs, mostly odorless, the ves- 
ture of simple hairs often interspersed with glandular. Leaves mostly petiolate, varying in 
shape, entire or toothed. 

Note. A very large and polymorphic genus, usually separated into two subgenera, Eu- 
scutell^ria Briq. and Scutellariopsis Briq. The latter does not occur in the USSR, and 
thus all Soviet species might be expected to belong exclusively to Eu-scutellaria. How- 
ever, investigation of Central Asian representativesofthegenusrevealsanumber of unusual 
types that fail to conform to the characteristics of any of the sections of this subgenus, 
and it would take some very forced interpretation to justify their inclusion. We were, there- 
fore, compelled to estabUsh three new subgenera of Scutellaria. With a more fragmentiz- 
ed approach to the category of genus, these subgenera could indeed be raised to generic 
status with no less vaUdity than the section Heteranthesia Briq., of the subgenus Eu- 
scutellaria (not represented in the USSR in wild state) which Grossgeim (in Izv. Azerb. 
fil. AN SSSR, No. 3, 1945, 89) proposed to set up as a separate genus. In fact, generic 

* Treatment by S.V. Yuzepchuk. 

** From the Latin scutellum, dish, referring to the shape of the appendage in the upper part of the 
calyx of most of the species of the genus. The Russian name of this genus, "shlemnik" [shlem = 
helmet], does not correspond to the now accepted Latin name, but to an older one - Cassida 
(from cassis or cassida, helmet) which is mentioned by Bock. 

50 



names have already been proposed in the literature for two of the subgenera in question. 
We refrain from such treatment, taking into consideration that all these groups are appar- 
ently derived from the subgenus Eu-scutellaria; being relatively recent, they do not 
deviate markedly from the genus Scutellaria s. str. 

Economic importance. It is interesting to note that, v^hile most of the Labiatae are 
plants yielding volatile oil and are mainly used as such, species of Scutellaria are a strik- 
ing exception to the rule. Most of them are presumably dye plants but very few have been 
investigated and evaluated in this respect. 
74 Many scullcaps are ornamental, but again few of the Soviet species have been introduced 
into cultivation. Some species are of medicinal value. 

See also notes to individual species. 

Key to Subgenera 

1 . Upper part of calyx with a dorsal appendage 2. 

+ Upper part of calyx unappendaged, sometimes with a small bulge on back ... 3. 

2. Appendage on the upper (deciduous) part of calyx forming a concave or flat fold, a 
so-called scutellum Subgenus 1. Euscutellaria Briq. (p. 51). 

+ Appendage on the upper part of calyx globose, bladderlike 

Subgenus 2. Cystaspis Juz. (p. 133). 

3. Flowers in a lax secund inflorescence; calyx not concealed by bracts, entire, early 
deciduous, scarious Subgenus 3. Anaspis(Reching. fil.) Juz. (p. 138). 

+ Flowers in dense quadrilateral inflorescences; calyx concealed by bracts, long persis- 
tent, conchoidal Subgenus 4. Apeltanthus (Nevski) Juz. (p. 144). 



Subgenus 1. Euscutellaria Briq. in Pflanzenfam. IV, 3a (1896) 225, prosect. - Flowers 
variously disposed; calyx accrescent, in fruit, both lips scarious or coriaceous, unequal, the 
appendage on the upper hp (scutellum) arched-concave, rounded at apex, functioning, 
at least in many species (of the sections Stachymacris andLupulinaria),as a kind of lever 
which, in maturity, causes the upper part of the calyx to break away at the slightest touch, 
thus promoting dispersal of the nutlets. Herbs (perennial or rarely annual), subshrubs or 
sometimes shrubs of very varied habit. 

1 . Flowers turned to one side to form a secund inflorescence; bracts herbaceous . 2. 
+ Flowers in quadrilateral inflorescences; bracts more or less membranous, rarely sub- 
herbaceous 25. 

2. Flowers in the axils of normal leaves which gradually diminish in size toward the 
summit, or forming a racemose inflorescence and then bracts leaflike though some- 
what smaller 3. 

+ Bracts distinguished from the crenate-dentate cauUne leaves by their small size and 
entire margin 18. 

3. Low, procumbent, West Caucasian high-mountain plants, with purple flowers; bracts 
75 entire, markedly smaller than the obscurely crenate or entire cauUne leaves . . 4. 

+ Plants different 5. 



51 



4. Leaves approximate, glabrous or ciliate in lower part; bracts ciliate-hairy along up- 
per margin; calyx shortly stipitate-glandular in addition to being sparsely hirsute . 
23. S.pontica C.Koch. 

+ Leaves distant, diffusely hairy above and on the veins beneath, the margin long- 
ciliate; bracts resembling cauhne leaves but more densely hairy; calyx densely 
hispid, the bristles interspersed with stipitate glands . . . . .24. S. helenae Alb. 

5. Leaves lanceolate, mostly attenuate at base, entire, firm, frequently subcoriaceous, 
glabrous or sparingly pubescent above, foveolate-punctate beneath; flowers rather 
large (2-2.5 cm long), forming a racemose inflorescence, the bracts shghtly smaller 
than the cauline leaves 14. S.baicalensis Georgi. 

+ Leaves differing in at least some characters, flowers axillary, the subtending bracts 
not differing in any way from cauline leaves, gradually decreasing in size. ... 6. 

6. Leaves not foveolate-punctate beneath, usually widest in lower part, gradually atte- 
nuate above, mostly subacute 7. 

+ Leaves foveolate-punctate beneath, usually widest at middle, with nearly parallel . 
margins, mostly obtuse 15. 

7. Rhizome monihform, consisting of white articulated tubers 

13. S. moniliorrhiza Kom. 

+ Rhizome not moniliform 8. 

8. Leaves with strongly developed lower teeth, at least some hastate 9. 

+ Leaves not hastate 11- 

9. Flowers large, 13-22 cm long; pedicels eventually declined, not drooping ... 10. 
+ Flowers very small, ca. 5 mm long; pedicels eventually drooping 

8. S. dependens Maxim. 

10. Leaves (at least the middle ones) distinctly hastate at base, often with 1-2(4) obtuse 
teeth on basal lobes, otherwise entire 6. S. hastifolia L. 

+ Leaves unevenly crenate-dentate below, with 2-10 teeth on each margin, entire 

above 7. S. dubia Tal. et Schir. 

11. Flowers usually ca. 1.5 cm, sometimes to 2 cm long. . .• 12. 

+ Flowers larger, to 2.5 cm long 13. 

12. Leaves dark green, mostly shallowly crenate-dentate, usually sparsely hairy beneath, 
sometimes glabrate 1. S. galericulata L. 

+ Leaves pale or bright green, rather deeply crenate-dentate in lower part, densely cov- 
ered beneath with short crisp hairs, often nearly velutinous (Far Eastern form) . . 
2. S. krasevii Kom. et L Schischk. 

13. Leaves 1-2 cm wide, usually broadly lanceolate or oblong-ovate, with 4-14 well 
developed teeth 3. S. yezoensis Kudo. 

+ Leaves to 1 cm wide, lanceolate, entire or with few shallow crenations in lower 

part 14. 

14. Plants 20-60 cm high; leaves densely short-hairy on both sides 

4. S. regeliana Nakai. 

+ Plants 10-35 cm high; leaves glabrous or with scattered hairs above, short-hairy on 

the veins and margins, with scattered small granular glands beneath 

5. S. ikonnikovii Juz. 

15. Leaves glabrous or sparingly hairy 16. 



52 



+ Leaves very hairy 17. 

16. Stem simple or branching; the branches rather short, declined; leaves not many, 
usually not attenuate, obtuse; flowers 1.8-2.2 cm w^ide . . 9. S. scordiifolia Fisch. 

+ Branches very long, erect; leaves numerous, widest at base and gradually attenuate 
above, often acutish; flowers small, 1.4-1.7 cm long . . . .12. S. poly phy 11a Juz. 

17. Stems mostly divaricately branching from base; leaves usually elliptic, rarely ovate 
or oblong, obtuse or rounded at apex, with slightly developed blunt teeth, densely 
hairy on both sides 10. S. taquetii Levi, et Vaniot. 

+ Stems usually simple or few-branched; leaves ovate or oblong-ovate, subacute to 
subobtuse at apex, with well developed sharp teeth, rather densely hairy on both 
sides. . 11. S. tuminensis Nakai. 

18. Bracts generally shorter than calyx; leaves rather thin-textured, deeply and coarsely 
crenate-dentate 19. 

77 + Bracts as long as or longer than calyx; leaves mostly firm, rather shallowly and 

usually finely crenate 23. 

19. Bracts ovate, rounded at base (European and Caucasian plants) 20. 

+ Bracts lanceolate, tapering at both ends (Far Eastern plants) 21. 

20. Corolla bicolor, whitish, with violet-blue upper lip; stems and leaves sparingly hairy 
15. S.altissimaL. 

+ CoroUa purple; plant wholly glabrous 16. S. tournefortii Benth. 

21. Flowers to 1 cm long (Sakhalin) 19. S. shikokianaMak. 

+ Flowers twice as long (Ussuri) 22. 

22. Leaves glabrous or with scattered hairs on the whole surface above, only on the veins 
beneath; calyx sparsely hairy 17. S. ussuriensis (Rgl.) Kudo. 

+ Leaves more or less hairy on both sides; calyx rather densely hairy 

18. S. pacifica Juz. 

23. Flowers whitish 24. 

+ Flowers dingy mauve 22. S. woronowii Juz. 

24. Stems and petioles covered with short, curved, recUnate or appressed hairs . . . . 
20. S. albidaL. 

+ Stems and petioles with long patent hairs 21. S. pallida M.B. 

25. Central Asian subshrubs; leaves small, entire, hairy, short-petioled; bracts leaflike 
but smaller 26. 

+ Bracts well differentiated, mostly scarious 28. 

26. Scutellum small, concave, raised across the upper lips of the calyx; leaves entire. 27. 
+ Scutellum large, flat, fringing the upper Up of the calyx; leaves finely sinuate-dentate 

27. S. oris tataM. Pop. 

27. Flowers solitary in axils of upper leaves, brownish-purple .25. S. colpodea Nevski. 

78 + Flowers in small subcapitate 3-7-flowered axillary inflorescences, creamy-yellow, 

purple-streaked or spotted 26. S. striatella Gontsch. 

28. Verticillasters very distant 29. 

+ Verticillasters closely approximate, forming a spicate inflorescence 33. 

29. Bracts ovate or lanceolate, usually acute, navicular, thinly tomentose 30. 

+ Bracts ovate or broadly ovate, obtuse or short-acuminate, nearly flat, covered with 

simple and glandular hairs 31. 



53 



30. Leaves glaucescent beneath with thin tomentum; bracts lanceolate, long-acuminate, 
usually all navicular 28. S. navicularis Juz. 

+ Leaves green on both sides; bracts ovate, rather short-acuminate, the lower often 
not navicular 29. S. chenopodiifolia Juz. 

31. Corolla yellow, not anthocyanin-colored 30. S. juzepczukii Gontsch. 

+ Corolla with intensely anthocyanin-colored lower lip 32. 

32. Lower lip of corolla nearly black-purple; calyx strongly accrescent in fruit, to 1 cm 
across 31. S. litwinowii Bornm. et Sint. 

+ Lower lip of corolla bright blue; calyx slightly accrescent in fruit 

32. S. luteo-coerulea Bornm. et Sint. 

33. Inflorescences 1-2 cm long, densely capitate. Subshrubs, with virgate stems and 
very remote small leaves 34. 

+ Inflorescences larger 37. 

34. Low, almost pulvinate subshrubs, with stout upright stems 7.5-10 cm high. . . . 
36. S. squarrosa Nevski. 

+ A higher and slimmer fruticose plant, with slender virgate stems 10-50 cm high 35. 

35. Stems covered all the way up with very short, curved, closely appressed hairs and a 
faint grayish bloom 34. S. intermedia M. Pop. 

+ Vesture of stems different 36. 

36. Stems short-hairy only in lower part, glabrous elsewhere; leaves narrow, lanceolate; 
flowers 2-2.5 cm long 33. S. ramosissima M. Pop. 

+ Stems glabrate or with sparse short hairs usually interspersed with longer ones; 
79 leaves wider, ovate or oblong; flowers 1 .5-2 cm long . . . .35. S. glabrata Vved. 

37. Leaves (at least initially) tomentose, white or gray beneath (and sometimes above) 
38. 

+ Leaves without tomantum, green on both sides or, if heavily pubescent, grayish- 
green 95. 

38. Bracts not scarious, rather thinly tomentose, essentially eglandular, usually lanceo- 
late or narrowly ovate, acute, navicular with pronounced keel, rarely wider and 
nearly flat, very rarely obtuse 39. 

+ Bracts usually scarious, rarely somewhat tomentose but invariably with numerous 
stipitate or subsessile glands, ovate, obtusish to acuminate, generally not distinctly 
navicular, at most slightly carinate 60. 

39. Bracts eglandular 40. 

+ Bracts with some glandular hairs 56. 

40. Flowers yellow, sometimes with greenish, dark yellow or brownish spots on lower 
lip or on lateral lobes of upper Up 4L 

-I- Flowers at least partially anthocyanin-colored 48. 

41. Crimean-Caucasian herbaceous plants 42. 

+ Central Asian subshrubs 47. 

42. Leaves oblong or lanceolate, cuneately tapering at base; petioles less than half as 
long as the blade 37. S. araxensis Grossh. 

+ Leaves ovate, with truncate base; petioles only slightly shorter than the blade . 43. 

43. Bracts rather narrowly lanceolate, navicular; leaves ovate 44. 

+ Bracts broadly lanceolate, not navicular; leaves oblong-ovate 46. 



54 



44. Stems and underside of leaves with patent short hairs 38. S. taurica Juz. 

+ Stems and underside of leaves appressed-tomentose 45. 

45. Leaves ovate; bracts lanceolate, acute 39. S. stevenii Juz. 

+ Leaves broadly ovate; bracts broadly lanceolate, acute or short-acuminate .... 

40. S. tatianae Juz. 

46. Bracts rather large, the lower 9-1 2 mm long 41. S. raddeana Juz, 

+ Bracts small, the lower 5-7 mm long 42. S. leptostegia Juz. 

47. Bracts lanceolate, navicular, acute 50. S. comosaJuz. 

+ Bracts ovate, concave but not navicular, obtuse 52. S. microdasys Juz. 

48. Subshrubs; leaves rather large; flowers essentially yellow, only the upper Up and its 
lateral lobes anthocyanin-tinged 49. 

+ Herbs; leaves small, up to 1 cm long, densely gray-tomentose on both sides; flowers 
mostly purple, but if yellow then spotted or streaked with purple on lower lip and 
on tube 50. 

49. Leaves green or grayish-green above, rather sparingly pubescent, white- or gray- 
tomentose beneath; bracts broadly lanceolate below, abruptly cuspidate, long- 

hairy-ciliate on the margin, otherwise fmely tomentose, all entire 

51. S. oxystegia Juz. 

+ Leaves grayish-green on both sides with rather dense tomentum; bracts ovate, acute, 

rather sparsely tomentose, the lower mostly with solitary teeth 

53. S. androssovii Juz. 

50. Corolla completely an thocyanin-colored 51. 

+ Corolla bicolor or variegated 53. 

51. Bracts ovate, not navicular 46. S. karkaralensis Juz. 

+ Bracts lanceolate, navicular 52. 

52. Leaves shallowly incised-crenate, with short teeth; corolla purple 

43. S.grandiflora Sims. 

-I- Leaves deeply incised-crenate-dentate, with oblong teeth; corolla blue 

44. S. tuvensis Juz. 

53. Bracts ovate, not navicular; corolla purple or yellow .... 45. S. turgaica Juz. 
+ Bracts lanceolate, navicular 54. 

54. Leaves villous-tomentose on both sides; corolla pink or purple, with a yellow spot 
on lower lip 47. S. nepetoides M. Pop. 

+ Leaves appressed-tomentose on both sides; corolla generally yellow, with purple 
and red spots and stripes on both Ups and on the tube 55. 

55. Stems softly patent-hairy; flowers to 2 cm long, cream-colored . .48. S. picta Juz. 
+ Stems covered with short curved hairs; flowers not more than 1 .5 cm long, dingy- 
yellow 49. S. flabellulata Juz. 

56. Stems patent-hairy; leaves in maturity very sparingly tomentose beneath, nearly 
green 57. 

+ Stems appressed-tomentose; leaves with distinct whitish or grayish tomentum 

beneath 58. 

57. Stems numerous (20-25), suberect; leaves crenate .... 55. S. kurssanoviiPavl. 
+ Stems usually fewer, ascending; leaves deeply cut 56. S. titovii Juz. 



55 



58. Central Asian subshrubs, with ascending or prostrate stems and deeply toothed 

leaves; flowers essentially yellow, but the upper lip connnonly anthocyanin-tinged 

at apex 59. 

+ Caucasian pereiniial, with erect stems and rather shallowly incised-crenate leaves; 

flowers yellow, not anthocyanin-tinged 7^^. S. rhomboidalis Grossh. 

50. Stems ascending, green or faintly colored; bracts to 1-2 cm long; flowers 2-2.8 cm 

long, with rather wide tube 54. S. mesostegia Juz. 

+ Stems prostrate, strongly anthocyanin-colored; bracts to 7 mm long; flowers 1.5- 

2.5 cm long, with very slender tube 57. S. catharinae Juz. 

60. Flowers pink. High-mountain plant of Talass Ala-Tau. . . . 1 14. S. talassica Juz. 

+ Flowers yellow or sometimes bicolor (yellow and anthocyanin) 61. 

b\. Glands on the bracts usually a nunor component of vesture, very small, usually dis- 
cernible only under high magnification, short-stipitate, sometimes sessile. Herbs or 
subshrubs of Crimea-Caucasus, West Siberia or Central Asia (northern regions) . 62. 
+ Glands on the bracts usually a dominant component of vesture, large, long-stipitate, 
82 easily perceived. Subshrubs of Central Asia, West Tien Shan and Pamir-Alai . . 87. 

62. Leaves rather shallowly crenate or dentate, the incisions not usually extending more 
than half way to midrib 63. 

+ Leaves pinnately incised-dentate, the incisions usually extending more than halfway 
to midrib 81. 

63. Bracts with varied vesture, the lower with short appressed hairs, others with long 
spreading hairs interspersed with subsessile glands; stems elongate, suberect, slender, 
more or less virgate; leaves very remote, small, shallowly incised-crenate with numer- 
ous small teeth (5-0 on each margin) 72. S. granulosa Juz. 

+ All bracts with same type of vesture. Plant with different habit (only S. daghesta- 
nica recalling the habit of S. granulosa, but its leaves more deeply incised-crenate) 

^ ^ 64. 

t»4. West Siberian and Baltic plant; stems covered with long, horizontally spreading 
hairs; loaves rather narrowly cuneate at base, sparsely pubescent beneath .... 
63. S. sieversii Bge. 

+ Plants differing in at least some characters 65. 

65. Bracts with short, curved, generally appressed hairs and sessile glands (Caucasian 
plants) 66. 

+ Bracts with long spreading hairs and stipitate glands 71. 

66. Ciscaucasian species; bracts ovate, long-pointed at apex 67. 

+ Transcaucasian species; bracts broadly ovate, scarcely longer than wide, short- 
pointed at apex 68. 

67. Leaves oblong-ovate or narrowly triangular, with 5-13 teeth on each margin . . . 
68. S. polyodon Juz. 

+ Leaves ovate or broadly ovate, with 3-8 teeth on each margin 

70. S. oschteiiica Juz. 

68. Bracts predominantly dentate 78. S. karjagiiiii Grossh. 

+ Bracts entire or only the uppermost dentate 69. 

60. Leaves thinly wliite-tomentose beneath with appressed hairs, the tomentum conceal- 
ini: the veins 73. S. orientalis L. 



56 



+ Leaves softly gray-tomentose beneath with spreading hairs, the network of veins 
usually distinct, not concealed by tomentum 70. 

70. Leaves incised-crenate or dentate with incisions less than half the half-blade width; 

leaves and bracts covered above with sparse appressed hairs 

75. S. platystegia Juz. 

+ Leaves deeply incised-dentate with incisions usually not less than half the half-blade 

width; leaves and bracts densely appressed-hairy above 

76. S. grossheimiana Juz. 

71 . Crimean-Caucasian plants, with short spreading hairs on stems and leaf petioles and 
frequently on the lower side of leaves 72. 

+ Stems, petioles and the lower side of leaves appressed-tomentose 74. 

72. Crimean-Novorossisk plants with incised-crenate or incised-dentate leaves, the inci- 
sions usually less than half the half-blade width 73. 

+ Transcaucasian plants, with pectinately incised leaves, the incisions at least half the 
half-blade width 74. S. sedelmeyerae Juz. 

73. Leaves broadly (wate or suborbicular, truncate at base, obtusely incised-crenate, the 
incisions 1-4 mm; bracts broadly ovate, acuminate or subacuminate, very densely 
glandular 66. S. hirtella Juz. 

+ Leaves often ovate, obtusely angled at base, more deeply incised, with mostly acute 
teeth; bracts narrower, usually ovate, more strongly acuminate at apex, faintly 
glandular 67. S. novorossica Juz. 

74. Bracts up to I cm long; flowers 1 .5-2.5 cm long 75, 

+ Bracts 1 .5-2 cm long; flowers 2.2-4 cm long 77. 

75. Dagestan plant, with suberect, elongate, more or less virgate stems; leaves usually 
1/4 as longas internodes 71. S. daghestanica Grossh. 

+ Crimean species, with rather short ascending stems; leaves not shorter than inter- 
nodes 76. 

76. Leaves ovate, to 2 cm long, with somewhat uneven but not spreading teeth; inflores- 
cence relatively few-flowered, to 10 cm long in fruit; bracts acuminate; flowers al- 
ways yellow 64. S. hypopolia Juz. 

+ Leaves suborbicular, usually not more than 1 .5 cm long, with slightly spreading 
teeth; inflorescence many-flowered, to 15 cm long in fruit; bracts obtuse or very 

short-acuminate; flowers pure yellow or purple or yellow-purple 

65. S. heterochroa Juz. 

77. Caucasian high-mountain plant; leaves subcordate or truncate at base, shallowly 
incised-crenate 69. S. oreophila Grossh. 

+ Central Asian plant; leaves truncate or broadly cuneate at base, deeply incised- 
dentate 78. 

78. Bracts broadly ovate, short-acuminate, with short appressed hairs only on margins 
and veins, rarely all over, covered with very small stipitate glands; stems usually 
strongly anthocyanin-colored 62. S. krylovii Juz. 

+ Bracts narrowly ovate or ovate, mostly long-acuminate, rather densely covered with 
long spreading hairs and few stipitate glands; stems usually green or faintly antho- 
cyanin-colored 79. 

79. Leaves triangular-ovate or oblong, subacute; bracts acuminate 80. 



57 



+ Leaves shorter, rather broadly ovate or elHptic, obtuse; bracts subacute, not acumi- 
nate 59. S. alberti Juz. 

80. Leaves large, with uneven, mostly subacute teeth; bracts usually long-acuminate; 
flowers to 3.7 cm long 58. S. transiliensis Juz. 

+ Leaves small, with even obtuse teeth; flowers 0.5-1.5 cm long 

61. S. soongorica Juz. 

81. Bracts very faintly glandular, even the lowermost entire (Tien Shan) 

60. S. przewalskii Juz. 

+ Bracts distinctly glandular, often slightly dentate, especially the lowermost (Trans- 
caucasian) 82. 

82. Leaves mostly dissected scarcely more than half way to midrib, the lobes usually 
slightly narrower than the undissected part of blade 83. 

+ Leaves mostly dissected more than half way, often nearly to midrib, the lobes usual- 
ly as wide as or even wider than the undissected part of blade 86. 

83. Bracts rather sparsely covered with short, closely appressed hairs and subsessile 
glands 73. S. orientalis L., forma. 

+ Bracts rather densely covered with long spreading hairs and stipitate glands . . 84. 

84. Leaves with horizontally spreading or even recurved lobes, appressed-hairy above, 
softly tomentose with patent hairs beneath, the network of veins not fully con- 
cealed by the tomentum 77. S. artvinensis Grossh. 

+ Leaves with obhquely ascending lobes and different vesture, the lateral and second- 
ary veins beneath concealed by the tomentum 85. 

85. Leaves predominantly covered on both sides with patent glandular hairs, these often 
completely concealing the sparse tomentum beneath . . 80. S. prilipkoana Grossh. 

+ Leaves with short appressed hairs above, sparsely tomentose beneath, without patent 
glandular hairs 84. S. araratica Grossh. 

86. Bracts at least partly dentate 81. S. darriensis Grossh. 

+ Bracts entire 82. S. sosnovskyi Takht. 

87. Rather large, large-leaved mountain plants with bracts to 1 cm long 88. 

-I- Low-growing, small-leaved, high-mountain plants with bracts usually less than 1 cm 

long ■. 93. 

88. Leaves generally not anthocyanin-tinged 89. 

+ Leaves at least partly anthocyanin-tinged beneath 91. 

89. Leaves green or grayish-green with scattered hairs above, white- or gray-tomentose 
beneath, rather evenly crenate-dentate, with 4-10 crenations on each margin . . 90. 

+ Leaves green at both sides, glabrous above, faintly tomentulose beneath, coarsely 
crenate-dentate, with 5-7 uneven obhquely antrorse teeth on each margin .... 
90. S. urticifolia Juz. 

90. Leaves deeply and finely crenate-dentate, densely tomentose beneath, the network 
of veins completely concealed by the tomentum; flowers 3-3.5 cm long .... 
85. S. adenostegia Briq. 

+ Leaves rather shallowly and coarsely crenate, loosely tomentose beneath, with clear- 
ly discernible network of veins 86. S.bucharica Juz. 

91. Plants of West Tien Shan; leaves with large, rather long subacute teeth . . . .92. 
+ Plants of Pamir-Alai; leaves with smaller, shorter and more obtuse teeth 

87. S. gontscharovii Juz. 

58 



86 



92. Leaves broadly ovate or ovate-rhombic, rounded or broadly cuneate at base; bracts 
ovate to broadly ovate, at least the lower abruptly terminating in a usually recurved 
point, more or less uniformly glandular-hairy 88. S. haematochlora Juz. 

+ Leaves ovate or narrowly rhombic, cuneate at base; bracts narrowly ovate, gradually 
tapering to a mostly straight point, slightly glandular or subglabrous between veins 
89. S. tschimganica Juz. 

93. Stems with more or less spreading hairs; leaves narrowly ovate, acute 

93. S. angrenica Juz. et Vved. 

+ Stems appressed-tomentose; leaves broadly ovate, obtuse 94. 

94. Leaves deeply, coarsely and unevenly toothed; bracts subacute to subobtuse, more 
or less violet; flowers yellow with purple lateral lobes, 2.2-3 cm long (Zeravshan . 
basin) 91. S. iskanderi Juz. 

+ Leaves rather shallowly, finely and evenly toothed; bracts obtuse, not usually color- 
ed; flowers plain yellow, 1.5-2.5 cm long (West Tien Shan). 92. S. pycnoclada Juz. 

95. Low shrubs, with woody stems and branches 96. 

+ . Herbs, rarely subshrubs 99. 

96. Leafy shoots and leaves with long-stipitate glands; leaf base mostly cordate . . . 
94. S. cordifrons Juz. 

+ Leafy shoots and leaves with vesture destitute of long-stipitate glands; leaf base 

mostly broadly cuneate or truncate, rarely subcordate 97. 

97. Central Asian species; lower lip of corolla mostly slightly pubescent at base, usually 
not longer than the upper lip 98. 

+ Transcaucasian species; lower hp of corolla glabrous, longer than the upper Up . . 
97. S. sevanensis Sosn. 

98. Leaves ovate or oblong, often subacute, with acute teeth, glabrous or sparsely short- 
hairy above, usually densely covered beneath with short curved hairs and minute 

87 glands; stems and petioles usually eglandular; bracts elHptic, acute, usually entire 

except sometimes the lowermost 95. S. adsurgens M. Pop. 

+ Leaves broadly ovate or suborbicular, obtuse, with obtuse teeth, usually densely 
short-hairy above, with thickish tomentum and short-stipitate glands beneath; stems 
and petioles very often with short-stipitate glands; bracts broadly ovate, usually ob- 
tuse, mostly crenate-dentate 96. S. phyllostachya Juz. 

99. High-mountain plants of Armenia (Mount Aragats), with leaves dissected nearly to 
midrib into very narrow linear segments; flowers as in species of the Orientales 
group (i.e. with long and slender, distally expanding tube and large lower lip). . . 
83. S. amphichlora Juz. 

+ Different from above 100. 

100. Plants of Kara Tau Mountains; leaves very rigid, brittle when dry, densely covered on 
both sides with very short hairs, subvellutinous; inflorescence congested .... 
122. S. karatavica Juz. 

+ Leaves not rigid, with different vesture; inflorescence rather lax 101. 

101. Plants mesophilous in habit; leaves rather sparingly hairy, rarely with fairly profuse 
hairs of moderate length 102. 

+ Xerophilous plants; leaves rather densely covered on both sides with long, mostly 
somewhat flexuous hairs 118. 



59 



102. Rather large plants of mountains and plains; leaves copiously toothed (in more 
developed leaves 7-1 1 teeth on each margin) 103. 

+ Lov^-growing high-mountain plants of Central Asia; leaves sparingly toothed (usual- 
ly not more than 5 at each side) 113. 

103. Flov^ers essentially violet 104. 

+ Flowers essentially yellow 106. 

104. Altai plants; leaves rounded or slightly cordate at base; flowers 2-2.5 cm long, vio- 
let, with paler lower lip and tube 104. S. altaica Fisch. 

+ Leaves cuneate at base; flowers 2.5-3 cm long, of different color 105. 

105. Plants of the Tuva ASSR, with dark azure corolla and very compact inflorescence . 
105. S. mongolica Sobolevsk. 

88 + Tien Shan plants, with bicolored corolla (bright yellow tube and violet-purple lips) 
and rather lax inflorescence 108. S. xanthosiphon Juz. 

106. Leaves deeply cordate, glandular 107. S. knorringiae Juz. 

+ Leaves not cordate, rarely subcordate, eglandulose 107. 

107. Uraland Asian plants with flowers 2.2-3.5 cm long 108. 

+ European plants with flowers 1-2.5 cm long 110. 

108. Ural plants with acute leaves; stems with fine, recurved hairs 

102. S. oxyphylla Juz. 

+ Siberian or Central Asian plants with obtuse leaves; stems with spreading hairs . . 
109. 

109. Stems covered with short rechnate hairs, these often interspersed with longer spread- 
ing hairs; inflorescence short and dense, 2.5-4 cm long; bracts broad, ovate, to 

1.2 cm long, scarious, sparsely long-hairy; corolla yellow . . . 103. S.supinaL. 
+ Stems covered with short and long, horizontally spreading hairs; inflorescence 
slightly looser, to 8 cm long; bracts narrower, oblong-ovate, to 8 mm wide, sub- 
herbaceous, densely long-hairy; corolla yellow, the lower hp lilac, the upper 
lilac-tipped 120. S. alexeenkoi Juz. 

1 10. Plants of western Ukraine; bracts early deciduous; petioles and base of leaves ciliate 
98. S. vema Bess. 

+ Plants different from above 111. 

111. Stems much branched, the branches long, flexuous; upper Hp of corolla with usually 
violet-tinged lateral lobes 100. S. chitrovoiJuz. 

+ Stems simple or sparingly short-branched; flowers plain yellow 112. 

112. Narrow-leaved plants (chalks of Ukrainian SSR) 99. S. creticola Juz. 

+ Broad-leaved plants (Central Povolzhye) 101. S. cisvolgensis Juz. 

113. Flowers essentially yellow 114. 

+ Flowers essentially anthocyanin-colored 117. 

114. Leaves unevenly toothed, the teeth long, divaricate (Saur Range) 

89 106. S. irregularis Juz. 

+ Leaves rather evenly crenate-dentate, the teeth not divaricate 115. 

115. Leaves cordate at base, as wide as long or even wider (Kirgiz Alatau) 

113. S. popovii Vved. 

+ Leaves tapering or rounded at base, longer than wide 116. 



60 



90 



1 16. Leaves to 2.4 cm long, 1 .4 cm wide; stems rather densely covered with long patent 
hairs; leaves rather densely hairy beneath; corolla to 3.5 cm long (Tien Shan) . . 
109. S. oligodonta Juz. 

+ Leaves to 1 .4 cm long, 0.9 cm wide; shoots with sparse very short hairs or subgla- 

brous; leaves with scattered hairs; corolla ca. 2.5 cm long. (Pamir-Alai) 

110. S. paulsenii Briq. 

117. Bracts scarious, entire; flowers violet, with yellowish tube . . 1 1 1 . S. filicaulis Rgl. 
+ Bracts subherbaceous, toothed; flowers lilac-purple .... 112. S. kugarti Juz. 

118. Corolla yeUow, the upper hp purple-tipped, the lower lip with 2 large round dark- 
brown spots (Zeravshan) 121. S. ocellata Juz. 

+ Corolla without spots on lower Hp (Tien Shan) 119. 

1 19. Leaves (at least the lower) subcordate to cordate at base, the lateral veins arising 
mainly close to leaf base, hence venation apparently flabellate 120. 

-I- Leaves broadly cuneate at base, the lateral veins arising mainly from higher up the 
midrib, venation essentially pinnate 121. 

120. Leafy shoots densely covered with rather long spreading hairs; leaves mostly grayish- 
green on both sides, with profuse, slightly crisp or nearly straight hairs of medium 
length (Talass Ala Tau) 115. S. linczevskii Juz. 

+ Leafy shoots less densely covered with rather short hairs; leaves green on both sides, 
with scattered, short, crisp hairs (Kirgiz Alatau) 116. S. subcordata Juz. 

121. Cespitose plants; stems and petioles with rather short hairs (Kara Tau) 

117. S. subcaespitosaPavl. 

+ Non-cespitose plants; stems (at least in their lower part) and petioles densely covered 
with long hairs 122. 

122. Stems in lower part, petioles and young leaves densely lanate-villous; leaves rugose, 
partly tomentose beneath, the veins strongly impressed above and very prominent 
beneath; bracts up to 1 cm long, mostly broadly ovate ... 1 18. S. lanipes Juz. 

+ Stems in lower part, petioles and young leaves with sparser and shorter hairs; leaves 
not rugose, not tomentose beneath, the veins not strongly impressed above or pro- 
minent beneath; bracts to 1 .7 cm long, elongate-elliptic 

119. S. toguztoraviensis Juz. 



Section 1. Galericularia A. Hamilt. Monogr. Scutell. in Ser. Bull. Bot. V(1832) 31.- 
Axillares Benth. in Bot. reg. 18 (1832) ad calcem. No. 1493. — Galericulatae Boiss. Fl. 
or. IV (1879) 681. — Flowers paired in the axils of normal leaves, these remote, gradually 
smaller toward summit, forming a secund inflorescence. Mostly upright, mesophilous pe- 
rennials; leaves lanceolate, ovate or eUiptic, sessile or the lower short-petioled, entire or 
slightly dentate. A group with holarctic distribution. 



Series 1. Eu-galericulatae Juz. in Bot. mat. gerb. Bot. inst. ANSSSR, XIV (1951) 
359. — Flowers medium-sized, usually ca. 1 .5, sometimes to 2 cm long; rhizome slender; 
leaves usually shallowly crenate-dentate, not sagittate, the lower surfaces not foveolate- 
punctate. 



61 



1. S. galericulata L. Sp. pi. (1753) 599; Ldb. Fl. Ross. Ill, 398; Shmal'g. Fl. II, 327; 
Kryl. Fl. Zap. Sib. IX, 2296. — S. galericulata a. pubescens et j3. vulgaris Benth. Lab. 
gen. et sp. (1834) 437. - S. galericulata oc. genuina Rgl. Tent. Fl. Ussur. (1861) 118. - 
S. pubescens Martr. Fl. Tarn. (1864) 572. — S. rosmarinifolia Adams in herb. — S. 
adamsii Spreng. Syst. 2 (1825) 701. — Cassida galericulata Scop. Fl. earn. ed. II, 

1 (1772)430; Moench, Meth. (1794) 413. - C. major Gilib. Fl. Lithuan. I (1781) 90. - 
Ic: Rchb. Ic. Fl. Germ. XVIII, tab. 1256; Syreishch. lU. fl. Mosk. gub. Ill, 101. - Exs.: 
PI. Finl. exs. No. 901; GRF, No. 683. 

91 Perennial, with creeping branching rhizome; stems 8-50 (70) cm long, ascending or 
erect, simple or branching (sometimes strongly so), with internodes mostly shorter than 
the leaves, covered mainly on the ribs with short spreading or reclined hairs; leaves 2-6 
(7) cm long, 0.5-2 cm wide, rather firm, oblong-ovate or broadly lanceolate, truncate or 
cordate at base, gradually attenuate above, subobtuse to obtuse, shallowly and obtusely 
crenate-dentate, with 2-8 very flat, often indistinct crenations on each margin, sparsely 
hairy or glabrous, dark green above, usually covered beneath with sparse short spreading 
curved or even crisp hairs, or hairy only on margins and veins or sometimes subglabrous, 
usually not punctate, pale green, often becoming lilac (like the stems); petioles short, 
broad, with vesture as on the stems; flowers solitary in the axils of middle or upper leaves, 
on pedicels 1-2 mm long, forming a lax secund inflorescence; calyx 3-4 mm long, in fruit 
to 5 mm, as long as or slightly shorter than the pedicel, with short curved hairs (mostly 
eglandular), usually colored; corolla 1.2-2.6 cm long, violet-blue, glandular-hairy outside; 
upper Up as long as or slightly shorter than the orbicular-reniform slightly cleft lower Up; 
nutlets 1 .5 mm across, angular-ovoid, light brown, finely tuberculate. June— September. 

Flooded and swampy meadows, bogs and marshy forests, shrubs, damp banks of rivers, 
brooks and lakes, ditches. — European part: Kar.-Lap., Dv.-Pech., V.-Kama, Bait., Lad.- 
Ilm., U.V., U. Dnp., M. Dnp., U. Dns., Bes., BL, V.-Don, Transv., L.V.; Caucasus: Cisc, 
E. and S. Transc, W. Transc. (see Note); West Siberia: ah regions; East Siberia: aU 
regions; Centr. Asia: Ar.-Casp., Balkh., Amu D., Dzu.-Tarb., T. Sh., Pam.-Al., Mtn. Turkm. 
Gen. distr.: Scand., Atl. and Centr. Eur., Med. (sporadically), Bal.-As. Min. (northern part 
of Balkans), Mong., Ch., Jap. Described from Europe. Type in London. 

Note. A highly variable plant, prone to produce poorly defined local races. Specimens 
from Western Transcaucasia are very uniform; they are distinguished by high growth, 
strongly branching stems and leaves glabrous on both sides, with hairs confined to the 
veins (S. phasiana Juz. ined.). A Far Eastern (Ussuri) race is described below. A North 
American race was described as S. epilobiifolia A. Ham. (Monogr. ScuteU., 1832, 32). 

Economic importance. In former times the plant was used medicinally, mainly in the 

92 treatment of malaria ("herba Tertianariae"). It is popularly used for its hemostatic action. 
A dyeing agent. 

2. S. krasevii Kom. et I. Schischk. in sched. (1927) ex Juz. in Bot. mat. gerb. Bot. inst. 
ANSSSR, XIV (1951) 357. - S. galericulata /J. pubescens Rgl. Tent. Fl. Ussur. (1861) 
1 18, non aUor. — S. scordifolia var. velutina Kom. in sched. (1924) sec. Juz. 1. c. 

Perennial, with creeping rhizome; stems 32-75 cm long, erect, simple or mostly branch- 
ing from base or above, the branches long, subappressed or obUquely ascending, covered 
on the ribs with short reclinate hairs, the internodes as long as the leaves or shorter; leaves 



62 




PLATE V. 1 - Scutellaria ikonnikovii Juz., general aspect; 2 - S. hastifolia L.,end of stem; 
3 - S. scordiifolia Fisch., leaf, flower, calyx; 4 - S. taquetii Levi, et Vaniot, general aspect, leaf; 
5 - S. baicalensis Georgi, summit of stem, calyx, upper and lower part of fruiting calyx, nutlet. 



63 



rather thin, 1.2-4.5 cm long, 0.5-2 cm wide, ovate or triangular-ovate, cordate or truncate 
at base, rather abruptly attenuate at apex, obtuse to subacute, rather deeply and unevenly 
crenate-dentate below, with 2-7 obtuse or acute teeth on each margin, entire toward apex, 
pale or bright green with short appressed hairs above, the undersurface mostly densely cov- 
ered with short crisp hairs, often nearly velutinous, gray-green, the veins with profuse short 
retrorse hairs; petioles short, covered with reclinate hairs; flowers small; calyx ca. 3 mm 
long, with profuse short patent hairs; corolla violet-blue, densely hairy outside, mostly ca. 
1.5 cm long; upper lip much shorter than the lower; nutlets ca. 1.5 mm across, globose- 
ovoid, yellowish, finely and closely tuberculate. July. 

Thickets of purple osier, cane, common reed and smallreed along banks of rivers and 
lakes, oxbow -lakes and backwaters. - Far East: Uss. (?), Uda. Endemic? Described from 
Gaivoron village in Spassk-Dalni region. Type in Leningrad. 

Note. The morphological differences between this race and the typical S. galericula- 
ta L. are negligible and difficult to discern. Nevertheless, we are presenting it here as an 
independent unit, chiefly because of some disjunction in its distribution area. 



Series 2. Regelianae Juz. in Bot. mat. gerb. Bot. inst. ANSSSR, XIV (1951) 359. - 
Like Eu-galericulatae Juz., but the flowers rather large, to 2.5 cm long. 
95 N ote. Even though Hulten questioned the specific status of one of the forms concern- 
ed (S. yezoensis Kudo) and argued (Fl. of Kamtch. IV, 1930, 88) th^t its distinguishing 
characters lie within the variation range of S. gale ricu lata L., there are strong indications 
for the distinctiveness of large-flowered East Asian forms of the S. galericulata— S. latis- 
sim a type. Most convincing is the fact that the distribution area of two of these forms 
(S. ikonnikovii Juz. and S. regeliana Nakai) partly overlaps the area of S. galericulata 
s, 1. (inch S. krasevii). See, however. Note to S. yezoensis. 

3 . S. yezoensis Kudo in Journ. Coll. Sc. Univers. Tokyo, XLIII, art. 8 (1921). - S. ga- 
lericulata Fr. Schm. Fl. Sachal. (1868) 165, non L. - Ic: Kudo, 1. c. tab. I, fig. 9. 

Perennial; stem 35-70 cm long, erect, simple or branching, often with elongate inter- 
nodes, sparsely crisp-hairy on the ribs, otherwise glabrous; leaves 2.5-5.5 cm long, 1-2 cm 
wide, thin, oblong-lanceolate, ovate-lanceolate or elongate-ovate, truncate or shallowly cor- 
date at base, acute at apex, distinctly crenate-dentate with 4-14 short teeth on each margin, 
with short crisp hairs or subglabrous or scabridulous above, paler, with rather profuse short 
crisp hairs beneath, densely and finely appressed-hairy on midrib; petiole to 4 mm long (in 
lower leaves), with hairs as on midrib; bracts like the leaves but gradually smaller toward 
apex, oblong-lanceolate, cordate at base, acuminate, obscurely and finely crenate-dentate 
or nearly entire; flowers axillary, remote, on pedicels approximately half the length of 
calyx; calyx 4-5 mm long, densely covered with short white slightly crisp hairs; corolla 
2-2.4 cm long, violet-blue, crisp-hairy outside; nutlets 1.5-1. 8 mm long, ellipsoid, densely 
covered with small tubercles. July— August. 

River valleys, shores of lakes. - Far East: Kamch. (?), Sakh. (Kuriles). Gen. distr.: 
Japan. Described from Japan (Kyushu). Type in Tokyo. 

Note. This species combines the characters of the series Eu-galericulatae (leaf shape 
and dentation) and Regelianae (flower size) and could be included in the former (but then, 
the characterization of the series would have to be modified). 

64 



4. S. regeliana Nakai in Bot. Mag. Tokyo, XXXV (1921) 197. - S. galericulata 7. 
angustifolia Rgl. Tent. Fl. Ussur. (1861) 118. - S. angustifolia Kom. in Tr. Bot. sada, 
XXV, II (1907) 345, p. p. non Pursh nee Adams. - Ic: Komarov, op. cit. Table IV, Fig- 
ure 1; Kom. and Alls. Opred. rast. Dal'nevost. kraya, II, Plate 271, 6-10. 

Perennial, with slender, creeping, rarely vertical rootstock; stems solitary, 20-60 cm 
long, erect, mostly simple, sometimes with long appressed branches from base, mostly few- 
brancled, with elongate internodes (especially in lower part), densely short-hairy on the 
ribs or all over, frequently becoming lilac (like lower side of leaves); leaves gray-green, 
oblong or lanceolate, 1-4 cm long, 0.2-0.8 cm wide, attenuate above but obtusely rounded 
at apex, rarely acute, entire or, especially in lower part, with few shallow and flat, often 
indistinct crenations, frequently slightly revolute-margined, short-petioled or subsessile, 
densely covered on both sides with short hairs, these short antrorse setiform above, softer 
curved or slightly crisp beneath, antrorsely appressed on the veins; flowers few, large, on 
short erect pedicels in the axils of middle and upper leaves; calyx always densely covered 
with short curved white hairs; corolla 1 .5-2.5 (usually 2) cm long, blue-violet, with spo- 
radic pubescence and short stipitate glands, the tube markedly curved, the throat ca. 1 cm 
wide; nutlets with slender papilliform tubercles over entire surface. July— September. 

Banks of rivers and oxbow-lakes, swampy meadows, sometimes also sphagnum bogs, 
more rarely birch groves, wastelands. — Far East: Uss. Gen. distr.: China (Manchuria), 
Korea. Described from the Ussuri River valley, Usachi Mountain and estuary of Iman 
River. Type and paratype in Leningrad. 

5. S. ikonnikovii Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 358. - S. an- 
gustifolia Kom. in Tr. Bot. sada, XXV, II (1907) 345, p. p. non Pursh nee Adams. — 

S. regeliana Ikonnikov in sched. ex Juz. I. c. non Nakai. — S. galericulata Ldb. Fl. 
Ross. Ill, 398, p.p. non L. — S. galericulata var. subintegerrima F. Schm. in sched. 
ex Juz. 1. c. — Ic: Sugawara, 111. fl. of Saghal. IV, tab. 725. 

Perennial, similar to S. regeliana Nakai but usually smaller; stem 10-35 cm long, the 
ribs covered with very short appressed hairs; leaves bright green, glabrous or with scatter- 
ed hairs above, with very short appressed hairs beneath only on the veins and margins, 
elsewhere diffusely and finely grainy -glandular; flowers disproportionately large for the 
size of the plant, to 2.5 cm long. July— August. (Plate V, Figure 1.) 

River banks, meadows and swamps. — East Siberia: Ang.-Say. (shores of Baikal, Kultuk 
settlement), Dau., Lena-Kol.; Far East: Uda, Ze.-Bu., Uss. (western part), Sakh. Gen. 
distr.: Mong. Described from Transbaikalia (Chitinskaya highway, Karymskoe). Type in 
Leningrad. 



Series 3. Hastifoliae Juz. - Similar to Eu-galericulatae but leaves broader at base, with 
lowermost teeth exceptionally well developed and thus more or less hastate, entire above. 

6. S. hastifolia L. Sp. pi. (1753) 599; Ldb. Fl. Ross. Ill, 399; Shmal'g. Fl. II, 328; 
Kryl. Fl. Zap. Sib. IX, 2298. - Cassida hastifolia Scop. Fl. Carn. I, ed. II (1772) 430. - 
Ic: Rchb. Ic. Fl. Germ. XVIII, 1256; Syreishch. III. Fl. Mosk. gub. Ill, 111. - Exs.: Fl. 
pol. exs. No.767(nom. S. galericulata). 



65 



Perennial, with a very slender creeping rhizome and short stolons; stems 15- 40 cm 
long, ascending, simple, rarely branching, glabrous or with short hairs mainly on the ribs; 
leaves 12-18 mm long, 6-12 mm wide, oblong-ovate or oblong-lanceolate, orbicular or 
truncate at base, the middle and often the lower hastate at base, often with 1-2 (3-4) 
obtuse teeth on the lobes, otherwise entire, obtuse at apex, glabrous or diffusely pillulose 
above, only on margins and sometimes on the veins with very short hairs, otherwise gla- 
brous, frequently becoming lilac beneath (like stems); petioles 2-5 (10) mm long; inflo- 
rescence a short loose secund spikelike raceme; flowers paired in the axils of upper cauline 
leaves which they exceed in length; pedicels short, covered (like the subtending leaves) 
with short glandular hairs; calyx 2.5-4 mm long, slightly longer than the pedicel, often 
densely glandular-hairy, usually violet; corolla (13) 17-22 mm long, blue-violet, glandular- 
hairy outside; lateral lobes of upper lip rounded, slightly shorter than the middle lobe; 
lower lip rounded-reniform, obtuse or scarcely cleft; nutlets 1-1.5 mm across, papillose- 
tubercled. June— August. (Plate V, Figure 2.) 

River banks, swampy meadows, coppices, oak forests. -European part: Balt.,Lad.-Ibn., 
U.V., U. Dnp., M. Dnp., U. Dns., Bes., Bl, V.-Don, L. Don; Caucasus: Cisc; West Siberia: 
Ob. Gen.distr.: Scand., Atl. and Centr. Eur., Bal.-As. Min. Described from Sweden. 
Type in London. 

7. S. dubia Tal. et Schir. in Sb. tr. Khar'kovsk. veterin. inst. VII, 6 (1906) 60 (rossice); 
Shiryaev in Tr. Bot. sada Nyur'evsk. univ. XII (191 1) 22 (diagn. lat.). - S. tatarica Fisch. 
in sched. — S. hastifolia f. wolgensis Schennikov in sched. - Ic: Taliev and Shiryaev, 
op. cit. 60; Stankov- Taliev, Opred. 860. 
98 Perennial; stems 20-55 cm long, erect or ascending, straight or slightly flexuous, simple 
or often branching, glabrous or very finely hairy, very leafy; leaves on distinct but fre- 
quently short, glabrous or finely hairy petioles, the lower to 3-4 cm long, 2 cm wide, the 
upper leaves gradually smaller; lower leaves of lateral branches 2-2.5 cm long, 0.8-1 .3 cm 
wide, others smaller, aU triangular-ovate, truncate or cordate at base, obtuse at apex, un- 
evenly crenate-dentate in lower part, with 2-10 teeth on each margin, entire in upper part, 
glabrous at both sides or sparsely hairy above, minutely puberulent on the margin or on 
the veins beneath; inflorescence secund; flowers paired in the axils of upper leaves; pedi- 
cels 1-2.5 mm long covered with short hairs interspersed with stipitate glands; calyx 2- 
3 mm long, to 7 mm long in fruit, declined or nearly pendulous, densely hairy and glandu- 
lar; corolla 1 .6-2 cm long, lilac-blue, glandular-hairy, with slightly curved tube; anthers 
. hairy; nutlets ellipsoid, densely papillose-tubercled, brown. June— August. 

* ""flood plains, forests and coppices, margins of swamps. — European part: V.-Don, 
Transv.,L. Don,L.V.; West Siberia: Ob, U.Tob., Irt.; Centr. Asia: Balkh. Endemic. 
Described from vicinity of Ust'-Medveditskaya station in Don region. Type in Khar'kov. 

Note. This species is apparently connected with S. hastifolia L. through intermediate 
(or transgressive?) forms. In their description Taliev and Shiryaev grouped it primarily 
with S. scordiifolia, which we find an unsuitable choice. 



Series 4. Minores Juz. — Like Eu-galericulatae or, more so, HastifoUae, but plants 
usually smaller; flowers very small, ca. 5 mm long, on finally drooping pedicels. 



66 



8. S. dependens Maxim. Prim. fl. Amur. (1859) 219; Komarov in Tr. Bot. sada, XXV, 
II, 341. - S. oldhami Miq. in Ann. Mus. Bot. Lugd.-Bat. Ill (1867) 197. - S. minor 
Ldb. Fl. Ross. Ill, 399; Turcz. Fl. baic.-dahur. 11,419, non L. — S. breviflora Turcz. in 
sched. - Ic: Sugawara, 111. Fl. of Saghal, IV, tab. 727. 

Perennial, with slender subfiliform rhizome and long capilliform stolons; stems 5-50 cm 
long, mostly erect, straight or flexuous, weak, sometimes decumbent, sulcate, glabrous, 
simple or branching, the branches appressed or ascending, sometimes nearly horizontally 
spreading, often very long; leaves 0.4-3 cm long, 0.2-1 .7 cm wide, ovate, oblong-ovate to 
narrowly triangular, with cordate, orbicular, truncate or nearly sagittate base, obtuse at 
99 apex, entire or very often crenate in lower part, with 1-3 uneven obtuse teeth on each 
margin, sometimes obscurely crenate, glabrous above or with very fine short hairs only 
along margins and on the veins beneath, the short petioles mostly with scattered short 
appressed hairs; flowers solitary in the axils of nearly all leaves (except the lowermost); 
pedicels scabrous, longer than petioles, erect at first, later drooping; calyx ca. 1 .5 mm 
long, pale green, prominently nerved, the margin and nerves slightly hispid; corolla small, 
ca. 5 mm long, twice the length of calyx, white or pale violet with blue spots, inflated at 
throat, pubescent outside; nutlets depressed-globose, very small, verrucose. June— August. 
(Plate VI, Figure 1.) 

Shady forest valleys, water meadows, swamps, osier-beds along river banks, forest 
streams, rice fields. — East Siberia: Ang. -Say. (Irkutsk), Dau.; Far East: Ze.-Bu., Uss., 
Sakh. Gen. distr.: Jap. and Ch. Described from lower Amur, slopes of Khekhtsir Moun- 
tain (near estuary of Ussuri River). Type and paratype in Leningrad. 

Note. The true West European S. minor L., reported for Lithuania by older authors 
(Gilibert, Jundzill - quoted by Ledebourg (Fl. Ross. Ill, 400)), does not occur in the USSR. 



Series 5. Scordiifoliae Juz. — Like Eu-galericulatae or Regelianae but leaves foveo- 
late-punctate beneath. 

9. S. scordiifolia Fisch. ex Schrank in Denkschr. Bot. Ges. Regensb. — II (1822) 55; 
Ldb. Fl. Ross. Ill, 398; Kryl. Fl. Zap. Sib. IX, 2298. - S. galericulata 5. scordifolia 
Rgl. Tent. Fl. Ussur.( 1861) 118. —S. adamsii auct. plur. in synon. non Spreng. (nee 
Hamilt.). — S. reptans Pall, in herb. sec. Ldb. 1. c. Ill, 398. — S. depressa Fisch. mss. ex 
Ldb. 1. c. — S. montana Rudolph in herb. — Ic: Komarov in Tr. Bot. sada, XXV, II, 
Plate IV, Figure II (non typica). 

Perennial, with slender vertical or creeping rhizome; stems 10-30 cm long, erect, nor- 
mally with decHned branches from base, rarely simple, covered mainly on the ribs with 
short curved hairs; leaves 1-3.5 cm long, 3-8 mm wide, oblong or lanceolate, the lower 
wider, sometimes oblong-ovate, rounded or broadly cuneate at base, obtuse, entire or 
sparsely denticulate, usually glabrous above, rarely with scattered short hairs, foveolate- 
punctate beneath, with appressed short hairs mostly confined to margin and veins, short- 
100 petioled; inflorescence a short, lax, secund raceme; flowers solitary in the axils of, and 
usually exceeding, the leaves; calyx 3.5-4 mm long, short-appressed-hairy and glandular, 
commonly colored; corolla 18-22 mm long, violet-blue, glandular-hairy outside; lateral 
lobes of upper Up scarcely as long as the middle lobe; lower lip equaling or slightly 



67 



exceeding the upper, slightly emarginate; nutlets 1 .5-2 mm long, largely papillose-tubercled. 
June -September. (Plate V, Figure 3.) 

Steppe meadows, wastelands, stony slopes. — West Siberia: Ob., Irt., Alt.; East Siberia: 
universal; Far East: Ze.-Bu., Uss. Gen. distr.: Mong., Ch., Jap. Described from Siberia. 
Type unknown. 

10. S. taquetii Levi, et Vaniot in Fedde, Repert. VIII (1910) 402. - S. scordifolia 
var. hirta Fr. Schm. Fl. Sachal. (1868) 165. — S. scordifolia var. pubescens Miq. in 
Ann. Mus. Bot. Lugd.-Bat. Ill (1867) 197; Kom. (cit. Regl.) in Tr. Bot. sada, XXV, II, 
344 (non S. galericulata |3. pubescens Rgl.). — S. scordifolia var. sachalinensis 
Matsum.et Kudo in Bot. Mag. Tokyo, XXVII (1912) 296. - S. scordifolia var. obtusi- 
folia Levi, in Fedde, I.e. 280. — S. pubescens Loes. in Beih. Bot. Centralbl. XXXVII, 
II (1920) 174. - S. schmidtii Kudo in Journ. Coll. Sc. Univers. Tokyo, XLIII, 8 (1921) 
13, saltem pro max. parte. - S. strigillosa Kom. in Kom. and Alls. Opred. rast. Dal'ne- 
vost. kraya, II (1932) 899, vix Hemsl. - Ic: Sugawara, 111. Fl. of Saghal. IV, tab. 724 
(nom. S. strigillosa Hemsl.). 

Perennial; rhizome slender, subvertical or oblique, rarely creeping, yellowish, jointed, 
rooting at nodes; stems 7-70 cm long, erect or flexuous, usually strongly branching from 
base, rarely simple, mostly covered all over, rarely only ^n the ribs, with shortish patent 
crisp hairs, pale yellowish-green, often lilac-tinged, the branches flexuous, branching in 
turn; leaves 1-5 cm long, 0.3-2.2 cm wide, elliptic, rarely ovate or oblong, rounded or 
slightly subcordate at base, obtuse or rounded at apex, shallowly crenate-dentate or cre- 
nulate, with few (usually 3-7, rarely to 10) teeth on each margin, or very often all or some 
(especially the upper) entire, densely covered on both sides with short hairs, these straight 
appressed above, straight or somewhat crisp, appressed or mostly spreading beneath, the 
petiole short or very short; flowers in axils of upper leaves, forming a short lax secund 
raceme; pedicels erect or recurved; calyx 3-5 mm long, with rather long appressed or 
spreading hairs and, especially at base, stipitate-glandular, usually colored; corolla 2- 
2.5 cm long, to 1.5 cm wide at throat, covered outside with short glandular hairs; lateral 
101 lobes of upper lip shorter; lower hp usually distinctly longer than the upper; nutlets ca. 
1 .8 (to 2) mm long, elliptic, densely papillose-tubercled, dark brown or blackish. June. 
(Plate V, Figure 4.) 

Seacoasts, shores of lakes and rivers (often near estuaries), mostly on sands, sandy 
embankments and hills, rarely on rocks, sometimes in fields. — Far East: Sakh. (Kuriles), 
Uda., Uss. Gen. distr.: Jap., Korea. Described from Korea, from Quelpart Island. Type 
in Paris, isotype in Leningrad. 

Note. This species is sometimes reported as S. strigillosa Hemsl. which was described 
from China (Sinkiang and Shantung provinces). We have never seen the material; Hem- 
sley's brief description shows little resemblance to our plant (there is notably a significant 
difference in vesture). Nevertheless, the name S. strigillosa has priority. 

11. S.tuminensisNakaiinBot.Mag. Tokyo, XXXV (1921) 198. - S. galericulata e. 
puberula Rgl. Tent. Fl. Ussur. (1861) 119, saltem p.p. 

Perennial; rhizome creeping, stout, white; stems 20-35 cm long, mostly simple or few- 
branched, becoming purple (especially in lower part), covered (usually sparingly) with fine 



68 



hairs and with dense bundles of longer hairs at nodes; leaves 1-5 cm long, 0.6-2 cm wide, 
ovate or oblong-ovate, with rounded, cordate or nearly sagittate base, acute or obtuse at 
apex, often serrate-dentate, with 7-12 rather high acute teeth, rather densely covered on 
both sides (especially the upper) with short white hairs, bright green above, pale beneath, 
with very prominent veins, obscurely foveolate-punctate, short-petioled or subsessile; 
flowers clustered at summit of stem, rather large, 1 .5-2.5 cm long, soUtary in the axils of 
sharply serrate-dentate mostly subacute or acute bracts, on rather patent-hairy pedicels 
2-4 mm long; calyx 4-5 mm long, patent-hairy, with short scutellum, usually colored; 
corolla violet-blue, the tube hairy and glandular outside, the broad lower hp longer than 
the upper; anthers hairy. June— August. 

Meadows, oak forests, coppices in valleys of rivers and streams, grassy and scrub-covered 
volcanic slopes, wastelands. - Far East: Uss. Gen.distr.: Ch. (Manchuria). Described 
from Kanto region, along Tumingan River. Type in Tokyo. 

Note. A little known, critical species of which we have not seen the type. It may be 
one of the intermediate forms between S. scordiifolia and S. taquetii which apparent- 
ly occur frequently where the distribution areas of these two species converge. 

^^^ 12. S. polyphylla Juz. in Bot. mat. gerb. Bot. inst. AN SSSR. - XIV (1951) 359. 

Perennial, 25-45 cm high; stem erect, branching from base, glabrous or short-hairy on 
the ribs below; branches very long, erect; leaves very many (25-30 pairs on stem and 
branches), approximately the length of internodes or often shorter, 1-3 cm long, 0.3- 
1 .6 cm wide, ovate or ovate-lanceolate, gradually attenuate from broad, truncate or sub- 
cordate base, obtuse or acute at apex, with 3-10 well developed, strongly inequilateral 
obtuse teeth on each margin, pale green, completely glabrous on both sides, or the upper 
surface with scattered short hairs, the lower with isolated minute hairs sometimes confined 
to the veins, obscurely foveolate-punctate; petioles well developed, 3-12 mm long; flow- 
ers few, small, in the axils of remote upper leaves, on pedicels 3-5 mm long; calyx 2-3 mm 
long, with short patent hairs; coroUa 1 .4-1 .7 cm long, sparingly short -hairy and glandular; 
nutlets ca. 1 mm wide, globose-ovoid, black, densely and finely tuberculate. July— August. 

Mixed and coniferous forests in valleys, riverbanks. — Far East: Ze.-Bu., Uss. Endemic? 
Described from Khabarov Territory. Khekhtsir. Type (Desulavi, No. 1481) in Leningrad. 

Note. We have only tentatively placed this little known species in the series Scordii- 
fohae Juz. It may perhaps prove to be related to S. komarovii Levi, et Vain, which was 
described from Korea and which we have never seen, and may eventually join it to make 
up a specific series. 



Series 6. Moniliorrhizae Juz. — Flowers large, 2-3 cm long; rhizome moniliform, 
consisting of white jointed tubers; leaves not foveolate-punctate beneath. 

13. S. moniiiorrhiza Kom. in Tr. Bot. sada, XXV, ed. II (1907) 346. - Ic: Komarov, 
op. cit. Plate IV, Figure III. 

Perennial; rhizome vertical or creeping, moniHform, consisting of white jointed tubers, 
rooting at the joints, with fihform stolons arising at the crown; stems 12-35 cm long, 
single, simple or with few appressed branches at base, rarely divaricately branched, more 



69 



or less white-hairy at nodes, glabrous elsewhere; leaves ovate or oblong-ovate, cordate 
or rounded at base, rounded at apex or sometimes acute, crenate-dentate, with 3-7 
103 large obtuse teeth on each margin, glabrous or with long patent hairs above, usually 
glabrous beneath or with few long hairs on veins and densely and finely granular- 
glandular; flowers few, large, in the axils of small but always crenate-dentate upper 
leaves, short-pediceled; calyx ca. 3 mm long, in fruit ca. 5 mm, sparingly pubescent 
(usually only on nerves and margins) or subglabrous; corolla 2-3 cm long, violet-blue, 
with whitish tube, short-hairy and glandular outside; nutlets 1-1.5 mm long, ellipsoid, 
densely papillose-tubercled, brownish. June —September. 

Mountain forests, stony and rocky placers, in thick grass around springs. — Far East: 
Uss. Gen. distr.: Korea. Described from N. Korea. Type and paratype in Leningrad. 



Section 2. Stachymacris A. Hamilt. Monogr. Scutell. in Ser. Bull. Bot. V(1832) 1.- 
Flowers in dorsoventral verticillasters, forming terminal racemose one-sided spikes; bracts 
herbaceous, resembling cauline leaves but smaller, or markedly distinguished by their shape, 
smaller size and entire margin, but leaflike in being distinctly petioled. Perennial, often 
comparatively large herbs of mesophytic aspect, with sessile or long-petioled, entire or 
dentate leaves. 

This section, like the preceding, has a holarctic type of distribution. 

Note. A very heterogeneous section, usually combined with the preceding under the 
name Vulgares Benth. (in Benth. et Hook, fil. Gen. II, 1876, 1202). There is some ground 
for this unification inasmuch as there is no clear-cut dividing line between Galericularia 
and Stachymacris; thus the subsection Angustifohae Benth., for example, could well be 
included in the section Galericularia if just a few slight changes were introduced in the 
characterization of the latter (c. f . Benth. in DC. Prodr. XII, 1 848, 423). However, for the 
purpose of our flora, we prefer to retain the subdivision of the subgenus Eu-scutellaria 
into three sections and thus avoid further increase of heterogeneity within the sections. 



Subsection 1. Angustifoliae Benth. in DC. Prodr. XII (1848) 423. - Bracts resem- 
bhng cauline leaves but smaller, always sessile; flowers large. Erect herbs, with lanceolate 
or linear, entire, somewhat coriaceous, sessile or short-petioled leaves. 

14. S. baicalensis Georgi, Bemerk. einer Reise im Russ. R. I (1775) 223; id. Beschr. 
Russ. R. Ill, 5, 1097; Ldb. Fl. Ross. Ill, 397. - S. macrantha Fisch. ex Rchb. Ic. Bot. 
104 PI. crit. V (1827) 52. - S. speciosa Fisch. ex Turcz. Cat. baic.-dahur. II (1838) 416. - 
S. grandiflora Adams ex Bge. Enum. pi. Chin. bor. (1831) 52. - S. adamsii A. Hamilt. 
Monogr. Scutell. (1832) 34, non Spreng. - S. lanceolaria Miq. ex Franchet et Sav. Enum. 
pi. Japon. I (1875) 377 (in syn.). — S. tinctoria Pall, in sched. - S. ammaniana Ldb. in 
sched. — Ic: Amman, Stirp. rar. Ruthen. ic. et descr. tab. VI; Rchb. 1. c. tab. 488, fig. 
681 ; A. Hamilt. 1. c. tab. 2, fig. 3. - Exs.: ex herb. H. Bot. Jurjev. sine No., sub nom. 
S. macrantha Fisch. 

Perennial; root vertical, long, thick, fleshy, branching; stems few, 15-35 (50) cm long, 
slightly ascending or erect, robust, straight or flexuous, simple or branching from base. 



70 



glabrous or short-hairy (especially on ribs), green or often suffused with purple (especially 
in lower part); leaves 1.5-4 cm long, 1 .5-13 mm wide, ovate-lanceolate to linear-lanceolate, 
orbicular or usually tapering at base, obtuse or acute, entire, glabrous or with short patent 
hairs above, ciliate on the margins, glandular-foveolate beneath, glabrous or short-hairy 
along midrib, firm, often subcoriaceous, often with slightly revolute margins; petioles 
obsolete or very short, ciliate; inflorescence a simple, one-sided, rather dense raceme, 
sometimes branching (especially in the Far East); flowers large, solitary in the axils of 
upper leaves, these slightly reduced in size, much shorter than the flowers; calyx ca. 3 mm 
long in flower, mostly rather densely short-hairy, long-hairy at apex, usually violet; corolla 
2-2.5 cm long, blue, densely covered outside with long glandular hairs, the tube strongly 
dilated above, the upper Up slightly longer than the lower; nutlets small, flattened-subglo- 
bose, black, aculeolate. June— July. (Plate V, Figure 5.) 

Stony and clayey dry mountain slopes, steppes, wastelands, osier beds along riverbanks. 
- East Siberia: Ang.-Say. (Lake Baikal), Dau.; Far East: Ze.-Bu.,Uss. Gen.distr.: Mong., 
Ch., Jap. Described from Transbaikalia. Type unknown. 

Economic importance. Used by the Chinese as pectoral emoUient and anthelmintic 
remedy. In Tibet it is appHed in the treatment of pneumonia, myocarditis, palpitation, 
acute rheumatism and as an antipyretic. It has valuable qualities as a hypotensive agent 
and should be introduced into medical practice in the treatment of hypertonia and func- 
tional disorders of the nervous system (see articles by E.M. Dumenova, A.S. Saratikov, 
A.M. Voronova and E.A.Toloknov in collection: Novye lekarstvennye rasteniya Sibiri i 
ikh lechebnye preparaty, Vol. II, Tomsk, 1946). An ornamental. Apparently a dye stuff. 



105 Subsection 2. Peregrinae Boiss. Fl. or. IV (1879) 681. — Bracts differing markedly 
from cauline leaves, small, entire, short-petioled; flowers usually of medium size. Com- 
paratively large herbs, with erect or rarely ascending stems and broad, frequently ovate, 
petiolate, crenate-dentate leaves. 



Series 1. Altissimae Juz. — Large plants (to 1 m high), with large, strongly crenate- 
dentate leaves; bracts generally not longer than calyx, ovate, short-petioled; flowers 
partly or completely anthocyanin-colored. 

15. S. altissima L. Sp. pi. (1753) 600; Ldb. Fl. Ross. Ill, 396; Shmal'g. Fl. II, 327. - 
S. commutata Guss. Fl. Sic. Prodr. II (1828) 136. - S. peregrina Waldst. et Kit. Ic. et 
descr. pi. rar. hung. II (1805) 132, non L. — S. peregrina var. altissima Fiori, Nuova Fl. 
anal. It. Ill (1900) 16. - Ic: Waldst. et Kit. 1. c. tab. 125; Reching. fll. in Bot. Arch. 43, 
tab. 1, fig. 11. - Exs.: Rehm. et Woloszcz. Fl. pol. No. 228; Callier, It. taur. Ill, No. 705. 

Perennial; rhizome oblique, nodose; stems 25-100 cmlong, erect, simple or with erect 
opposite branches, 4-angled, covered with small curved hairs, shortly stipitate-glandular 
above, especially in inflorescence; leaves petiolate, 4-8 cm long, 2.5-7 cm wide, ovate, with 
truncate or cordate base, obtuse or acuminate, with (3) 6-12 large obtuse crenate teeth on 
each margin, glabrous or sparsely short-hairy above, more densely and finely curved-hairy 
beneath (especially on the veins); bracts short-petioled, broadly ovate, acute, entire, equaling 



71 



or slightly exceeding the calyx; inflorescence spikelike, to 30 cm long; calyx curved, cov- 
ered with long soft and short stipitate-glandular hairs, often more or less violet; corolla- 
tube geniculately curved in lower part, glandular-hairy, bluish-white, the back of the tube 
and upper lip violet-blue; upper lip longer than the lower; nutlets flattened-ovoid, ca. 
2.5 mm long, finely verrucose, with sohtary stellate 3-5-rayed hairs on the warts. May — 
July. (Plate VI, Figure 2.) 

Open deciduous forests (especially oak), scrub, meadow slopes. — European part: V.- 
Kama (S. Ural), V.-Don, M. Dnp., Bl., Bes., Crim.; Caucasus: Cisc, E. and W. Transc. 
Gen. distr.: Centr. Eur. (usually escaped), Med., Bal.-As. Min. Described from Georgia 
("Iberia"). Type in Paris (Tournefort Herbarium). 

1^^ 16. S. toumefortii Benth. Bot. Reg. XVIII (1832) ad calcem. No. 1493; id. apud DC. 
Prodr. XII (1848) 419; Ldb. Fl. Ross. Ill, 396. - S. altissima var. ? coroUis 4-plo majori- 
bus C. A. M. Verzeichn. (1831) 89. - S. altissima var. grandiflora C. A. M. in sched.? - 
Ic: Reching. fil. in Bot. Arch. 43, tab. 1, fig. 19. 

Perennial; stems 20-85 cm long, erect, straight or somewhat flexuous, simple or shghtly 
branching, 4-angled, covered with fine curved hairs; leaves 2.5-8.5 cm long, 1.4-5.5 cm 
wide, ovate-triangular, truncate or subcordate at base, obtuse or acute, coarsely and obtuse- 
ly crenate with 5-10 teeth on each margin, sparsely covered on both sides with short- 
appressed or close-appressed hairs, on petioles 0.5-2.5 cm long; inflorescence 7-25 cm 
long, racemose, 1-sided, lax, the axis covered with short curved hairs; bracts short-petioled, 
ovate-rhombic, acute, entire, shorter than calyx; calyx 4-5 mm long, with short patent 
hairs and stipitate glands, corolla 6 times the length of calyx, 1 .7-2.2 cm long, purple, 
sparsely covered with short hairs and short stipitate glands; nutlets ca. 1 mm long, dark 
brown, scabrous with short papilliform hairs or warts. May — September. (Plate VI, 
Figure 3.) 

Forests, shrubs on mountain slopes, ravines. — Caucasus: Tal. Gen. distr.: Iran 
(Hircan province). Described from Iran. Type in London (Lambert Herbarium). 



Series 2. Indie ae Juz. - Like Altissimae but smaller (usually to 40 cm high) and more 
slender; bracts lanceolate. 

17. S. ussuriensis (Rgl.) Kudo in Kudo et Yoshimi Rep. Veg.Tomakomai Forest (1916) 
53; Nakai in Bot. Mag. Tokyo, XXXV(1921) 198; Kom. and Alls. Opred. rast.Dal'nevost. 
kraya, II, 896. - S. japonica j3. ussuriensis Rgl. Tent. Fl. Ussur. (1861) 1 18, p. p. (s. 
str.). — S. japonica Maxim. Prim. fl. Amur. (1859) 218 et auct. plus. Ross. p. p., non 
Morr. et Decne. - S. indica Kom. in Tr. Bot. sada, XXV, II (1907) 339 (nee alibi) p. max. 
parte, non L. -S. indica var. japonica auct. Ross. p. p. non Miq. — Ic: Kom. and Alls., 
op. cit . Plate 271, Figures 1 -5 . 

Perennial; rhizome very slender, long, white, with slender underground shoots; stems 
5-40 cm long, sohtary, erect, straight or slightly flexuous, with few internodes, simple or 
rarely branching, with erect or arched or flexuous branches, 4-angled, deeply sulcate, gla- 
brous or shghtly patent-hairy (especially under the nodes), often dark purple; leaves dark 
107 green, 0.8-6.5 cm long, 0.6-4.5 cm wide, orbicular, broadly ovate or ovate, with rounded, 



72 



cordate or truncate base, obtuse, coarsely and unevenly crenate with 5-12 irregular, round- 
ed or obtusish teeth, glabrous on both sides or with sparse, frequently long, multicellular 
(articulate) white hairs above and on the veins beneath; petioles slightly shorter to rarely 
longer than blade; inflorescence racemose, terminal, erect, rather lax, mostly one-sided; 
floral leaves at base of inflorescence resembling upper cauline leaves but much smaller; 
bracts small, usually lanceolate, attenuate at both ends, petiolate, acute, entire; pedicels 
as long as bracts, finely pubescent; calyx green or dark violet, with few hairs; corolla 10- 
20 mm long, partly white, partly blue (upper lip), with arched tube; nutlets ca. 1 mm long, 
angled-ovoid, brownish, muricate-tubercled. June— September. 

Coniferous and mixed forests, riverbanks, in the shade of trees or among shrubs, on 
Uttered soil, on rocks or decayed tree trunks. — Far East: Uss. Gen. distr.: Korea. 
Described from Ussuri River valley (estuary of Ussuri River near Turme). Type in 
Leningrad. 

18. S. pacifica Juz.inBot. mat.gerb.Bot.inst.ANSSSR,XIV(1951)362.-S.indica 
Kom. in Kom. and Alis. Opred. rast. Dal'nevost. kraya, II (1932) 895 et auct. plur. non L. 
fil. — S. japonica Maxim. Prim. fl. Amur (1859) 218 et auct. plur. Fl. Ross. p.p. non 
Morr. et Decne. — S. indica var. japonica auct. Ross. p.p. non Miq. — S. japonica j3. 
ussuriensis Rgl. Tent. fl. Ussur. (1861) 1 18, p. p. - ? S. ussuriensis var. transitra (sic) 
Nakai in Bot. Mag. Tokyo, XXXV (1921) 199, non S. transitica (sphalm. transitra) Mak. 

Perennial; stems 8-35 cm long, mostly erect, with patent curved hairs at least on the 
ribs; leaves 0.6-5.5 cm long, 0.7-4.3 cm wide, rather profuse, bright green, paler and some- 
times purple beneath (especially the lower leaves), crenate with 5-10 large teeth, covered 
on both sides with long multicellular hairs, always hairy on veins; upper leaves usually 
acute, or subacute; floral leaves usually with angled, acute or acuminate teeth; bracts 
slightly larger than in S. ussuriensis (Rgl.) Kudo, the lower often with few teeth; calyx 
mostly densely patent -hairy and glandular; corolla 15-18 (20) mm long; nutlets curved- 
ovoid, dark brown to blackish. Otherwise similar to S. ussuriensis (Rgl.) Kudo. June- 
July. (Plate VI, Figure 4.) 
108 Deciduous and mixed forests, coppices. — Far East: Ze.-Bu., Uda, Uss. Gen. distr.: 
Korea. Described from Maritime Territory, Lien Ch'ih Ho River valley. Type in Leningrad. 

19. S. shikokiana Mak. in Bot. Mag. Tokyo, VI (1892) 54 (nomen) et XVIII (1904) 
44. - Ic: Sugawara, 111. Fl. of Saghal. IV, tab. 726. 

Perennial; plant with long filiform underground shoots; stems 5-28 cm long, erect, 
simple or slightly branching, glabrous; leaves 1-5 cm long, 1-4 cm wide, triangular-ovate, 
with truncate or subcordate, rarely angled base, obtuse at apex, coarsely dentate, with 
4-9 beveled semiovate or triangular, obtuse teeth on each margin, thin, sparsely patent- 
hairy or subglabrous above, glabrous, sometimes reddish beneath; petioles glabrous or 
patent -hairy, 0.5-3 cm long; inflorescence lax, 1-6 cm long, erect, the slender axis with 
scattered hairs above; bracts lanceolate to ovate-lanceolate, short-petioled or sessile, longer 
than pedicels, entire, very sparsely patent-hairy, the lower usually dentate; flowers small, 
5-10 mm long, white or with faintly azure upper lip; pedicels much shorter than flowers, 
suberect-divaricate, short-hairy; calyx 1 .5-2 mm long, to 3 mm in fruit, campanulate, 
sparsely and thinly patent-glandular and hispid; corolla with slightly curved tube, the limb 



73 



(109) 




PLATE VI. 1 - Scutellaria dependens Maxim., general aspect, flower, upper and lower part of fruit- 
ing calyx; 2 - S. altissima L., summit of stem, corolla, upper and lower part of fruiting calyx, nutlet; 
3 - S. tournefortii Benth., corolla, upper and lower part of fruiting calyx; 4 - S. pacifica Juz., gen- 
eral aspect, corolla, upper and lower part of fruiting calyx; 5 - S. pallida M.B., general aspect, calyx, 
corolla. 

74 



much shorter than the tube; lower Up larger than the upper, with small spots: lateral lobes 
erect, ovate-triangular, slightly shorter than the emarginate middle lobe and much shorter 
than the subpendulous 3-lobed lower one; nutlets ca. 1 mm long, slightly flattened, black, 
tuberculate. July— August. 

Grassy mountain slopes. — Far East: Sakh. (S). Gen. distr.: Japan. Described 
from Kochi prefecture. Type in Tokyo. 

Note. We have not been able to compare the Sakhalin plant with the type or any other 
material on S. shikokiana from Japan. 



Series 3. A Ibidae Juz. — All parts slightly smaller in comparison with Altissimae; 
leaves less deeply crenate-dentate; bracts longer than calyx, long-petioled; flowers whitish 
or cream-colored. 

20. S.albidaL.Mant. altera (1771) 248, s. str.; Shmal'g. Fl. II, 327, saltem p. p. - 
S. cretica Mill. Gard. Diet. ed. VIII (1768) No. 2, non L. - S. albida ssp. eu-albida 
Reching. fil. in Bot. Arch. 43 (1941) 6. - S. albida var. typica Stoj. et Stef. Fl. Bulg. ed. 
2 (1937) 854. - S. peregrina Ldb. Fl. Ross. Ill, 397, non L. - Ic: Sibth. et Sm. Fl. 
Graeca, VI, tab. 581 ; Reching. fil. 1. c. tab. I, fig. 23. 

Perennial; rhizome creeping, flexuous, light brown; stems 23-50 cm long, erect or 
slightly flexuous, mostly with obUquely ascending opposite branches, obtusely 4-angled, 
densely covered with curved recUnate to subappressed hairs, in inflorescence with sparse 
long patent and very profuse long-stipitate glandular hairs; leaves 2-5 cm long, 1-4 cm 
wide, broadly ovate, with truncate or subcordate base (in upper leaves often broadly cu- 
neate), each margin with 3-12 shallow, obtuse, frequently rounded teeth, both surfaces 
covered with fine curved hairs, the lower more densely so; petioles long, densely covered 
with short curved hairs; bracts long-petioled, broadly elliptic or ovate-lanceolate, rounded 
at apex, mucronate or acuminate, entire, mostly at least twice the length of calyx, at an- 
thesis about equaling the flowers; inflorescence 5-16 cm long, at first contracted, later 
elongating but not lax; pedicels 2-3 (4) mm long; calyx 3-4 mm long, covered with few 
long reclinate hairs and numerous stipitate glands, in fruit 7-8 mm long; corolla to 1 5 mm 
long, whitish; nutlets depressed-globose, tuberculate, the tubercles surmounted by solitary 
stellate hairs. June— August. 

Forests, riverbanks, stony taluses, limestone. — European part: Crim.; Caucasus: W. 
Transc. (Abkhazia). Gen. distr.: Med. (Italy), Bal. -As. Min. (Greece, Vifinia). Described 
from the "East." Type in London. 

Note. Rechinger (op. cit.) does not report this plant for the Crimea but vaguely refers 
the Western Caucasian (Abkhazia) plant to S. pallida M.B. from which, however, it differs 
markedly. We can find no substantial differences between our plant and the genuine S. al- 
bida L. whose sporadic distribution is also noted by Rechinger (1. c. 9). More detailed in- 
vestigation of the various vicarious races will result in subdivision of the species. 

21. S. paUida M.B. Fl.taur.-cauc. 11(1808)65. - S. albida Ldb. Fl. Ross. 111,397, 
non L.; Shmal'g. Fl. II, 327, pro max. parte. — S. albida ssp. pallida Reching. fil. in 
Bot. Arch. 43 (1941) 8. 



75 



112 Perennial; close to preceding species but leaves more deeply cordate, the dentation 
deeper and more acute; hairs on stems and petioles longer and straighter; leaves densely 
covered on both sides with longer hairs, these spreading beneath especially on the veins, 
stems more densely glandular-hairy (glands present here not only in inflorescence but also 
on upper half of sterile part of stem); bracts narrower and more acute. June — July. 
(Plate VI, Figure 5.) 

Stony slopes and taluses, roadsides in conifer (juniper and pine) forests. — European 
part: Crim. Endemic. Described from the Crimea. Type in Leningrad. 

Note. The two Crimean races of this series (S. albida and S. pallida) do not appear 
to be clearly separated from each other and are connected by intermediate (probably hy- 
brid) forms. We know the perfectly genuine S. pallida only from a few localities on the 
southern Crimean coast (for example, Yalta and Nikita). In certain respects it recalls the 
Balkan-Rumanian S. velenovskyi Reching. fil. (S. pichleri Velen. non Stapf) rather than 
S. albida, but is distinguished by its exceptionally well developed glandular hairs and its 
large, obtuse bracts. 



Series 4. VaciUantes Juz. — Like the preceding series but bracts with shorter petioles; 
flowers anthocyanin-colored. 

22. S. woronowii Juz. in Bot. mat. gerb. Bot. inst. ANSSSR, XIV (1951) 360. - S. pe- 
regrina var. sibthorpii Woron. in Busch. Marc, et Woron. Fl. Cauc. exs. fasc. XI (1909) 
8, non Boiss. et Rent. - S. sibthorpii Grossh. Fl. Kavk. Ill (1932) 287, non Halacsy nee 
Reching. fil. - S. vacillans ssp. colchica Reching. fil. in Bot. Arch. 43(1941) 15. -Exs.: 
Fl. Cauc. exs. No. 274. 

Perennial, 20-40 cm high, with an obUque or creeping rhizome, stems erect, straight or 
flexuous, usually branching nearly from base or in upper part, with obliquely ascending 
branches, distinctly 4-angled, densely covered with short spreading hairs, often almost velu- 
tinous, occasionally with some long hairs, in inflorescence with spreading long-stipitate 
glands; leaves 2-4 cm long, 1 .4-3 cm wide, 1 Vi times as long as wide, broadly ovate, sub- 
cordate or truncate at base, obtuse or acute, with 5-7 usually shallow obtuse teeth on each 
margin, densely covered on both sides with short spreading hairs, mostly subvelutinous 
beneath, the petioles densely covered with short spreading hairs; inflorescence 7-15 cm 
113 long, lax; bracts short-petioled, ovate to ovate-lanceolate, acute, entire, as long as or sUght- 
ly longer than calyx, at anthesis much shorter than corolla, densely long-hairy on both 
sides; calyx 3-4 mm long, to 6 mm in fruit, with long patent hairs and stipitate glands; 
corolla 1.4-1.7 cm long, dingy mauve; nutlets covered with stellate hairs. June. 

Stony places among shrubs. - European part (?): Crim. (see Note); Caucasus: W. 
Transc. (rarely). Gen. distr.: As. Min. (Artvin, Trabzon). Described from former Artvin 
district, Svetibar village near Artvin. Type in Leningrad, isotypes (Fl. Cauc. exs. No. 274) 
in many herbaria. 

Note. Zefirov identified with this species the Crimean plant referred by Syreishchikov 
(no doubt erroneously) to S. columnae All. We have not so far had an opportunity of 
studying Syreishchikov's plant. 



76 



Subsection 3. Salviifoliae Boiss. Fl. or. IV (1879) 681. — Bracts entire, smaller than 
cauline leaves. Low prostrate herbs, with obscurely crenate or entire leaves. 



Series 5. Ponticae Juz. — Flowers purple or violet; leaves not rugose (as opposed to 
the Eastern Mediterranean series Eu-salviifoliae Juz. with yellow flowers and conspicuously 
rugose leaves). 

23. S. pontica C. Koch in Linnaea, XXI (1848) 701 ; Boiss. Fl. or. IV (1879) 686. 
Low herbaceous perennial, with numerous slender procumbent and rooting glabrous 

brownish or dingy -violet caudices; flowering shoots 4-1 2 cm long, ascending at base, simple, 
often lilac, covered with very short (sometimes longer at the angles) recurved white hairs, in 
inflorescence with long spreading hairs and short stipitate glands, mostly with very short in- 
ternodes and hence densely leafy; leaves small, obovate, 8-1 4 mm long, 4-9 mm wide, round- 
ed or subcuneate at base, obtuse, obscurely and remotely crenate with 1-6 crenations on each 
margin, sometimes almost entire, glabrous or short- or rather long-ciliate on the margins below; 
petioles to 14 mm long, covered with short curved reclinate hairs; uppermost leaves short- 
petioled or subsessile; bracts subsessile, 3-8 mm long, 2-4 mm wide, elliptic to narrowly 
ovate, entire, longer than calyx but much shorter than corolla, ciliate on the margin, cover- 
114 ed in lower part with long, and in upper part with short hairs; flowers few, opposite, 

crowded in short one-sided loose raceme 2.5-5 cm long; calyx ca. 3 mm long, covered with 
short-stipitate glands and few long coarse white hairs; corolla 2.2-2.5 cm long, short- 
stipitate glandular and sparingly pubescent outside, the tube scabrous, curved at base, ca. 
5 mm wide at throat, hps approximately equal, purple-violet. June — July. 

Stony places and pastures in the alpine belt. — Caucasus: W. Transc. (Adzharia, moun- 
tains Khino and Trial). Gen. distr.: Bal.-As. Min. (Lazistan). Described from "Pontus 
Lazicus." Type was in Berlin. 

24. S. helenae Alb. in Otch. and Tr. Odessk. otd. Ross, obshch. sadov. 1890 (1891) 
14. - S. ponticai3.abchasica Alb. in Bull. Herb. Boiss. (1893) 261.- Ic: Alb. 1. c. 
(1891) tab. (sine No.); Kolakovskii, Fl. Abkhaz. IV, Plate V. 

Perennial; caudex semiligneous, rather slender, prostrate, branching, subglabrous, joint- 
ed, grayish-brown; leafy branches 5-12 cm high, rather slender, ascending, simple, usually 
turning Ulac, covered with short spreading recurved hairs, these interspersed above with 
stipitate glands; leaves rather distant, ofteji much shorter than internodes, 6-20 mm long, 
3-12 mm wide, larger than in S. pontica, ovate or eUiptic, truncate or often subcuneate 
at base, obtuse, with scattered short appressed hairs above, diffusely patent-hairy on the 
veins beneath, the margin with few obscure crenations, long-ciliate; lower petioles 5-1 7 mm 
long, covered with long spreading hairs; inflorescence a short lax to fairly dense one-sided 
raceme 3-5 cm long; bracts 5-10 mm long, 2.5-5 mm wide, on distinct slender petioles 
1-2 mm long, herbaceous, narrowly elliptic, longer than calyx (including pedicel), with 
scattered hairs on upper surface, densely long-cihate at margins, glandular-punctate be- 
neath, the midrib rather densely covered with long spreading hairs; calyx ca. 2-3 mm long 
with pedicel 1-1.5 mm, both densely hispid and stipitate-glandular; corolla 1.5-2.5 cm 
long, purple, hairy and glandular outside, the tube slightly curved at base. July— August. 



77 



Rocks, stony places in the iilpino bolt, alpine raitges. Caucasus: W. Transc. (Abkhazia, 
Lesser Adzharia, Karchkhal Mountain). Hndenuc. Described from Vzyb Range (Chipshir 
Mountain) in Abklia/.ia. Type (or isotype) in Leningrad. 
115 Section 3. Nevskianthe Ju/.. Flowers opposite in a.vils of ordinary leaves, turned in 
different directions or aggregated in small axillary capitate few-fiowered inflorescences and 
then bracts leaflike but much smaller. Xeromorphic subshrub,with numerous erect virgate 
stems; leaves small, entire or slightly dentate, short-petioled; stems and leaves covered with 
short hairs. A Central Asian oligotypic group. 

Note. Species of this group were connected by the authors concerned with S. multi- 
caulis Boiss. This was clearK a mistaken approach since, notwithstanding a certain super- 
ficial resemblance to that plant, they do not fit at all into the section Lupulinaria, to 
which S. multicaulis is commonK referred on account of undifferentiated leaves. Ab- 
sence of the characteristic one-sided intlorescence, not to mention differences of habitat, 
should preclude inclusion of these species in any of the preceding sections. 



Series 1. Colpodeae Juz. - Scutellum small, concave, perpendicular to upper calyx 
lip; leaves entire. 

:5. S. colpodea Nevski in Tr. Bot. inst. AN SSSR, 1, 4 (1^)37) 322. 
Subshrub with a stout woody caudex; stems many, 35-50 cm long, slightly woody at 
base, erect, slightly tlexuous, obscurely 4-angled to subcylindric, velutinous with very short 
hairs, strongly branching; lower leaves 1-1 .5 cm long, 4.5-9 mm wide, ovate, obtuse, cune- 
ate at base and tapering to short petiole; upper leaves and bracts smaller, 2-9 mm long, 
1-4.5 mm wide, ovate-lanceolate, obtuse, reclinate; all leaves grayish-green, puberulent 
(especially beneath), entire, thickish; flowers solitary in axils of bracts, forming a very 
loose inflorescence; pedicels very short, reaching 2.5 mm in fruit, erect, pubescent; calyx 
ca. 2 mm long, short-tomentose, accrescent to 3-3.5 mm; scutellum on upper lip 1.7-2 mm 
long. 1.3 mm wide, oblong-tetragonal, scarcely attenuate at apex, notched and obtusely 2- 
toothed; lower lip orbicular-obovate, very obtuse, ca. 2.5 mm across, persistent; corolla 
1.8-2.3 cm long, brownish-purple, densely hairy outside, with a dingy yellow spot on lower 
lip, the very slender purple-yellow tube slightly curved at base, ca. 1 mm thick, nutlets 
grayish-blackish, sparingly pubescent. July, 
lib Dry gypseous hills. - Centr. Asia: Pam.-Al. (Kugitang). Endemic. Described from foot- 
hills of Kugitang Mountain near the village of Kugitang. Type in Leningrad. 

26. S. striatella Gontsch. in Tr. Bot. inst. AN SSSR, 1, 1 (1933) 172. 

Subshrub; caudex thick, woody, to 3 cm across, covered with brown bark, branching; 
branches 2-4 cm long, stoutish. erect or ascending; annotinous stems of two kinds, sterile 
and flowering, numerous, erect or ascending, slenderh virgate, obscurely 4-angled, branch- 
ing in lower part, very densely gray-tomentose. with distant leaves; flowering stems (27) 
32-40 cm long, to 1 .5-2 mm across in lower part; lower leaves 8-12 mm long, 4-6 mm 
wide, rhombic-oblong-ovate, rarely ovate-lanceolate, cuneate at base, obtuse at apex, on 



78 



petioles 3-4 mm long; upper leaves smaller, ovate-lanceolate, gradually tapering to very 
short petiole; all leaves entire, slightly revolute-margined, with very short hairs above, 
canescent-tomentulose beneath, thickish; inflorescences short, spicate to subcapitate, 
3-7-flov/ered, axillary, paired, on short peduncles or subsessile, forming an interrupted 
compound inflorescence, this (\4) \ 8-26 cm long, consisting of 5-7 (9) very distant pairs 
of spicules; bracts small, ca. 4 mm long, obovate, frequently long-white-villous; pedicels 
very short; calyx 1-1 .5 mm long, densely villous; scutellum 1 mm long, to 3 mm in fruit, 
slightly emarginate, with very short hairs; corolla ca. 2.4 cm long, pale yellow or cream- 
colored, the inside and the lateral lobes of upper lip purple, the lower lip purple or yellow, 
the tube purple-streaked on the nerves inside, hairy outside; nutlets ca. 1 mm long, black- 
ish-purple, minutely puberulent. August. (Plate VII, Figure 1.) 

Stony slopes. Centr. Asia: Parn.-AI. (Tadzhikistan). Endemic. Described from Kara 
Tau Mountain at right bank of Vakhsh River north of Kurgan-Tepe near Zarangbulak, 
1 1 km north of Khodzha-Mastan Mountain. Type in Leningrad. 



Series 2. Cristatae Juz. - Scutellum large, ca. 1 cm wide, flat, forming a winglike ex- 
tension of the upper lip; leaves finely sinuate-dentate. 

27. S. cristata M. Pop. in Bot. mat. gerb. Gl. bot. sada RSFSR, V, 10 (1924J 155. - 
S. reniformis Gontsch. et Zapr. in Izv. Tadzhiksk. bazy ANSSSR, I, 1 (1933) 74. - Ic: 
Gontsch. et Zapr. 1. c. 
] 1 7 Perennial; caudex rather robust, crooked, strongly branching, covered with peeling 
brownish-gray bark; stems ca. 30 cm long, numerous, woody in lower part; branches few, 
opposite, slender, virgate, faintly 4-angled, grayish with profuse short slightly reclinate 
hairs; leaves 1-1.3 cm long, 0.9-1 cm wide, remote, much shorter than internodes, ovate- 
orbicular, rounded at base or cuneately tapering to petiole, apically rounded, obtuse, 
sparsely denticulate, prominently veined beneath, covered on both sides with short or 
very short hairs, grayish with denser pubescence beneath, short-petioled ; upper leaves 
and bracts gradually but strongly decreasing in size, the uppermost almost entire; inflores- 
cence loose, elongate (at least in fruit); bracts herbaceous, resembling cauhne leaves (see 
above); flowers 1-2 in axils of leaves, 1.2-1.3 cm long; calyx small at flowering, sparsely 
tomentose, the hps rounded-ovate, ca. 2 mm long, to 2.5 mm in fruit; scutellum narrow at 
first (1 mm), expanding into a large, reniform, coriaceous, nerved crest, 5-6 mm long, 8.5- 
10 mm wide; corolla ca. 1 cm long, greatly exceeding the calyx, brownish-green, with 
yellow lower lip, pubescent outside the tube ca. 1 mm wide at»base, to 2.5 mm long to 
throat, the upper lip obtuse, ca. 3 mm long, the lower lip ca. 2.5 mm long, broadly ovate; 
nutlets nearly globose or slightly oblong, brownish, smooth, sparingly pubescent. June- 
July. (Plate VII, Figure 2.) 

Conglomerate rocks, outcrops of red clays, red sand ravines, sandstone detritus. -Centr. 
Asia: Pam.-Al. (S. Tadzhikistan). Endemic. Described from Bal'dzhuan, rocks along 
Kizyl-Su River. Type in Tashkent. 

Note. We have not seen the type of S. cristata but, on the basis of its description, we 
have referred to it the plant collected by Zapryagaev and Tekut'ev and described in Zaprya- 
gaev's article "Novinki flory Tadzhikistana" (Novelties in the Flora of Tadzhikistan); it 



79 



was also collected in the Shuroabad area by Linchevskii and Maslennikov. Following a 
study of this material, we have made some additions to Popov's description of S. cristata. 



Section 4. Lupulinaria A. Hamilt. in Mem. Soc. Linn. Lyon, I (1832) 11. - Spicatae 
Benth. in Bot. Reg. 18 (1832) ad calcem, No. 1493. - Flowers usually in more or less 
approximate, rarely distant, verticillasters; inflorescences usually terminal, 4-angled, spi- 
cate or rarely subracemose, not one-sided, at most obscurely dorsiventral; bracts usually 
118 scarious, conspicuously differing from cauline leaves in shape, consistency, color and ves- 
ture, always sessile. Perennials or subshrubs, sometimes shrubs, with petiolate, mostly 
crenate-dentate or deeply dissected leaves. More or less xerophytic, rarely mesophytic, 
alpine plants. 

The largest section of the genus; indigenous to the Eastern Mediterranean, in particular 
to countries lying east of the region (the so-called "Ancient Mediterranean"). Only one 
subsection, Alpinae, penetrates into the Euro-Siberian holarctic subregion. 



Subsection 1 . Fruticosae Juz. — Subshrubs, with numerous, usually virgate stems; 
leaves slightly tomentose beneath or green on both sides; inflorescence spikelike, inter- 
rupted, the verticillasters (especially the lower) remote; bracts varying in shape (see char- 
acteristics of series) and in this respect similar to the bracts of Orientales Juz. An Irano- 
Turanian group. 



Series 1. Naviculares Juz. — Bracts subherbaceous, ovate-lanceolate, acute, navicular, 
with prominent keel, sparsely tomentose, eglandular. 

28. S. navicularis Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 377. 

Many-stemmed subshrub branching from base, 17-35 cm high; root long, woody, flex- 
uous, branching; perennial shoots woody, to 3 mm in diameter, covered with brownish 
bark, producing short sterile shoots and much longer fertile ones; fertile shoots many, to 

1 mm in diameter, subterete, tough, with a sparse pruinose cover of very fine hairs, gradual- 
ly passing above into inflorescence; leaves distant, much shorter than internodes (some- 
times half as long), 0.7-2 cm long, 0.3-1 .2 cm wide, ovate or the upper narrowly ovate, 
truncate or obtusely angled at base, obtuse or acute at apex, deeply or above shallowly 
and unevenly crenate with (0) 1-5 teeth on each margin, often apically entire, bright green, 
glabrous above, glaucescent beneath, with faint appressed tomentum of minute fine hairs, 
the veins indistinct, scarcely prominent beneath; petioles 0.1-1 cm long, with bloomlike 
pubescence on the underside; flowers soUtary in the axils of bracts; inflorescence 3.5- 

6 cm long, lax, the distance between pairs of flowers 0.7-1 .5 cm, the vesture of the axis 
hke that of the stems; bracts about twice the length of calyx (including pedicel), much 
119 shorter than flowers, 0.6-1 cm long, expanded 2.5-4 mm wide, ovate-lanceolate, acute, 
navicular, entire, sparsely and finely appressed-tomentose, glaucescent; pedicels ca. 1- 

2 mm long, calyx 2-3 mm long, finely appressed-hairy, whitish; scutellum ovate, in fruit 
to 6 mm long, broadly ovate, subglabrous; corolla ca. 2.5 cm long, covered outside with 



80 



sparse, very short hairs and short-stipitate glands, the tube strongly curved at base, 1.5 mm 
in diameter, rapidly expanding above and to 1 cm v^ide at throat, the upper lip ca. 9 mm, 
the lower ca. 7 mm long; nutlets 1 .5 mm long, brown, scarcely pubescent. June— August. 
(Plate VII, Figure 3.) 

Sands, sand and loess slopes. — Centr. Asia: Balkh. Endemic. Described from an area 
between Karatal River and Uch-Kul' lakes (ravine and spring at Turanglyk). Type in 
Leningrad. 

Note. Originally we referred this plant to group Oxystegiae Juz., subsection Orien- 
tales Juz. (on account of the similarity of the bracts), but we find it more appropriate to 
include it in the subsection Fruticosae. Among the species of this section known to us, 
most closely comparable would be the Iranian S. nepetifolia Benth. and, possibly, S. 
fruticosa Desf. 

29. S. chenopodiifolia Juz. sp. nov. in Addenda XIX, 344. 

Subshrub; shoots of the preceding year persistent, woody, very strong; annotinous 
shoots 20-35 cm long, pale green with a yellowish tint; leaves 1-2.5 cm long, 0.5-1.5 cm 
wide, ovate to broadly ovate, obtuse, shallowly and coarsely crenate-dentate or the upper 
almost entire, thickish, green on both sides, subglabrous above, covered beneath with scat- 
tered fine papilliform hairs; bracts ovate, the lower obtuse, nearly flat, the upper short- 
acuminate, navicular, very sparingly short-tomentose, pale green; calyx pale green in fruit, 
not turning red; corolla up to 2 cm long, yellow. Otherwise resembling the preceding 
species. June. 

Sands. — Centr. Asia: Balkh. Endemic. Described from the left bank of Kurtu River 
between Sary-Kemir and Karasu. Type in Tashkent. 



Series 2. Multicaules Juz. — Bracts scarious, ovate or broadly ovate, obtuse or acumi- 
nate, nearly flat, covered with simple or glandular hairs. 
Southwest Asian and partly Central Asian group. 

30. S. juzepczuki Gontsch. in Bot. mat. gerb. Bot. inst. ANSSSR, VII, 5 (1938) 105. 

Strongly branching many-stemmed subshrub ca. 30 cm high; perennial shoots (ramifi- 
120 cations of caudex) woody, long, ascending or suberect, to 3 mm in diameter, covered with 
brown bark, producing short sterile shoots and long flowering ones; fertile shoots many, 
slender, 1-1 .5 mm in diameter, 4-angled, covered with very fine short crisp hairs, gradually 
passing above into interrupted inflorescence; leaves remote, 13-17 mm long, 9-10 (1 l)mm 
wide, much shorter than internodes, ovate to subrhombic-ovate, rounded or angled at base, 
obtuse at apex, deeply and unevenly dentate with 3-6 teeth on each margin, green, sub- 
glabrous above, glaucescent with short appressed hairs beneath, the veins impressed above, 
prominent beneath, the petioles ca. 3-4 mm long; flowers soUtary in axils of bracts; in- 
florescence spikelike, short, loose, 14-18-flowered, 6-8 cm long, rather dense at onset of 
flowering, becoming interrupted, the pairs of flowers ca. 1 cm apart; axis of inflorescence 
covered with spreading white hairs interspersed with stipitate glands; bracts shorter than 
flowers, to 1 cm long, 8 mm wide, sessile, ovate to broadly ovate, acuminate, the lower 
crenulate, green, the upper entire, purple-green, all sparsely covered with longer simple 



81 



white spreading and shorter glandular hairs; pedicels ca. 3 mm long, pubescent; calyx 
(not including scutellum) ca. 2 mm long, short-villous; scutellum 2 mm long, 3 mm wide, 
round, slightly pubescent; corolla bright yellow, ca. 2.8 cm long, the tube ca. 1-1.6 mm in 
diameter, curved at base, dilating from about the middle to throat, the upper lip ca. 7 mm 
long, dingy green at apex, the lower ca. 5 mm long, copiously glandular outside. June. 
Scrub in the rosarium belt (altitude ca. 2100 m). - Centr. Asia: Pam.-Al. (Sanglak 
Range). Endemic. Described from Pekhe, a village on the crest of Sanglak Range. Type 
in Leningrad. 

31. S. litwinowii Bornm. et Sint. in Sintenis, It. transcasp.-pers. 1900-1901, No. 1726 
(nomen seminudum); 0. and B. Fedch. Perech. rast. Turk. V (1913) 160 (nomen); Bornm. 
in Russk. Bot. zhurn. 1 (1914) 8. Exs.: Sintenis, 1. c. No. 1726. 

Subslirub, persistent old woody shoots (ramifications of caudex) erect, stout, covered 
v^th grayish bark; herbaceous stems many, 17-25 cm long, simple or slightly branching, 
stoutish, densely covered with short fine hairs, in inflorescence velutinous with spreading 
glandular hairs; leaves commonly in 3 or 4 remote pairs, 0.3-2 cm long, 0.3-1 .2 cm wide, 
one-third to half the length of internodes, ovate, truncate at base, obtuse, shallowly crenate 
121 with 2-6 teeth on each margin, sometimes almost entire, green, sparsely covered with short 
hairs above, glaucous, heavily pubescent and densely beset with small sessile glands be- 
neath, prominently veined, on petioles 1-3 mm long, the uppermost sessile; inflorescence 
6-12 cm long, rather dense, with verticillasters 1-2.5 cm apart; flowers solitary in the axils 
of bracts, these slightly wider than cauline leaves, 0.6-1 cm long, 0.5-0.9 cm wide, much 
shorter than the flowers, broadly ovate or suborbicular, subacute or subacuminate, entire, 
green or sporadically dull purple, somewhat scarious, short-hairy and glandular; pedicels 
very short, hairy; calyx ca. 2 mm long, densely covered with rather short spreading hairs 
and glands, strongly accrescent in fruit; scutellum strongly enlarging, to 8 mm long and 
1 cm wide, rounded, emarginate-crenate, often reddish, sparingly pubescent; corolla 2- 
3 cm long, yellow, covered outside with spreading hairs and scattered short-stipitate glands, 
the tube curved at base, 1-1 .3 mm in diameter, gradually broadening to Umb ca. 1 cm in 
diameter, the upper lip ca. 6 mm long, the lower lip ca. 4 mm, dark brownish-purple. 
April-May. (Plate VII, Figure 4.) 

Sandy hiUs. - Centr. Asia: Mtn. Turkm. Endemic? Described from Kizil-Arvat, Kara- 
kal near Kutenak. Isotype in Leningrad. 

32. S. luteo-coerulea Bornm. et Sint. in Sintenis, It. transcasp.-pers. 1900-1901 (1903) 
No. 948 (nomen seminudum); Bornm. in Russk. Bot. zhurn. 1 (1914) 7. - Exs.: Sintenis, 
I.e. No. 948. 

Subshrub, with short, erect or sUghtly ascending, persistent, slender, virgate, woody, 
strongly branching, brownish shoots: herbaceous stems (or branches) numerous, 20-40 cm 
long, simple or branching, rather slender, very finely and densely short-hairy, in inflores- 
cence with longer hairs and stipitate glands; leaves many, 0.5-2 cm long, 1-5 mm wide. 
yi to ^4 the length of internodes, oblong-lanceolate, cuneately tapering to petiole, subacute, 
with 2-5 acutish teeth on each margin, dull, sparsely covered on both sides (more densely 
beneath), with short hairs and short-stipitate glands, the veins strongly impressed above, 
prominent beneath; petioles 0.2-1 cm long, finely pubescent or glandular; inflorescence 



82 



6-1 5 cm long, loose, the verticillasters to 3 cm apart; bracts 4-7 mm long, 3-5 mm wide, 
ovate, entire, tapering to a short point, green, somewhat scarious, patent-hairy, densely 
stipitate-glandular; pedicels very short or flowers subsessile; calyx ca. 2 mm long, densely 
122 patent -hairy and glandular, slightly enlarging in fruit (approximately twofold); scutellum 
ca. 2 mm long, 3 mm wide, deeply furrowed, remotely crenate, usually becoming reddish, 
sparingly pubescent; corolla 1.8-2.4 cm long, patent-hairy and sparsely stipitate-glandular 
outside, the tube slender (ca. 1 mm in diameter at base), curved, gradually dilating above, 
the limb to 5 mm in diameter, yellow, the lower lip bright blue. June-July. 

Mountain slopes. — Centr. Asia: Mtn. Turkm. (Kopet Dagh). Endemic. Described 
from vicinity of Ashkhabad. Isotype in Leningrad. 



Subsection 2. Ramosissimae Juz. - Subshrubs, in habit like Fruticosae but generally 
with more numerous stems; inflorescences terminal, small, dense, 4-angled, spicate to sub- 
capitate; bracts scarious, usually ovate, carinate. 

Note. We have only provisionally placed this unique group alongside. Fruticosae, with 
which it has always been associated (its leading species has even been acceptp'^ as a variety 
of S. multicaulis). It should be noted that the flowers of the species included here show 
greater similarity to those described for the preceding section, notably to the flowers of 
S. colpodea Nevski, than to those of Fruticosae and other Lupulinaria. Their inflo- 
rescences also suggest more affinity with S. striatella Gontsch. of the same section (i.e. 
Nevskianthe Juz.) than with Lupulinaria. The species of the Ramosissimae group 
should perhaps be excluded from Lupulinaria to which they have so far been referred 
on purely formal grounds. 

All the known species of this subsection are very closely interrelated; they vicariate 
geographically and form one natural series. 

33. S. ramosissima M. Pop. in Bot. mat. gerb. Bot. sada, V, 10 (1924) 155. - S. mul- 
ticaulis 7. glabrescens Rgl. in Tr. Bot. sada. VI, II (1880) 365. - Exs.: HFAM, No. 203. 
Subshrubs, caudex woody, strongly branching, the ramifications long, usually flexuous, 
covered with light brown bark, procumbent or ascending; stems numerous, up to 100 
(hence plant very bushy), 20-50 cm long, slender, virgate, erect, simple or often opposite- 
branched, the branches slender, obliquely ascending, blackish-lilac, 4-angled, slightly short- 
hairy in lower part, glabrous above, softly short-hairy under the inflorescence; leaves 
small, 5-20 mm long, lanceolate, acute or acuminate, the lower unevenly and acutely 
incised-dentate, tapering to short petiole, the upper entire, sessile, glabrous or at base 
123 sparingly pubescent; inflorescences many, at ends of stems and branches, 4-angled, 1-2 cm 
long (not including flowers), dense, spicate to subcapitate; bracts 5-10 mm long, ovate or 
oblong, navicular, tapering at base, acute, scarious, with prominent longitudinal ner/es, 
densely patent-hairy and glandular, sometimes becoming purple; calyx short-pediceled, 
appressed to axis of inflorescence, small, ca. 3 mm long, the lower lip short-hairy, the up- 
pe' hairy and glandular, with small hairy scutellum; corolla rather large, 2-2.5 cm long, 
purpUsh-vinaceous, with yellowish tube and a small white spot on lower lip, glandular- 
hairy outside; nutlets 1.25 mm long, gray, finely appressed-hairy. July-September. 
(Plate VII, Figure 5.) 



83 



Stony places in subalpine mountain belt. — Centr. Asia: T. Sh. (W.). Endemic. De- 
scribed from Khumson along Ugamu River. Type in Tashkent. 

34. S. intermedia M. Pop. in ByuU. Sr.-Az. Gos. univ. XII, dop. (1926) 16. - Exs.: 
HFAM,No.201. 

Subshrubs; caudex woody, strongly branching, nodose, its ramifications subimplexed, 
slender, flexuous, procumbent, covered with grayish-brown bark; stems many, 10-30 cm 
long, slender, virgate, erect, opposite-branched, densely covered with very short curved 
appressed hairs, apparently grayish-pruinose, with slender ascending or spreading branches 
nearly from base, almost leafless in upper part; leaves small, the lower and middle 0.8- 
1 .2 cm long, 6-8 mm wide, ovate or oblong, rounded or cuneate at base, acute, crenate- 
dentate with few teeth (usually 3-4 per margin), sparsely covered with fine short curved 
appressed hairs, with strongly impressed veins above, seemingly plicate, short-petioled; 
uppermost leaves much reduced, sessile, subentire; inflorescences many, at ends of stems 
and branches, 0.8-1.5 mm long (not including flowers), capitate; bracts small, 4-6 mm 
long, ovate, acuminate, herbaceous, resembling uppermost leaves, densely covered with 
stip'' ' glands interspersed with longer coarser hairs; calyx short-pediceled, 1 mm long, 
densely glandular-hairy, hairy above, with small scutellum; corolla 1.5-2 cm long, pubes- 
cent, with yellow tube and vinaceous-purple lips; nutlets 1 mm long, grayish with small 
stellate hairs. June. 

Pebbles at foot of mountains. - Centr. Asia: Syr D. (Mogol Tau), Endemic. Described 
from vicinity of Katar-Bulak. Type in Tashkent, isotypes in Leningrad and other places. 

35. S. glabrata Vved. nom. nov. — S. multicaulis i3. patens Rgl. in Tr. Bot. sada, VI, 
II (1880) 365. — S. intermedia var. glabriuscula M. Pop. in M. Popov and N. Androsov. 
Rasteniyazapovedn. Guralash i Zaaminskogo lesnich. (1936) 34. 

Perennial, similar to S. intermedia but stems with longer, more spreading branches; 
stems and leaves bright green, glabrous or sparingly pubescent, the vesture including isolat- 
ed longer hairs; corolla more intensely colored, both tube and limb slightly wider than in 
S. intermedia. June— August. 

Dry gravelly slopes. - Centr. Asia: Pam.-Al. (Turkestan Range, Zeravshan). Endemic. 
Described from Guralash forest reserve. Type in Tashkent 

36. S. squarrosa Nevski in Tr. Bot. inst. AN SSSR, 1, 4 (1937) 324. 

Subshrub, densely cespitose or pulvinate; caudex stout, woody, strongly branching, 
covered with brownish bark, the branches flexuous, crowded, subimplexed; annotinous 
stems 7.5-10 cm long, very numerous, obscurely 4-angled, erect, simple or rarely slightly 
branching, densely covered with very short and soft spreading hairs, light grayish-green; 
leaves (4) 5-9 mm long, (1.75) 3-6 mm wide, suberect to spreading, ovate or ovate-lanceo- 
late, obtuse, sparsely and shall owly crenate-dentate, slightly deflexed, appressed-tomentu- 
lose above, patent-tomentulose beneath, grayish-green, thickish, short-petioled or (the 
upper) subsessile; flowers few, aggregated in 2's-4's in short depauperate terminal spicules 
0.5-1.2 cm long (not including flowers); bracts 3-4 mm long, to 3 mm wide, ovate, acute, 
entire, distinctly keeled, densely patent-hairy and stipitate-glandular, commonly purple; 
pedicels very short, erect, to 2 mm long after flowering, hairy; calyx ca. 2 mm long, later 



84 



(125) 




PLATE VII. 1 - Scutellaria striatella Gontsch., summit of stem, flower; 2 - S. cristata M. Pop. 
summit of stem, flower, upper and lower part of fruiting calyx; 3 - S. navicularis Juz., summit of 
stem, bract; 4 - S. litwinowii Bornm. et Sint., summit of stem, upper and lower part of fruiting 
calyx, nutlet; 5 - S. ramosissima M. Pop., general aspect, bract, flower. 



85 



slightly accrescent, the upper lip with long glandular hairs, dorsally rounded-galeate, the 
lower Up very obtuse, almost eglandular; corolla 1.8-2.3 cm long, purple, the tube yellow 
127 beneath, ca. 1.5 mm wide, nearly straight, with purple spots at throat; lower lip with yel- 
low spot. July. 

Upper limit of Central Asian juniper zone. — Centr. Asia: Pam.-Al. (Kugitang). Endem- 
ic. Described from Kugitang Mountains above Kugitang village. Type in Leningrad. 

Note. In 1932, Massagetov and Masal'skii collected in Uzbekistan (Tatar-Kishlak, 
mountains south of the village) a plant very similar to S. squarrosa, but differing slightly 
in the longer, arachnoid vesture of the stems (especially in their lower part), petioles and 
underside of leaves. We have named it S. arachnoidea Juz., but the material required 
for reliable substantiation and description of this species is obviously lacking. 



Subsection 3. Orientales Juz. in Bot. zhurn. SSSR, 24, No. 5-6 (1939) 432; Juz. in 
Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 377. - Subshrubs or perennial herbs; 
leaves with white or gray tomentum beneath; flowers in spikelike uninterrupted inflores- 
cence, bracts varying in shape, subherbaceous or scarious; corolla-tube rather long, very 
slender in lower part, gradually enlarging above; lower lip of corolla frequently longer 
than the upper. 



Group 1. Oxystegiae Juz. in Bot. zhurn. SSSR, 24, Nos. 5-6 (1939) 434 (pro serie). - 
Bracts rather narrow, generally lanceolate or ovate-lanceolate, rarely ovate, very often 
more or less strongly incurved, conduplicate, navicular or at least distinctly keeled, often 
gradually tapering above to a longish point, usually tomentose without admixture of long 
hairs or glands, only very rarely long-hairy at margins (see characteristics of series Meso- 
stegiae Juz.). 

Note. When first publishing the characteristics of this group, we conceived it, as well 
as the group Platystegiae (see below), as a series. Grossgeim promoted them to the rank 
of subsections. While assuming that they differ in derivation and that the subsection Ori- 
entales (corresponding to "S.orientalis" of earlier authors sensu latissimo) is apparently 
not monophyletic, we must emphasize that, for all practical purposes, it is very difficult, 
if not impossible, to estabUsh a dividing line between these two groups. There is a certain 
number of well-defmed forms (most of them presented below) which, in our opinion, are 
distinctly hybrid in origin; their inclusion in one or the other of these groups is quite arbi- 
trary and provisional. Under the circumstances, we are obliged to retain the group Orien- 
tales and we are not in a position to raise the groups Oxystegiae and Platystegiae to 
higher status. 

While agreeing in principle with Grossgeim's division of Oxystegiae into more narrow- 
128 ly conceived series, we find it possible to accept only two of the three series that he pro- 
posed for the Crimean-Caucasian forms; to these we have added three Siberian-Central 
Asian series. 



Series 1. Araxenses Grossh. in Izv. Azerb. fil. AN SSSR, 3 (1945) 79. - Subshrubs; 



86 



stems woody in lower part, erect, virgate; leaves oblong or lanceolate, cuneately tapering 
at base, the petiole less than half as long as the blade. Southwest Asian species. 

37. S. araxensis Grossh. in Beih. z. Bot. Zentralbl. XLIV, 2 (1928) 233. - S. orienta- 
lis var. chamaedryfolia N. Pop. in Mat. fl. Kavk. IV, 3 (1916) 73, p. p. non Rchb. - 
Ic: Grossg. in Izv. Azerb. fil. ANSSSR, 3, 83, Figure 1. 

Subshrub 14-40 cm high; caudex ascending or erect, branching; leafy shoots (stems) 
many, ascending at base, then erect or curved, virgate, simple or branching, rather densely 
short-tomentose, grayish, often faintly lilac, remotely leafy, the internodes as long as 
leaves or 2-4 times as long; leaves 0.8-2.4 cm long, 3-10 mm wide, oblong or lanceolate, 
cuneately tapering at base, obtuse at apex, incised-crenate-dentate with 4-9 obliquely 
semi-ovate or oblong, obtuse or rounded teeth on each margin, the uppermost subentire, 
green or gray-green, copiously and finely appressed-hairy above, densely and thinly ap- 
pressed-gray-tomentose beneath, with shghtly prominent nerves, on petioles to 1 cm long; 
inflorescences 2.5-6 cm long, ovoid or oblong, rather loose; bracts 7-12 mm long, 1 .5- 
7 mm wide, in fruit only slightly exceeding calyx, herbaceous, ovate-lanceolate, keeled and 
commonly navicular-pHcate and curved, gradually acuminate, sparsely appressed-tomentu- 
lose, pale grayish-green, in fruit sometimes slightly suffused with lilac; calyx ca. 2 mm 
long, with short coarse tomentum interspersed with stipitate glands, ca. 5 mm long in fruit 
(including scutellum); corolla long, ca. 2.5 cm, yellow, with slender, slightly curved tube 
and short, greenish-brown lower lip, covered outside with long spreading hairs; nutlets 
1.3 mm long, gray with appressed pubescence. May -June, again in August. 

Dry stony slopes, limestone, gravelly taluses. - Caucasus: S. Transc. Gen. distr.: N. 
Iran. Described from vicinity of Dzhul'fa between Negram and Daroshan railroad stations. 
Type probably in Baku, topotypes in Leningrad. 



129 Series 2. Tauricae Grossh. in Izv. Azerb. fil. ANSSSR, 3 (1945) 79, ampl. (incl. ser. 
Boissierianae Grossh. 1. c). -Perennials; stems ascending to procumbent, not or scarce- 
ly woody at base; leaves ovate, usually truncate at base; petioles typically rather long 
(often nearly as long as blade); flowers pale yellow. Crimean-Caucasian forms. 

Note. Grossgeim included only the one S. taurica Juz. in the series Tauricae, but 
this species is so closely allied to S. stevenii Juz., with which it undoubtedly vicariates 
and from which it differs only in vesture, that there is no reason for placing these two 
species in different senes. Although the name Boissierianae Grossh. (proposed by Gross- 
geim for the series which we have combined with Tauricae Grossh. s. str.) has strict pri- 
ority, we cannot accept it as such since our interpretation of S. boissieri Sosn. differs 
from Grossgeim 's (see Note to this species). 

38. S. taurica Juz. in Bot. zhurn. SSSR, 24, Nos. 5-6 (1939) 434. - S. orientalis 
Schmalh. Fl. II, 326 et auct. fl. taur. p. p. - S. orientalis var. chamaedryfolia N. Pop. 
in Mat. fl. Kavk. IV, 3 (1916) 73, p. p. non Rchb. - S. orientalis var. taurica N. Pop. 
81, p. p. - Ic: Grossg. in Izv. Azerb. fil. ANSSSR, 3, 85, Figure 4. 

Subshrub 10-25 cm high; caudex woody, strongly branching, creeping; stems numer- 
ous, ascending at base, flexuous, branching; stems and pefioles densely covered with long 



87 



soft crisp spreading hairs; internodes short to medium; leaves on petioles 3-9 mm long, 
small, 5-14 mm long, 3-10 mm wide, ovate, w^ith truncate or slightly tapering (broadly 
cuneate) base, with 3-7 long, usually narrow, rather unequal, obtuse, often retrorse teeth 
on each margin, grayish above, softly white-tomentose-villous on both sides, very densely 
so beneath, with rather distinct veins; inflorescences ca. 3 cm long in flower, ca. 6 cm in 
fruit, rather dense at first, becoming loose; lower bracts ca. 1 cm long, 4 mm wide, narrow- 
ly ovate to ovate-lanceolate, incurved and condupHcate, gradually tapering to a longish 
point, densely tomentose, grayish-green; calyx ca. 2 mm long; corolla to 2 cm long, yellow, 
densely hairy outside. (June) July— September. (Plate VIII, Figure 1.) 

Stony slopes and placers, outcrops of limestone, steppes. — European part: Crim. (west- 
ern part of peninsula; region of Sevastopol' and southern coast to Simeniz). Endemic. 
Described from Kikeneiz. Type in Leningrad. 
130 Notes. 1) S. orientalis var. taurica N. Pop., a "variety" established by Popov, does 
not in any way conform to our species; it merely represents specimens of three different 
species, S. taurica Juz., S. stevenii Juz. and S. heterochroa Juz., mostly encrusted 
with limestone or chalk. 

2) Grossgeim reported S. taurica for the Caucasus (Novorossisk area), but the report 
was based on a misunderstanding. Apparently for this reason he set up this species as a 
monotypic series (see above). 

39. S. stevenii Juz. in Bot. zhurn. SSSR, 24, Nos. 5-6 (1939) 435. - S. orientalis 
Schmalh. Fl. II, 326 et auct. fl. taur. p. p. non L. — S. orientalis var. chamaedryfolia 
(nonRchb.)et var. taurica N. Pop. in Mat. Fl. Kavk. IV, 3 (1916)73 et 81, p.p. 

Perennial; similar to the preceding species, but stems and petioles appressed-tomentu- 
lose; leaves 0.6-1 .4 cm long, 3-9 mm wide, sparsely grayish-green-tomentose above, dense- 
ly white-tomentose beneath, with completely concealed veins and usually short straight 
broad teeth; bracts 5-13 mm long, 2-4 mm wide, densely appressed-tomentose, grayish; 
calyx tomentose. Otherwise, as in S. taurica Juz. July— August. 

Dry clayey and stony slopes, limestone outcrops. — European part: Crim. (foothills in 
vicinity of Simferopol and Bakhchisarai). Endemic. Described from Simferopol. Type 
in Leningrad. 

40. S. tatianae Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 380. - S.hypo- 
polia et S. steveni Grossh. in Izv. Azerb. fil. AN SSSR, 3 (1945) 80, non Juz. 

Perennial; alHed to S. stevenii Juz.; differing in the slightly broader leaves, 0.7-1. 7cm 
long, 0.5-1.1 cm wide (usually twice as long as wide); bracts also somewhat wider (fully 
expanded 4-8 mm wide, 8-1 1 mm long), acute or short-acuminate. 

Gravelly mountain slopes. - Caucasus: W. Transc. (Novorossisk). Endemic. Described 
from Markotkh Range. Type in Leningrad. 

41. S. raddeana Juz. in Bot. zhurn. SSSR, 24, Nos. 5-6 (1939) 431 (in adnot.). - S. 
boissieri Grossh. in Izv. Azerb. fil. AN SSSR, 3 (1945) 79, non Sosn. in sched. neque 
Grossh. Fl. Kavk. Ill (1932) 286; Kharadze in Tr. Tbil. Bot. inst. XII (1948) 1 1, p. p. - 
S. orientalis var. chamaedryfolia N. Pop. in Mat. Fl. Kavk. IV, 3 (1916) 73, p. p. non 
Rchb. - Ic: Grossg., op. cit. (1945) 85, Figure 2. 



Subshrub 12-22 cm high, with woody creeping caudex; stems numerous, ascending at 
131 base, often flexuous; leafy shoots mostly reddening, covered with short crisp appressed 
hairs, rather densely above, in lower part frequently subglabrous; internodes generally 
elongate; leaves medium or small, 0.7-1.8 cm long, 0.4-1 cm wide, ovate to oblong-ovate, 
with truncate or rarely broadly cuneate base, coarsely and rather deeply crenate-dentate 
with 4-6 unequal obtuse teeth 1-3 mm long on each margin, green, glabrous above, finely 
white-tomentose with indistinct lateral veins beneath; petioles 1-5.7 mm long; inflores- 
cences rather dense at first, 2.5-3.5 cm long, later slightly interrupted; bracts loosely 
disposed, small, the lower 0.9-1 .2 mm long, 4-6 mm wide, ovate, tapering at apex to a 
fine sometimes curved point, green, finely puberulent; calyx densely hairy or tomentose; 
corolla large, to 3 cm long, yellow, densely hairy outside. 

Mountain steppes and subalpine meadows. - Caucasus: Cisc. (center of Main Range). 
Endemic. Described from Dzhvari-VozeU village in Tushetia. Type in Leningrad. 

N o te. The quesfion of the correct name of this species and the real meaning of the 
name S. boissieri Sosn. is so complicated that we are devoting a separate article to it. 
Here it should only be stressed that Grossgeim, the first to describe this species, combined 
under this name two plants that have a certain resemblance in habit, but differ in other 
respects. In his "Flora Kavkaza" (Flora of the Caucasus), Grossgeim reported this species 
for "Ter." (where our S. raddeana and S. leptostegia grow) and "Dag." (where another 
species occurs). In his very brief descripfion he does not mention the bracts. In establish- 
ing our S. raddeana in 1939, we preserved the name S. boissieri for the Dagestan plant 
(which we now call S. granulosa Juz.) and placed the latter in the series Platystegiae. We 
were prompted in our decision by Grossgeim's indication of the height of the stems of S. 
boissieri as being 30-50 cm (in our material on S. raddeana the stems did not exceed 
22 cm but in the Dagestan plant very high stems were observed). It should also be men- 
tioned that Grossgeim himself (ibid.) placed S. orientalis f. elongata N. Pop. in the 
synonymy of S. boissieri Sosn. This form includes as type the Dagestan plant and bears 
no relationship to our S. raddeana Juz. However, in spite of established usage in botani- 
cal nomenclature, Grossgeim did not accept our point of view in his later work and appHed 
the name S. boissieri Sosn. to our S. raddeana (including the latter name in the synony- 
my of S. boissieri) and (incorrectly, in our view) referring S. boissieri of our conception 
to his S. daghestanica (see below). We must categorically oppose Grossgeim's position 
132 which only introduced confusion into the nomenclature of the two species under discussion. 

In order to avoid this confusion we refrain from using the name S. boissieri Sosn. ex 
Grossh. 

42. S. leptostegia Juz. in Bot. mat. gerb. Bot. inst. ANSSSR, XIV (1951) 379. - S. 
boissieri Sosn. in sched. Herb. Inst. Bot. Tbilis. p. p. vix autem Grossh. Fl. Kavk. Ill 
(1932) 286; Kharadze in Tr. Tbil. Bot. inst. XII (1948) 10 s. str. 

Subshrub 12-25 cm high, with short woody caudex and 1-6 ascending stems; shoots 
dark violet, subglabrous, lustrous in lower half, with scattered, very short hairs above, 
rather densely hairy and whitish at summit; internodes of medium length; leaves on 
petioles 3-5 mm long, medium-sized, 6-20 mm long, 3-10 mm wide, ovate or narrowly 
ovate, mostly truncate at base, incised-crenate-dentate with 3-5 obtuse teeth to 3 mm long 
on each margin, glabrous above, thinly white-tomentose beneath, with indistinct lateral 



89 



veins; inflorescence loose, 3-5 cm long at the onset of flowering; bracts loosely disposed, 
small, the lower 5-7 mm long. 1 .5-2.5 mm wide, lanceolate, long-acuminate, green, with 
very sparse tomentum of short fine appressed hairs; corolla rather small, 2-2.5 cm long, 
lemon-yellow, with a greenish-brown spot on lower lip, short-hairy and glandular outside. 
August. 

Stony mountain slopes. — Caucasus: Cisc. Endemic. Described from Terek River 
valley between Chumi and Lars. Type in Leningrad. 



Series 3. Pulchellae Juz. — Rather low perennials, leaves small, mostly tomentose on 
both sides, more densely beneath; flowers disproportionately large; corolla anthocyanin- 
colored or variegated. Siberian and Central Asian species. 

43. S. grandiflora Sims, Curt. Bot. Mag. XVII (1803) tab. 635; Kryl. Fl. Zap. Sib. IX, 
2299. - S. pulchella Bge. Suppl. alt. (1836) 42; id. in Ind. sem. hort. Dorp. (1840) 
VIII. - S. orientalis 7. microphylla Ldb. Fl. Ross. Ill (1849) 395. - Ic: Sims, 1. c. 

Perennial; root (or caudex) woody, flexuous, more or less branching; stems numerous, 
10-20 cm long, flexuous, spreading, branching, short-crisp-hairy, often turning lilac; leaves 
5-15 mm long, 3-16 mm wide, ovate or broadly ovate to suborbicular, truncate or slightly 
cordate at base, obtuse, rather shallowly incised-crenate-dentate at apex with 2-5 orbicular 
or oblong teeth on each margin, finely appressed-tomentose on both sides, grayish-green 
133 above, whitish beneath with short fine crisp hairs, with petioles to 1 .2 cm long, approxi- 
mately as long as or shorter than blade, the veins impressed above, inconspicuous beneath; 
inflorescences dense, commonly few-flowered, nearly 4-angled, 2.5-4 cm long; bracts 0.5- 
1 cm long, 1.5-4 mm wide, 3-4 times the length of calyx, ovate-lanceolate or lanceolate, 
navicular, attenuate toward base, acuminate, slightly keeled, entire or often with few small 
teeth (mostly one on each margin), with short crisp appressed hairs on both sides, often 
violet; calyx ca. 2 mm long, densely white-hairy, with subreniform concave scutellum; 
corolla (1 .5) 2-2.5 cm long, sparsely hairy outside, pink-violet or purple, the paler tube 
curved at base through nearly right angle, the upper lip twice as long as its lateral lobes, 
barely as long as the lower lip; nutlets ca. 1.5 mm long, trigonous-ovoid, very finely tuber- 
culate, black but appearing grayish because of the dense cover of stellate hairs. June— July. 
(Plate VIII. Figure 2.) 

Stony and gravelly slopes, rocks, pebble-beds. - West Siberia: Alt.; East Siberia: Ang.- 
Say. (mouth of Kemchik). Gen. distr.: Mong. Described from a specimen grown from 
seed obtained in Siberia. Type possibly in London (or lost?). 

44. S. tuvensis Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 389. 
Perennial, 10-25 cm high, closely allied to S. grandiflora Sims (with which it has so 

far been confused); root very sturdy, to 1 .5 cm in diameter; stems ascending to nearly 
erect, curved; leaves mostly oblong-ovate, 0.5-2 cm long, 2.5- 10 mm wide, usually some- 
what attenuate at base, often acute at apex, deeply incised-crenate-dentate with oblong 
teeth, often slightly recurved, gray-green, prominently veined beneath, flowers in a loose 
inflorescence; bracts lanceolate, grayish-green (not lilac); corolla blue; nutlets minutely 
tuberculate, black. Otherwise, similar to S. grandiflora Sims. June— August. 



90 



Stony river valleys, steppes. - East Siberia: Ang.-Say. (Tuva ASSR). Endemic? 
Described from Ulu-Khem valley near Kyzyl. Type in Leningrad, isotype in Tomsk. 

45. S. turgaica Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (i951).390. 

Perennial; root woody, vertical, branching; stems many, 12-25 cm long, procumbent, 
strongly and repeatedly branching, crisp-tomentulose, usually turning lilac; leaves small, 

134 0.4-1 cm long, 2-8 mm wide, ovate to broadly ovate, truncate or obtusely angled at base, 
generally acute at apex, incised-crenate-dentate, with 3-5 irregular rounded or oblong, 
often recurved teeth on each margin, fmely gray- or white-tomentose on both sides with 
short crisp hairs, the veins strongly impressed above, prominent beneath; petioles 1-5 mm 
long, appressed-tomentulose; inflorescences few-flowered, lax, 2-3 cm long; bracts 8- 

12 mm long, 2-5 mm wide, ovate, obscurely navicular to nearly flat, keeled, long-acumi- 
nate, entire, appressed-tomentulose, with short spreading hairs on the margin, usually 
violet; calyx ca. 2 mm long, tomentose; corolla 1.5-2 cm long, covered outside with short 
simple mostly appressed hairs and stipitate glands, the upper lip, its lateral lobes and the 
back of tube purple, the lower Up and the rest of tube yellow; nutlets ca. 1 mm long, 
curved-ovoid, grayish, fmely foveolate. May— June. 

Limestone and clayey-gravelly slopes. - Centr. Asia: Ar.-Casp., Balkh. (western 
border). Endemic. Described from Espe-Sai ravine in Turgai region. Type in Leningrad. 

Note. The shape of the bracts of this species is not characteristic of Oxystegiae. It 
may be a hybrid; the strikingly bicolored flowers would suggest an intermediate position 
between S. grandiflora Sims and S. sieversii Bge. 

46. S. karkaralensis Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 391. 
Perennial; root woody, angled; stems many, 7-38 cm long, procumbent, branching, 

slightly geniculate, often flexuous, frequently becoming slightly purple, densely covered 
with short spreading white hairs; leaves 3-13 mm long, 2-10 mm wide, ovate to triangular- 
ovate, truncate or sUghtly tapering at base, obtusely or acutely angular, obtuse, irregularly 
crenate-dentate, with 2-4 triangular, semi-ovate or oblong, often slightly recurved, obtuse 
or acute teeth on each margin, grayish-green, finely appressed-tomentose, with deeply im- 
pressed veins above, slightly white-tomentose, with prominent veins beneath; petioles 
villous-tomentose with spreading hairs, half as long as to nearly equaling the blade; inflo- 
rescence at first ca. 2-3 cm long, few-flowered, in fruit to 5 cm long, lax; bracts to 9 mm 
long, 5 mm wide, ovate, subobtuse to short-acuminate, carinate, entire, densely covered 

135 with short thin tomentum interspersed with long white spreading hairs, commonly dark 
lilac, loosely imbricate; flowering calyx ca. 2 mm long, densely covered with spreading 
white hairs; corolla 1.8-2.7 cm long, bright purple, puberulent and short-glandular out- 
side; tube paler, slender, slightly curved; upper Up longer than the lower; nutlets more 
than 1 mm long, ovoid, cinereous. May— July. 

Slopes of stony coniform hills, mainly on sandy soil. — West Siberia: Irt.; Centr. Asia: 
Balkh. Endemic. Described from hills near Atasu River and Bektau-Ata Mountain. Type 
and paratype in Leningrad. 

Note. Like the preceding species, S. karkaralensis Juz. has bracts of intermediate char- 
acter; it is thus distinguished from typical Pulchellae and from Platystegiae. In this feature, 
and also in shape and dentition of its leaves as well as vesture of its stems and petioles, it 



91 



is somewhat reminiscent of S. sieversii Bge. We consider it more than probable that both 
S. karkaralensis and S. turgaica are the result of hybridization between S. grandiflora 
Sims and S. sieversii Bge. (see also Note to S. sieversii Bge.). 

47. S. nepetoides M. Pop. in Bot. mat. gerb. Bot. sada, V (1924) 155 (invaHde publ); 
Yuz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 395. 

Perennial with slender branching rhizome; stems slender, weak, procumbent, distaUy 
simple, 5-15 cm long, gray, densely patent-hairy, with remote nodes; leaves small, 5- 
10 mm long and wide, broadly ovate, subcordate at base, coarsely crenate with 3-5 crena- 
tions on each margin, gray-villous on both sides with rather long subappressed hairs, with 
impressed nerves above; petioles short, white -villous; spikes terminal, at first short, sub- 
capitate, in fruit to 5 cm; bracts shorter than corolla but much longer than calyx, ca. 
7 mm long, 3 mm wide, herbaceous, oblong-lanceolate, navicular, acute, slightly tomen- 
tose, in young fruit to 1 cm long, later falling away; pedicels erect, as long as the small 
calyx, both short-hairy; scutellum small, erect, ovate, obtuse; corolla 1.2-1 .5 cm long, 
reddish-yellow, hairy outside, lower lip with a yellow dark-punctate spot; nutlets small, 
136 ca. 1 mm long, obovoid, rounded dorsally, with an oblong ventral protuberance, covered 
with appressed stellate hairs. June— August. 

Stony mountain slopes, taluses (in Central Asian juniper zone). - Centr. Asia: Pam.-Al. 
(northern slope of Alai Range and Sary-Tau Mountain). Endemic. Described from basin 
of Shakhimardan River, Kichkin River. Type in Tashkent, isotypes in Leningrad. 

Note. This and the following three species, while similar in habit to the preceding, 
differ from them in corolla color and are unlikely to be closely related to them. They 
should perhaps be segregated as series Pictae Juz. 

48. S. picta Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 393. 
Perennial, with woody root; stems many, procumbent, branching, 10-20 cm long; 

shoots (and petioles) densely covered with soft spreading hairs; internodes mostly abbre- 
viated; leaves small, 5-12 mm long, 4-9 mm wide, ovate or broadly ovate, broadly cuneate 
to subtruncate at base, coarsely crenate with 3-5 obtuse unequal often deeply cut (1- 
2 mm) crenations on each margin, coarsely gray-tomentose on both sides, often almost 
villous beneath, the nerves impressed above, prominent and patent-hairy beneath; petioles 
1 .5-6 mm long; inflorescence 2-3 cm long, at first dense, becoming looser; lower bracts 
ca. 10 mm long, 3 mm wide, lanceolate or narrowly ovate, tapering at base, acuminate at 
apex, conduplicate, slightly incurved, tomentulose, long-hairy at margin, grayish-green or 
dark purple, loosely imbricate; calyx at flowering ca. 2 mm long, very densely white- 
villous-tomentose; corolla ca. 1.8 cm long, pale yellow, the upper lip vinaceous inside, 
with a pair of darker longitudinal yellow-rimmed stripes, the lower lip minutely vinaceous- 
striped and spotted, the tube finely streaked and punctate inside, densely covered with 
rather long slightly crisp hairs. June -July. (Plate VIII, Figure 3.) 

Habitat unknown. -Centr. Asia: Pam.-Al. (Zeravshan). Endemic. Described from 
KuU-Kalon Lake. Type in Leningrad. 

49. S. flabellulata Juz. sp. nov. in Addenda XIX, 344. 

Perennial; root vertical, woody; caudex short, nodose; stems many, branching. 



92 



137 flexuous, pale, passing above into ascending, lilac- tinged, leafy, annotinous shoots, 4- 
10 cm long, densely covered with short curved hairs; internodes short; leaves crowded, 
small, 0.4-1 cm long, 0.3-1 cm wide, broadly ovate to orbicular-rhombic, with broadly 
cuneate base and obtuse apex, incised-crenate-dentate with 3-6 small crenations on each 
margin, gray with short coarse tomentum on both sides, the veins very faintly impressed 
above, prominently flabellate beneath; petioles 1-5 mm long, very densely covered with 
short spreading hairs; inflorescence 1.5-2 cm long, dense in flower; bracts ovate-lanceo- 
late, obtuse, arched, navicular-conduplicate, lilac-purple, the lower ca. 1 cm long, 4 mm 
wide, with isolated acute teeth, the upper entire, densely covered with thick short curved 
hairs, the margin beset with long spreading white cilia; calyx small, villous-hairy, in fruit 
ca. 4 mm long; corolla 0.7-1 .5 cm long, very plain, dull yellowish (cream-colored), dense- 
ly covered outside with short spreading hairs and stipitate glands, the upper Up violet- 
purple in upper part, the lower Up with violet-purple spots and lines, the tube finely lined. 
August. 

Mountain slopes, limestone, taluses. — Centr. Asia: T. Sh. (Talass Ala-Tau). Endemic. 
Described from upper reaches of Ul'kun-Kaindy. Type in Leningrad. 



Series 4. Eu-oxystegiae Juz. — Genuine subshrubs, the mostly ascending stems 
strongly lignified in lower part; leaves ovate; flowers mostly pale yellow. Central Asian 
forms (W. Tien Shan and Pamir-Alai). 

50. S. comosa Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV, (1951) 381 . 

Subshrub, with woody vertical flexuous root; stems few, 10-32 cm long, arched- 
ascending to suberect, strongly branching at base, with few usually short branches above, 
grayish-green with sparse short curved or slightly crisp hairs, sometimes turning lilac (es- 
pecially in lower part); leaves 0.5-3 cm long, 0.4-2 cm wide, ovate or triangular-ovate, 
with truncate or sometimes subcordate base, obtuse (the lower) to acute, shallowly 
crenate-dentate with 3-9 uneven rounded angular mostly obtuse teeth on each margin, 

138 green or grayish-green above, covered with very short, fine appressed hairs, evenly densely 
and finely white- or gray-tomentose beneath, the tomentum completely concealed but the 
veins usually prominent; petioles 0.2-2 cm long, with short crisp hairs; inflorescence few- 
flowered, 2-4 cm long at the onset of flowering, with a conelike tuft of nonflowering 
bracts at summit, finally elongating to 10 cm; bracts 7-12 mm long, 2.5-4.5 mm wide, 
lanceolate or ovate-lanceolate, navicular, gradually tapering at apex to a long point, gray- 
green with rather dense fine appressed tomentum, eglandular, often turning lilac; calyx 
tomentose, in flower 2-2.5 mm long; corolla 1.2-3 cm long, yellow, the lateral lobes of 
upper Up dark yeUow, these and the lower lip finally turning dark brown, the tube slender, 
curved, densely covered outside with dense tomentum interspersed with few short-stipitate 
glands; nutlets ca. 1.5 mm long, angled-ovoid, densely white-hairy. April— June. (Plate 
VIII, Figure 4.) 

Stony slopes, rocks, taluses, pebble-beds. — Centr. Asia: T. Sh., Syr. D., Pam.-Al. 
Endemic. Described from Arslanbob. Type in Leningrad. 

51. S.oxystegia Juz. inBot.zhurn. SSSR, 24, Nos. 5-6 (1939) 431. -S. orientalis 
var. mollis Briq. in Bot. Tidskr. 28 (1907) 233. 



93 



Perennial, very close to S. comosa Juz., 10-20 cm high; leaves 0.6-1.5 cm long, 0.5- 
1.2 cm wide; inflorescence short, 3.5-7 cm long, more compact, without a distinct brac- 
teal tuft at summit; bracts 7-12 mm long, 5-7 (8) mm wide, strongly convex, often obs- 
curely navicular, rather abruptly attenuate from broadly ovate base to a slender point 
(this shorter than in S. comosa), loosely and softly villous-tomentose, the margins beset 
with rather long, horizontally spreading, almost straight hairs; coroUa 2.5-2.7 cm long, 
essentially pale yellow but upper side of tube more or less suffused with purple. Other- 
wise, like S. comosa Juz. May — beginning of June. 

Stony slopes. - Centr. Asia: Pam.-Al. (near Osh). Described from Osh. Type in 
Leningrad. 

Note. As strictly conceived, this species is very stenotopic and is so far known from 
only two localities: Suleiman-Takhta Mountain (0. and B. Fedchenko, Korzhenevskii, 
Butryakov and probably O.W. Paulsen) and the highway leading to Naukat, 12 km south 

139 of Osh (0. Knorring). Apparently these habitats lie at the periphery of the very extensive 
distribution area of S. comosa Juz. which we did not originally distinguish from the true 
S. oxystegia. 

52. S. microdasys Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 382. - Ic: 
Juz., op. cit. 383, Figure 2. 

Subshrub, with vertical woody flexuous branching root; stems few, 10-25 cm high, 
ascending, erect or suberect, usually subvaricately branching from base, woody in lower 
part, profusely branching above, the branches erect, leafy, often lilac-tinged, sparsely cov- 
ered with very short curved white hairs; leaves 0.7-2.6 cm long, 0.4-1 .8 cm wide, ovate to 
subrhombic, the lower sometimes faintly cordate or truncate at base or mostly all leaves 
cuneately tapering to petiole, obtuse or (the upper) acute, very shallowly crenate-dentate 
with 4-7 short strongly beveled obtuse teeth on each margin, green or grayish-green, sparse- 
ly and very finely tomentulose above, covered beneath with very dense fine appressed 
white or gray tomentum, a lilac tinge sometimes showing in lower part, the prominent 
veins diverging almost flabellately from base; petioles 1-8 mm long, sparsely appressed- 
tomentose; inflorescence ca. 3 cm long, in fruit to 7 cm; bracts ovate or narrowly ovate, 
concave but not navicular, tapering at base, entire, obtuse or slightly acuminate at apex, 
finely appressed-tomentulose, eglandular; calyx small, accrescent in fruit to 7 mm long 
(including scutellum), tomentulose, eglandular; corolla ca. 2.5 (to 2.8) cm long, sparsely 
covered outside with short tomentum interspersed with few indistinct short-stipitate 
glands, yellow, the lateral lobes of upper lip often reddish, the lower lip with a large 
reddish-brown spot; nutlets to 1.5 mm long, angled-ovoid, densely white-appressed- 
pubescent. May. 

Low plains and steppe ridges. - Centr. Asia: Syr D. Endemic. Described from near 
Saryagach station in S. Kazakhstan. Type and paratypes in Leningrad. 

Note. The bracts of this and the next species have the characteristic vesture of Eu- 
oxystegiae, but in shape they approach Mesostegiae Juz. 

53. S. androssovii Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 397. 

140 Perennial, with flexuous, woody root; caudex short, nodose, ascending or procumbent, 
with woody ramifications; stems numerous (more than 12), herbaceous, procumbent or 



94 



ascending, 10-18 cm long, partly sterile, sparsely covered with very short slightly crisp 
hairs, usually turning lilac; leaves 7-18 mm long, 6-12 mm wide, subrhombic, cuneately 
tapering at base to short and broad petiole, subacute to mostly obtusish, in upper part 
with few (2-5 on each margin) large, uneven, rather deeply incised, obtuse teeth, gray- 
green on both sides, covered with long, slightly crisp hairs (more densely beneath), the veins 
rather indistinct; inflorescence 2-2.5 cm long, few-flowered; bracts to 1 1 mm long, 9 mm 
wide, ovate, distinctly point-tipped, sparsely appressed-tomentulose, eglandular, sporadi- 
cally lilac, the lower with occasional large teeth; calyx rather densely short-hairy, eglan- 
dulose; corolla ca. 2.5 cm long, yellow, covered outside with short hairs and very short- 
stipitate glands, the upper hp and lateral lobes lilac-tipped, the tube long and curved; 
nutlets unknown. May— June. 

Sandy and gravelly hills, pebble-beds, dry riverbeds. - Centr. Asia: T. Sh. (foothills of 
Kara-Tau Mountains). Described from Balamurun survey mark near ChiiU station. Type 
in Leningrad. 

Note. This species could be included in the next series; it closely approaches S. kurs- 
sancvii Pavl. and S. titovii Juz. 



Series 5. Mesostegiae Juz. — Bracts more or less distinctly keeled, navicular, as in 
the preceding series (although usually slightly broader), but with vesture consisting of 
three components: imperfect tomentum of rather coarse crisp hairs; simple, long, soft 
hairs (these sometimes nearly absent); small stipitate glands (usually very few). 

Note. In this series we have provisionally set apart a number of Central Asian species 
which vicariate geographically with each other and are morphologically intermediate be- 
tween the series Eu-oxystegiae and Eu-platystegiae. They may be products of hybridiza- 
tion, of the kind represented in the Caucasus by such forms as S. granulosa Juz. and 
S. rhomboidalis Grossh. In Central Asia, however, these hybridization processes were 
of far greater scope and in places where supposedly hybrid species of the series Mesoste- 
141 giae occur, it is impossible at present to find the typical "parent" forms (with the excep- 
tion of forms that grow alongside S. titovii Juz. in the Chu-IU Mountains and also border 
upon S. transiliensis Juz. 

54. S. mesostegia Juz. in Hot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 385. - S. 
subpinnatifida Iljin in sched. non S. orientalis var. pinnatifida Ldb. 

Perennial; root woody, vertical, flexuous; stems few, 10-25 cm long, ascending, usually 
slightly flexuous, strongly branching at base, sparingly above, sparsely covered with very 
short crisp hairs, often turning lilac; leaves 0.7-2.5 cm long, 0.4-1 .5 cm wide, ovate or 
oblong, usually tapering at base, obtuse or acute at apex, deeply (to 1/3 leaf width) incised- 
dentate with 4-8 long digitate obtuse teeth on each margin, sparsely covered above with 
short fine flexuous hairs, densely gray-appressed-tomentose beneath, distinctly though not 
prominently veined; petioles 1-17 mm long; inflorescence at first 3-5 cm long, elongating 
to 10 cm; bracts to 0.8-2 cm long, 0.4-1 cm wide, narrowly ovate, attenuate at apex to a 
rather long point, with prominent keel and lateral nerves, usually curved and often navicu- 
lar, with sparse short tomentum, usually turning lilac (especially in upper part), the margin 
with long simple hairs interspersed with occasional short stipitate -glands; calyx in flower 



95 



(143) 




PLATE VIII. 1 - Scutellaria taurica Juz., general aspect, flower, fruiting calyx, nutlet; 2 -S gran- 
diflora Sims, general aspect, flower; 3 - S. picta Juz., flower; 4 - S. comosa Juz., flowering stem, 
bract, flower, fruiting calyx, nutlet. 



96 



to 3 mm long, hairy and abundantly glandular; corolla 2-2.8 cm long, hairy and densely 
stipitate-glandular outside, yellow, the upper (and lower) lip very often turning purple, 
the lateral lobes of upper lip brown; nutlets to 2 mm long. May— July. 

Stony and gravelly slopes, steppes, pebble-beds. - Centr. Asia: T. Sh. (foothills of 
Kirgiz and Talass Ala-Tau, Ichkele-Tau Range). Described from northern foothills of 
Kirgiz Ala-Tau near Tort-Kul Lake and from the vicinity of Dzhambul. Type and para- 
type in Leningrad. 

55. S. kurssanovii Pavl. in Byull. Mosk. obshch. isp. prir. XL VII (1938) 82. 
Perennial, with robust woody root; caudex woody, with short procumbent or ascend- 
ing ramifications; annotinous shoots (stems) numerous, 20-25, herbaceous, slightly as- 

142 cending at base or some suberect, 10-20 cm high, densely patent-hairy, the hairs short, 
crisp; leaves 7-15 mm long, 3-10 mm wide, ovate or triangular-ovate, cuneate or rarely 
obtuse at base, acute at apex, with 3-6 large obtuse or rounded crenations on each mar- 
gin, grayish with very profuse crisp hairs, more densely and coarsely tomentose beneath 
and here sometimes whitish, with slightly protruding midrib and lateral nerves; petioles 
slender, shorter than blade, patent-hairy; inflorescence 2-3 cm long, 6-10-flowered, dense 
or finally rather loose; bracts ovate or oblong, short-acuminate or acute, entire, often col- 
ored in upper half, with long spreading crisp hairs and few short stipitate glands, nearly 
flat; calyx small, subsessile, with short glandular and long hairs; scutellum large, rounded, 
glabrous; corolla large, 25-30 mm long, tube long, slender, curved, yellow, on the outside 
intensely dark purple (hke the appendages), puberulent; upper lip covered outside with 
long hairs; lower lip yellow, with a dark brown spot at base or plain dark brown; anthers 
exserted, hairy; nutlets ca. 1 mm long, whitish-puberulent. May. 

Stony steppes. - Centr. Asia: T. Sh. (Kara-Tau Mountains). Endemic. Described from 
Kainar terrace (from Arkhireev collections). Type (Pavlov, Number 303 1) in Moscow. 

56. S. titovii Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 388. 
Perennial; root woody, flexuous; caudex strongly branching; stems 15-25 cm long, 

numerous, ascending, imperfectly tomentose with rather profuse long spreading hairs, 
green or faintly anthocyanin-colored; leaves 0.6-2.3 cm long, 0.4-1.2 cm wide, ovate, 
with obtusely angled or nearly cuneate base, tapering to long slender hairy petiole to 2 cm 
long, deeply incised-dentate with 2-6 (commonly 4) oblong obtuse and often recurved 
teeth on each margin, with short scattered very fine slightly crisp hairs above, imperfectly 
tomentose beneath with sparse fine appressed hairs, green at both sides or young leaves on 
145 lateral branches usually whitish; inflorescence at first 3 cm long, in fruit to 6 cm, loose; 
bracts small or medium, after flowering to 1.5 cm long, 7 mm wide, slightly scarious, cov- 
ered with spreading incomplete tomentum and numerous spreading simple hairs of moder- 
ate length, more profusely along margins, with few intermixed short-stipitate glands, pale 
green or sometimes faintly lilac; calyx at first small, rather densely and short-hairy, glan- 
dular, in fruit approximately as long as bracts; corolla to 2.5 mm long, covered outside 
with fine short hairs and stipitate glands, yellow, the lateral lobes and the tip of upper lip 
dark purple, the tube slender, slightly curved; nutlets to 2 mm long, grayish- white. 
June— July. 



97 



Gravelly and stony places. — Centr. Asia: T. Sh. (western part), Balkh. (Chu-Ilii 
Mountains). Endemic. Described from gorge of Aktasty Mountain and natural boundary 
Ak-Dzhar. Type and paratype in Leningrad. 

57. S. catharinae Juz. in Bot. mat. gerb. Hot. inst. AN SSSR, XIV (1951) 386. 
Perennial, with almost simple vertical woody flexuous root; caudex very short, branch- 
ing; stems numerous, 12-25 cm long, ascending or nearly procumbent, branching all the 
way up, with scattered minute crisp hairs, dark lilac; leaves small, 6-15 mm long, 2.5- 
12 mm wide, ovate or narrowly ovate, generally short-cuneate, tapering at base to slender 
petiole 1-7 mm long, the uppermost subsessile, subobtuse or subacute, incised-crenate- 
dentate with oblong or subtriangular obtuse or subacute teeth, yellow-grayish-green above, 
very finely and sparsely pubescent, glaucescent-gray-green beneath, rather densely tomen- 
tose with very short fine completely appressed hairs, prominently veined; inflorescence 
short, at the onset of flowering ca. 2 cm long, in fruit to 5 cm long, loose; bracts small, to 
7 mm long, 6 mm wide, subherbaceous, ovate to broadly ovate, convex, distinctly keeled, 
short-acuminate, with sparse and fine tomentum of very short appressed crisp hairs, some 
longish hairs especially along margins, and isolated glands, dark lilac; calyx small in flower, 
densely short-hairy, slightly glandular, in fruit equaUng or slightly exceeding bracts; corolla 
1.5-2.5 cm long, abundantly covered outside with •^'^nrt-stipitate glands and hairs, yellow, 
the lower lip and the lateral lobes of upper lip intensely colored, the tube slender, curved; 
nutlets ca. 1 .5 mm long, grayish-white. June. 
146 Sandy steppes, stony and gravelly slopes, pebble-beds. - Centr. Asia: Balkh. (south- 
eastern part). Endemic. Described from Taldy-Kurgan. Type in Leningrad. 



Group 2. Platystegiae Juz. in Bot. zhurn. SSSR, 24, Nos. 5-6 (1939) 433 (pro serie). - 
Bracts usually rather broad, ovate or broadly ovate, slightly concave or nearly flat, not 
navicular (only rarely keeled), at apex angled or shortly and usually abruptly acuminate, 
hairy and stipitate-glandular. 

Note. As mentioned before, we originally gave this group the rank of series. Grossgeim 
in "Izvestiya" Azerb. fil. AN SSSR (3, 1945, 80) made an attempt to classify it as a sub- 
section composed of a number of series; these, however, lack in clear characteristics and 
are completely arbitrary in content. In the present volume we have subdivided the group 
Platystegiae into two series and we have completely rejected the classification proposed 
by Grossgeim. 



Series 1. Eu-platystegiae Juz. — Series Orientales, Oreophilae, Karjaginae, Platyste- 
giae, Artvinenses, Darrienses Grossh. in Izv. Azerb. fil. AN SSSR, 3 (1945) 80-83. - Peren- 
nials; stems not woody or sUghtly woody at base; bracts covered mainly with simple hairs 
and with short-stipitate glands visible only under magnification. 

Note. Besides the regular species of this extensive series, we have tentatively incorpo- 
rated two forms that are apparently hybrids (these were referred to in the Note to the 
series Mesostegiae of the group Oxystegiae). 



98 



58. S. transiliensis Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 397. 
Perennial with vertical woody root; caudex branching, procumbent; stems numerous, 

12-40 cm long, ascending or suberect, curved or slightly flexuous, with short spreading 
crisp hairs, usually not colored or faintly lilac only at base; leaves 0.7-3 cm long, 0.3- 
2 cm wide, triangular-ovate or sometimes oblong, mostly with truncate or broadly cuneate 
base, acute, deeply (to 1/4-1/3 leaf width) incised-dentate with 4-8 irregular oblong 
rounded or often acute teeth on each margin, green, sparsely short-hairy above, gray be- 
neath with very fine appressed tomentum, the midrib and lateral veins prominent; petioles 
slender, 0.2-1.8 cm long, pubescent; inflorescence initially dense, capitate, 3.5-5 cm long, 

147 elongating to 12 cm; bracts almost scarious, to 1-1.8 cm long, 0.6-1 cm wide, ovate to 
broadly ovate, long-acuminate, covered all over with long hairs interspersed with few short 
stipitate glands, usually pale green (uncolored); flowering calyx 3-4 mm long, hairy and 
glandular; corolla large, 2.5-3.7 cm long, hairy and stipitate-glandular outside, yellow, the 
tip of upper lip and lateral lobes often suffused with purple; nutlets ca. 1.5 mm long, 
angled-ovoid, bluish-white with dense pubescence. May— July. (Plate X, Figure 1.) 

Mountain slopes, taluses. - Gentr. Asia: T. Sh. (mainly Zailiiskii Ala-Tau). Endemic. 
Described from vicinity of Alma-Ata. Type in Leningrad. 

59. S. alberti Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 399. 

Perennial, with woody flexuous root; stems several to many, 9-32 cm long, mostly as- 
cending, straight or curved, sometimes flexuous, branching at base, sparsely tomentulose, 
often turning lilac; leaves 0.6-2.2 cm long, 0.5-1 .5 cm wide, rather broadly ovate or ellip- 
tic, truncate or obtusely angled at base, at apex mostly obtuse, rather deeply incised-den- 
tate (to about 1/4 leaf width), with 4-6 large oblong round-tipped teeth on each margin, 
green or usually grayish-green above, with fine scattered tomentum, densely white- or 
grayish-tomentose beneath, the veins not or slightly prominent; calyx 0.2-1 .3 cm long, 
sparsely tomentulose; inflorescence 2.5-5 cm long in flower, later reaching 12 cm, loose; 
bracts almost scarious, to 1 .5 cm long, 1 cm wide, ovate to broadly ovate, attenuate at 
both ends, acute at apex, with somewhat prominent nerves, covered with long hairs inter- 
spersed with few minutely stipitate glands, usually not colored; calyx densely hairy and 
glandular; corolla rather large, 2.2-3 cm long, hairy and short-stipitate-glandular outside, 
plain yellow or sometimes the upper lip with purple lateral lobes and lower lip darker 
(sometimes brown); nutlets ca. 1 .5-2 mm long. May-July. 

Dry pebbly riverbeds, sandy and stony river shoals, stony and gravelly slopes. - Centr. 
Asia: Dzhu.-Tarb. (southern part of Dzhungaria Ala-Tau). — Gen. distr.: Dzhu.-Kash. 
Described from the vicinity of Dzharkent. Type in Leningrad. 

148 60. S. przewalskii Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 400. - 
S. orientalis var. pinnatifida auct. nonnuU. non Rchb. — Ic: Yuz., op. cit. 401, 
Figure 3. 

Perennial; root woody, obUque or vertical, flexuous; stems few, 6-20 cm long, ascend- 
ing or nearly procumbent, curved or flexuous, mostly strongly branching at base, other- 
wise simple, rarely branching above, sparsely tomentulose, often suffused with lilac; leaves 
0.6-2.2 cm long, 0.4-2 cm wide, ovate or eUiptic, truncate or rarely obtuse-angled at base, 
obtuse at apex, very deeply pinnatisect-dentate (width of the middle part of leaf often 



99 



scarcely exceeding the teeth), with 4-7 long digitate teeth 2-8 mm long on each margin, 
the lower leaves usually horizontally directed or reclinate, the upper obliquely ascending, 
dull green or cinereous-green above with sparse fine tomentum and with impressed veins, 
white- or gray-tomentose beneath with fine appressed hairs, the veins not or slightly pro- 
minent; petioles 1 .5-14 mm long, pubescent; inflorescence 2.5-4 cm long, elongating in 
fruit to 7 cm; bracts to 8-15 mm long, 6-10 mm wide, broadly ovate, acute or acuminate, 
long-villous, or without isolated short-stipitate glands, usually turning lilac (especially in 
upper half), with prominent midrib and rather faint lateral nerves; calyx in flower to 4mm 
long, villous-hairy or glandular; corolla 2-4 cm long, hairy and short-stipitate-glandular 
outside, plain yellow or the upper lip and its lateral lobes turning purple; nutlets ca. 
1.75 mm long, angled-ovoid, densely white-hairy. June— October. 

Steppes, dry sandy and clayey-stony exposed slopes, riparian and lacustrine terraces, 
banks and dry riverbeds. — Centr. Asia: T. Sh. (central part). Endemic. Described from 
Lake Issyk-kul' near mouth of Karakolki River. Type in Leningrad. 

61. S. soongorica Juz. sp. nov. in Addenda, XIX, 345. — ? S. orientalis Sievers in 
Pall. Neueste nord. Beitr. Ill (1796) 262, non L. - S. orientalis j3. pinnatifida Ldb. Fl. 
Ross. Ill, 1 (1847-1849) 395, p. p. (non Rchb.) et e. erecta Ldb. 1. c. 

Perennial; root woody, flexuous; caudex short, strongly branching; stems numerous, 
15-30 cm high, ascending or suberect, flexuous, with sparse short crisp hairs, green or lilac; 
leaves small, 0.5-1 .5 cm long, 0.2-0.8 cm wide, narrowly triangular-ovate or oblong, with 
149 truncate or mostly cuneate base, acute at apex, deeply incised-dentate (to about the middle 
of half-blade width) with 4-7 subequal oblong obUquely antrorse apically rounded slightly 
recurved teeth on each margin, grayish-green, diffusely tomentulose above, whitish be- 
neath with very fine appressed tomentum, prominently veined; petioles slender, to 1 cm 
long, tomentose; inflorescence at flrst short, becoming strongly elongated, loose; bracts 
almost scarious, 0.6-1 cm long, 0.4-0.8 cm wide, ovate to narrowly ovate, acuminate, 
strongly convex, rather densely covered (especially along margins) with long spreading 
hairs interspersed with short-stipitate glands, pale green, often suffused with lilac; calyx 
in flower 2-3 mm long, hairy and glandular, in fruit with scutellum ca. 3-4 mm long; 
corolla rather small, 1 .4-1 .7 cm long, pubescent and glandular outside, yellow, usually 
with a violet spot on lower Up; nutlets ca. 1 mm long, ovoid, whitish, with dense soft 
pubescence. June— July. 

Pebble-beds and sands along banks of streams, rarely on loess. — Centr. Asia: Dzu.- 
Tarb. Endemic? Described from Lepsa River. Type in Leningrad. 

Note. For Sievers' plant from Dzhar-Gruban (near sources of Irtysh), provisionally 
placed here in the synonymy, see Note to S. sieversii Bge. 

62. S. krylovii Juz, in Sist. zametk. mat. gerb. Kryl. Tomsk. Gos. univ. 10, No. 8 
(1936) 4; Kryl. Fl. Zap. Sib. IX, 2301. 

Perennial, ca. 25 cm high; stems woody in lower part, procumbent or ascending, 
branching near base, the branches strongly elongated; stems and branches flexuous, 
sparsely and very finely crisp-puberulent and glandular, dark purple to almost blackish- 
purple; leaves 1-2 cm long, 0.8-1 .6 cm wide, rather hard, ovate to mostly broadly ovate, 
slightly tapering at base, obtusely angled, at apex acute, coarsely and unevenly 



100 



incised-dentate or crenate with 4-6 obliquely antrorse oblong straight acute or mostly 
obtuse teeth 1 .5-3.5 mm long on each margin, light green, sparsely and very finely hairy 
above, grayish with very fine appressed tomentum beneath, the midrib prominent, petioles 
3-12 mm long, half as long to nearly as long as blade; inflorescence 3-3.5 cm long in 
flower; bracts broadly ovate, acuminate, with slightly protruding longitudinal nerves, pale 

150 green, with very short hairs along margins and nerves; lower bracts 1.5 cm long and 0.9 cm 
wide; calyx ca. 3 mm long in flower, very finely pubescent and glandular; corolla to 

2.5 cm long, yellow, hairy outside, with short glandular tube. July— August. 

Sands. — West Siberia: Alt. (southern part); Centr. Asia: Balkh. (eastern part), Dzu.- 
Tarb. (Tarbagatai, northwestern slope of Dzhungaria Ala-Tau). Gen. distr.: Dzhu.-Kash. 
Described from Blandy-Kum sands between Nikolaevka and Aleksandrovka (Zaisansk area). 
Type in Leningrad, isotype in Tomsk. 

63. S. sieversii Bge. in Ldb. Fl. Alt. II (1830) 394, sensu stricto; Kryl. Fl. Zap. Sib. IX, 
2300. - S. orientalis 5. adscendens Ldb. Fl. Ross. Ill, 395. — Ic: Ldb. Ic. pi. Fl. Ross. 
II, tab. 123. 

Perennial; rhizome branching, woody; stems numerous, 7-15 (20) cm high, forming 
loose tufts, ascending or suberect, mostly strongly branching at base, simple or weakly 
branching above, covered with soft somewhat crisp spreading hairs, usually greenish or 
lilac in lower part; internodes usually few, rather short; leaves 0.5-2.5 cm long, 0.2- 
1.5 cm wide, ovate to rhombic, obtuse or cuneate at base, mostly subacute, coarsely den- 
tate with 3-7 long subdigitate acute often recurved teeth 1-5 mm long on each margin, 
short-tomentose on both sides, more copiously beneath, grayish-green or the underside 
often canescent, with somewhat prominent veins; petioles as long as or slightly shorter 
than blade, covered with long spreading hairs; flowers short-pediceled; inflorescences 
capitate, loose, 2-6 cm long, slightly elongating after flowering; bracts broadly ovate or 
ovate, tapering at base, densely covered outside with long spreading hairs, finely glandular, 
grayish-green or sometimes sHghtly purple, with slightly protruding midrib and a few late- 
ral nerves, the lower ca. 1 .5 cm long, ca. 0.6 cm wide; calyx ca. 2 mm long, densely hairy 
and glandular; corolla 1 .8-2 cm long, glandular-hairy outside, pale yellow, sometimes the 
lateral lobes of upper Up or the whole upper lip violet and the lower lip with a greenish- 
brown spot; middle lobe of upper Up twice as long and wide as the lateral lobes; lower lip 
orbicular, slightly cleft; corolla-tube strongly elongated, very slightly curved; nutlets ca. 
2 mm long, trigonous-ovoid, densely covered with short grayish hairs. June-July. (Plate X, 
Figure 2.) 

151 Dry stony or gravelly slopes, clayey steppes and semideserts. — West Siberia: Irt.; East 
Siberia: Ang.-Say. (Minusinsk); Centr. Asia: Balkh. Gen. distr.: Mong. (northwestern 
part). Described from rocky places on Arkaul Mountains "Dolenkaraut Tschingistau." 
Type in Leningrad. 

Notes. 1) The original plant of Sievers, to whom the species is dedicated, was included 
here by Bunge, but it differs from Meier's plant which was actually described by Bunge 
and, in our opinion, does not belong to S. sieversii Bge. s. str. At least the Sievers speci- 
men preserved in the herbarium of the Botanical Institute of the Academy of Sciences of 
the USSR would be more appropriately referred to S. soongorica Juz. 

2) It is of considerable interest that forms occur in nature which are very similar to 



101 



S. sieversii Bge., but have a purple corolla. We were not inclined to refer such forms to 
S. sieversii and we have therefore omitted any reference to such coloring in describing 
the flowers of this species. In the herbarium, we have named one of these forms S. ulu- 
tavica Juz. (from its habitat in the Ulu-Tau Mountains), but the few specimens are old 
and in a poor state of preservation and we have therefore refrained for the time being from 
offering a description. We assume that the existence of such forms is evidence of hybridi- 
zation between species of the series Pulchellae and Platystegiae that has given rise to such 
species as the above-described S. turgaica and S. karkaralensis, or S. ulutavica. 

64. S. hypopolia Juz. in Bot. zhurn. SSSR, 24, Nos. 5-6 (1939) 433. - S. orientalis 
a. chamaedryfolia Ldb.Fl.Ross. Ill, 395, p.p.; N. Pop. in Mat. fl. Kavk. IV, 3, 73, 
p. p. non Rchb. 

Perennial, plant 6-25 cm high; stems numerous, ascending at base or nearly procum- 
bent, curved or flexuous, tomentulose in lower part, densely covered above, especially in 
inflorescence, with short spreading hairs; leaves 5-1 7 mm long, 3.5-1 2 mm wide, ovate, 
usually obtusely angled or sometimes cuneate at base, much more rarely truncate, irregu- 
larly and deeply incised-crenate-dentate with 5-7 obtuse to subacute often revolute 
medium-sized teeth (1-2.5 mm long) on each margin, green and finely puberulent above, 
very densely white-tomentose beneath, the often prominent veins completely hidden by 
tomentum; petioles 1.5-10 mm long; inflorescences 2.5-4 cm long, dense, in fruit up to 
10 (?)cm long, rather lax; bracts broadly ovate, concave, subacuminate to acuminate, the 
lowermost ca. 10 mm long, 7-9 mm wide, covered with dense or very dense rather long 
152 and soft hairs interspersed with numerous stipitate glands, greenish or later slightly red- 
dening, with prominent nerves; calyx hairy and glandular; corolla 1 .5-2.5 cm long, yel- 
low, hairy and glandular outside. April-June. 

Stony slopes, limestone and chalk outcrops, steppes. — European part: Crim. (foot- 
hills). Endemic. Described from Karasubazar, Kara-Kush. Type in Leningrad. 

65. S. heterochroa Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 402. - S. 
orientalis Schmalh. Fl. sr. i yuzhn. Ross. II, 326 et auct. fl. taur. p. p. non L. — S. orien- 
talis var. chamaedryfolia N. Pop. in Mat. fl. Kavk. IV, 3 (1916) 73, p. p. non Rchb. - 
S. orientalis var. t auric a id. ibid. (1916) 81, p.p. 

Perennial, 16-26 cm high, very similar to S. hypopolia Juz.; distinguished primarily 
by the shorter, often suborbicular leaves, these usually not more than 1 .5 cm (rarely to 
2 cm) long, to 1 .5 cm wide, usually truncate at base, the teeth more obtuse and slightly 
divaricate, inflorescences more floriferous, finally much elongated, reaching 15 cm in fruit; 
bracts somewhat smaller (the lowermost approximately to 6-8 mm long and to 8-9 mm 
wide), obtuse to short-acuminate, usually turning lilac, flowers pure yellow, purple or 
purple-yellow (wdth purple Ups and upper part of flower and yellow tube). June-July. 

Stony places. — European part: Crim. (mainly eastern part). Endemic. Described 
from vicinity of Karadag biological station. Type in Leningrad. 

66. S. hirtella Juz. in Bot. zhurn. SSSR, 24, Nos. 5-6 (1939) 434. - S. orientalis var. 
chamaedryfolia N. Pop. in Mat. fl. Kavk. IV, 3(1916) 73, p. p. non Rchb. 

Perennial, 5-20 cm high; leafy stems numerous, ascending at base, curved, very densely 



102 



covered, like petioles, with spreading or reclinate soft crisp hairs; internodes of medium 
length or short; leaves 0.5-2 cm long, 0.45-1 .9 cm wide, broadly ovate to suborbicular, 
usually truncate at base, coarsely and deeply (to 1-4 mm) incised-crenate with 3-6 equal, 
obtuse teeth on each margin, grayish, densely hairy, rugose, with impressed veins above, 
very densely white- or usually gray-tomentose to almost villous, with very distinct veins 
beneath; petioles 0.5-1.5 cm long; inflorescences 2.5-4 cm long, in fruit to 6 cm, dense 

153 or very dense; bracts closely imbricated, broadly ovate, convex or nearly flat, acuminate 
greenish, covered with simple long hairs and copious stipitate glands, prominently nerved, 
the lower ca. 1cm long, 0.8 cm wide; flowering calyx ca. 2.5 mm long; corolla medium to 
fairly large, 2-3 cm long, yellow, densely covered outside with glandular and simple hairs. 
May— June. 

Stony mountain slopes, rocks. — European part: Crim. (Crimean State Reserve). En- 
demic. Described from rocks near the sources of Sary Su River. Type in Leningrad. 

67. S. novorossica Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (195 1) 403. - S. 
hirtella Grossh. in Izv. Azerb. fil. AN SSSR, 3 (1945) 80, non Juz. 

Subshrubs, resembling S. hirtella Juz. in habit and in vesture of stems and leaves, but 
leaves (especially the upper) narrower, often ovate, obtuse-angled at base, deeply incised- 
dentate with mostly acutish teeth; bracts narrower, frequently ovate, more distinctly acu- 
minate, densely covered with moderately long spreading simple hairs interspersed with 
rather few substipitate glands. May— June. 

Mountain slopes. — Caucasus: W. Transc. (Novorossiisk district). Described from the 
vicinity of Novorossiisk. Type in Leningrad. 

Note. The species is known only from very scanty material and needs further study. 

68. S. polyodon Juz. in Bot. zhurn. SSSR, 21 , Nos. 5-6 (1939) 433, in adnot. - S. 
orientalis Schmalh. Fl. II (1897) 326 et auct. fl. Cauc. p.p. non L. — S. orientalis a. 
chamaedry folia Ldb. Fl. Ross. Ill (1847-1849) 395, p.p.; N. Pop. in Mat. fl. Kavk. IV, 
3, 73, p.p. non Rchb. -? S. caucasica A. Hamilt. Monogr. (1832) 282 (nomen prius!). - 
Ic: Grossg. in Izv. Azerb. fil. AN SSSR, 3 ( 1945) 85, Figure 6 - Exs: GRF, Number 524. 

Subshrub; caudex very short, sturdy, usually horizontal, rarely erect, nodose; stems 
mostly numerous, ascending or often erect, robust, slightly curved, 10-32 cm long, sparse- 
ly covered with short divergent or divaricate curved hairs; internodes as long as or shorter 
than leaves; leaves medium to fairly large, 0.8-3 cm long, 0.3-1 .5 cm wide, oblong-ovate 
or narrowly triangular, with subcordate, truncate or (the upper) obtuse-angled base, ob- 
tuse or (especially the upper) acute at apex, rather deeply incised-crenate-dentate or the 

154 upper usually acutely incised-dentate, with 5-13 obliquely antrorse teeth to 4 mm long 
on each margin, green, sparsely covered with short fine appressed hairs above, densely 
white-tomentose with fine appressed hairs beneath, the lateral veins not or slightly pro- 
truding, often hidden by tomentum; petioles sturdy, suberect, short-tomentose, 0.2- 
2 cm long, distinct even in upper leaves, these tightly enveloping the lower part of inflo- 
rescence; inflorescence 3.5-6 cm long, in fruit to 8 cm, elongate-ovoid or conical, dense; 
bracts large, to 2 cm long, 1 .5 cm wide, of rather firm consistency, ovate, rarely broad- 
ovate, long-acuminate, entire, pale green or sometimes bright lilac, diffusely covered with 
short appressed hairs and with scattered punctate or short-stipitate glands, the margins 



103 



with spreading short hairs, the interspaces between the prominent longitudinal nerves 
indistinctly reticulate; flowering calyx ca. 3 mm long, patent-hairy and densely stipi- 
tate-glandular; corolla large, to 3-3.5 cm long, to 8 mm wide at throat, yellow, covered 
outside with short spreading hairs and stipitate glands, the lips approximately equal in 
length. May -August. (Plate IX, Figure 1.) 

Steppe slopes, limestone, hmestone and rocky slopes, rocks, gravels, taluses. — 
Caucasus: Cisc. (foothills of Main Range). Endemic. Described from the vicinity of 
Kislovodsk. Type in Leningrad. 

69. S. oreophila Grossh. in Izv. Azerb. fil. AN SSSR, 3 (1945) 80. - S. orientalis var. 
chamaedryfolia N. Pop. in Mat. fl. Kavk. IV, 3 (1916) 73, p. p. non Rchb. - Ic: Grossg., 
op. cit. 85, Figure 8, 26, b. 

Perennial; caudex procumbent, flexuous, rooting at nodes; stems few, 10-30 cm long, 
usually ascending, rather robust, curved or slightly flexuous, simple or (mostly slightly) 
branching, sparsely covered with short fine crisp hairs or subglabrous; leaves medium to 
large, 1-4 cm long, 0.7-2.3 cm vdde, half as long (often the middle and upper) to about as 
long as internodes, broad-ovate, ovate or oblong-ovate, with subcordate or truncate or the 
upper sometimes with obtusely angled base, mostly obtuse at apex, shallowly incised- 
crenate with 3-8 mostly oblique semielliptically rounded crenations on each margin, green 
with scattered fine short hairs, densely covered beneath with fme white or grayish tomen- 
155 tum, the veins not prominent; petioles to 2 cm long, densely tomentulose; inflorescence 
3-6.5 cm long, dense, in fruit often loose, 9 cm long; bracts usually large, (1) 1 .5-2 cm 
long, 9-12 mm wide, scarious, ovate or broad-ovate, convex or almost flat, acute or long- 
acuminate, entire, densely covered on the back with long spreading soft hairs copiously 
interspersed with short stipitate glands, often suffused with lilac; corolla very large, 3- 
4 cm long, 6-9 mm wide at throat, yellow, rather sparsely covered outside with fme hairs 
and glands. June— September. 

Alpine pastureland, meadows, mountain-steppe slopes, rocks and taluses in alpine and 
subalpine mountain belts. — Caucasus: Cisc, Dag., E. Transc. (Greater Caucasus). En- 
demic. Described from Kryz in Kuba district. Type in Baku. 

Note. Since the distribution areas of S. oreophila Grossh. and S. polyodon Juz. are 
contiguous, one might expect that intermediate forms between these species would be 
found in nature. But so far we have not found in the herbarium any obvious transitional 
forms between these species, except for a single specimen from Mount Oshten. Consider- 
ing the isolated nature of the location, we can assume that this is a special race, a brief 
description of which we tentatively present below. 

70. S. oschtenica Juz. sp. nov. in Addenda XIX, 346. 

A perennial plant of an apparently intermediate character, with the overall habit of 
S. oreophila Grossh., but with almost completely glabrous bracts which, in shape, are 
more like those of S. polyodon Juz.; inflorescence very short, few-flowered, nearly 
globular, not changing in fruit. July. 

Grassy slopes, taluses. — Caucasus: Cisc. (Oshten Mountain). Endemic. Described 
from slopes of Mount Oshten. Type in Leningrad. 



104 



71. S. daghestanica Grossh. in Izv. Azerb. fil. AN SSSR, 3 (1945) 80, sensu stricto. - 
S. orientalis var. chamaedry folia (non Rchb.) cum f. elongata N. Pop. in Mat. fl. 
Kavk. IV, 3 (1916) 73 et 79, p. p. - Ic: ? Grossg., op. cit. 85, Figures 9, 26, a. 

Subshrub 15-40 cm high; caudex short, branching; stems few, ascending or suberect, 
elongate, more or less virgate, simple or slightly branching, stout, flexuous, diffusely white- 
tomentulose with fine-short-white-spreading hairs, usually reddening in lower part, remote- 
ly leafy; leaves usually 1/4 the length of internodes, 0.8-2 cm long, 0.4-1 .2cm wide, ovate 
or oblong, obtuse, deeply incised-crenate with 5-7 semieUiptic or oblong obtuse crenations 

156 on each margin, grayish-green, sparsely covered with fine spreading hairs above, densely 
white-tomentose beneath, the midrib and lateral veins usually very prominent; petioles 
1.2 cm long; inflorescence dense, 2.5-5 cm long, becoming loose in fruit, to 7 cm long; 
bracts scarious but thickish, 7-12 nmi long, 6-9 mm wide, broadly ovate to suborbicular, 
obtuse to short-acuminate, entire, convex, with prominent longitudinal nerves, densely 
covered on the back, especially on the margins and nerves, with long spreading hairs and 
short-stipitate glands, pale green, often dull purple in upper half; corolla rather small, ca. 
1 .5-2.2cm long, ca. 4mm wide at throat, yellow, densely hairy and glandular outside. July. 

Central mountain belt, rocks. — Caucasus: Dag. Endemic. Described from Ikhrek 
(Radde collections). Type in Leningrad. 

Note. We are interpreting this species in the narrow sense, and we base our approach 
on an investigation of the "type" selected by Grossgeim himself. However, Grossgeim con- 
fused it with some other plants, in particular with S. granulosa Juz. and low-growdng, 
high-mountain Dagestan forms of S. oreophila Grossh. In other words, he referred all 
"S. orientalis" of Dagestan derivation to S. daghestanica Grossh., and this, in our 
opinion, is quite incorrect. 

72. S.granulosa Juz. nom.nov. — S. orientalis var. chamaedry folia f. elongata N. 
Pop. in Mat. Fl. Kavk. IV, 3 (1916) 79, sensu stricto. — S. boissieri Sosn. in scheb. Herb. 
Inst. Bot. Ac. Sc. URSS (1929) nee aUbi; Grossg. Fl. Kavk. Ill (1932) 286, p. p. nee alibi; 
emend. Juz. in Bot. zhurn. SSSR, 24, Nos. 5-6 (1939) 431 (nomen ambiguum!). - S. da- 
ghestanica Grossh. in Izv. Azerb. fil. AN SSSR, 3 (1945) 80, p.p. 

Subshrub, 20-45 cm high; caudex ascending or erect, slender, woody, branching; an- 
notinous stems elongate, slender, more or less virgate, straight or curved, simple or branch- 
ing, with fine curved hairs, usually lilac, remotely leafy; leaves generally small, 1/2 to 1/3 
the length of internodes, (0.7) 1.2-2 cm long, (2) 5-10 mm wide, the lowermost broad- 
ovate to ovate, others oblong-ovate, acute, shallowly incised-crenate with (3) 5-9 obtuse 
or rounded smallish crenations on each margin, diffusely short-hairy above, finely white- 
tomentose beneath, the veins not prominent; petioles to 1 cm long, tomentulose; inflo- 
rescence short or oblong, 2.5-6 cm long, dense, becoming loose in fruit; bracts scarious, 

157 8-10 mm long, ca. 7 mm wide, convex, broadly ovate-rhombic, acuminate, entire, the 
lower with short appressed hairs, others with predominantly long spreading hairs inter- 
spersed with sessile and short-stipitate glands, with prominent longitudinal nerves and 
often distinctly carinate; corolla rather small, ca. 2 cm long, 4 mm wide at throat, yellow, 
covered outside with spreading hairs and short-stipitate glands. June— July. 

Central mountain belt, gravelly places. — Caucasus: Dag. Endemic. Described from 
Gimri and Gunib. Type and paratype in Leningrad. 



105 



Note. A distinctive and characteristic species known only from certain parts of Dage- 
stan (Gunib district: between Gunib and Chokli, between Gunib and Khunzakh; Avarskii 
Dagestan, soutli of Gimri along Koi-Su). Althougli we have included S. granulosa in the 
Platystegiae, this species clearly gravitates toward Oxystegiae in some characters (bracts 
with two types of vesture and a strongly developed keel). It may possibly be a hybrid 
which, on the strength of its derivation, constitutes a link between Platystegiae, to which 
we have referred it, and Oxystegiae. Grossgeim, in arbitrarily attempting to apply the 
name S. boissieri to our S. raddeana (see note to S. raddeana), clearly confounded 
the genuine plant with other forms, notably with his S. daghestanica s. str. 

73. S. orientalis L. Sp. pi. (1753) 598; Ldb. Fl. Ross. Ill, 395. - S. orientalis |3. cha- 
maedry folia Rchb. Ic. bot. pi. crit. I (1823) 10; Ldb. Fl. Ross. Ill, 395, p.p.; N. Pop., 
in Mat. fl. Kavk. IV, 3, 73, p.p. - S. orientalis var. genuina Boiss. Fl. or. IV (1879) 
682, p.p. — S. orientalis var. pinnatifida auct. p.p.; N. Pop., op. cit. 79, p.p. non 
Rchb. - S. orientalis var. chamaedryfolia f. elongata N. Pop. op. cit. 79, pro min. 

p. - ? S. bractealis Stev. ined. (ex sched. Herb. Gay). — ? S. caucasica A. Hamilt. 
Monogr. (1832) 282. - Ic: Tournef. Voy. II, tab. ad pag. 306; Rchb. 1. c, tab. VIII, 
fig. 17. - Exs.: Fl. Cauc. exs. No. 216. 

Subshrub; caudex mostly short, procumbent to ascending, woody, usually flexuous 
and strongly branching, leafy stems fairly numerous, ascending at base, curved or flexuous, 
rarely suberect, 7-25 cm long, densely covered with short spreading slightly crisp hairs; 
internodes mostly short, not exceeding leaves; leaves 0.4-2 cm long, 0.4-1 .5 cm wide, 
rounded-triangular, broad-ovate to ovate, truncate, subcordate or obtusely angled at base, 
158 usually obtuse or rounded at apex, deeply (sometimes more than half the half-blade width) 
incised-crenate or dentate with 3-7 semielliptic or usually oblong to sublinear teeth 1.5- 
3 mm long on each margin, dark green or gray-green, covered above with short fine appres- 
sed hairs, white-tomentose with fine oppressed hairs beneath, the veins not prominent or 
sometimes prominent, mostly concealed by tomentum, without distinct network of vein- 
lets; petioles 0.4-1 cm long, patent-pilulose; inflorescence 2.5-6 cm long, dense or fairly 
lax; bracts small or more often rather large, the lower 0.8-2 cm long, 0.6-1 .4 cm wide, 
scarious, ovate or often broad-ovate, subacute or subacuminate, entire or sometimes with 
solitary teeth, sparsely covered with short appressed hairs, finely punctate-glandular, most- 
ly pale green, rarely suffused with reddish-purple, with prominent longitudinal nerves, very 
obscurely reticulate; calyx densely covered with short spreading hairs, stipitate-glandular; 
scutellum to 4mm long in fruit, sparsely short-hairy; corolla medium to large, 2.5-3 cm 
long, 6-8 mm wide at throat, yellow, short-hairy and stipitate-glandular outside; nutlets 
gray with dense appressed pubescence. May— July. 

Dry grassy places and pastures, exposed stony and calcareous slopes, clayey bluffs, 
gravels. — Caucasus: E. Transc. Endemic. Described from Tbilisi. Type in Paris (or 
London?). 

Note. This species varies greatly in the degree of dissection of the leaf blade, and for 
tliis reason we have included it twice in the key. 

74. S. sedelmeyerae Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (195 1) 403. - S. 
orientalis var. chamaedryfolia N.Pop.in Mat. tl.Kavk.IV,3(1916) 73,p.p. non Rchb. 



106 



(159) 'i; 




PLATE IX. 1 - Scutellaria polyodon Juz., general aspect, leaf, bract, flower, anther; 2 - S. platy- 
stegia Juz., general aspect, leaf, bract, flower; 3 - S. karjaginii Grossh., summit of stem, leaf, bract, 
flower; 4 - S. sosno vskyi Takht., general aspect, leaf, bract, flower, anther, fragment of fruiting 
inflorescence, nutlets. 



107 



Perennial, 10-20 cm high; caudex very short, nodose; stems several to many, ascending, 
usually curved or slightly flexuous with very short spreading hairs, sometimes slightly lilac- 
tinged; leaves 0.8-2 cm long, 0.6-1.2 cm wide, ovate-triangular or oblong-ovate, usually 
truncate at base, acute, pectinately incised (to about half the half-blade width) with 5-7 
large remote obtuse or rounded slightly antrorse teeth (to 4 mm long) on each margin, 

161 green above, sparsely or densely covered with fine curved hairs, grayish beneath with dense 
short imperfect tomentum, with distinct slightly prominent yellowish veins; petioles to 
1.3 cm long, as long as or usually shorter than blade, covered with short curved hairs; in- 
florescence 3-7 cm long, at first rather dense, later fairly lax; bracts 1-1.5 cm long, 0.8- 

1 cm wide (the lower), rhombic, broadly ovate, tapering at both ends, subacute or short- 
acuminate, densely covered with short or rather long spreading hairs and scattered stipitate 
glands, pale green or mostly turning dark lilac at least in upper part, the lowermost with 
few large teeth (to 4 on each margin); calyx densely covered with long spreading white 
hairs and short-stipitate glands; corolla large, 3-3.7 cm long, 6 mm wide at throat, yellow, 
with a large brown spot on lower lip, patent-hairy and finely stipitate-glandular. June-July. 
Dry mountain slopes. - Caucasus: S. Transc. Endemic. Described from the vicinity of 
Elenovka. Type in Leningrad. 

75. S. platystegia Juz. in Bot. zhurn. SSSR, XXiV, Nos. 5-6 (1939) 431 (in adnot.). - 
S. orientalis a. chamaedryfolia Ldb. Fl. Ross. Ill (1847-1849) 395, p. p.; N. Pop. in 
Mat. 11. Kavk. IV, 3, 73, p. p. non Rchb. - Ic: Grossg. in Izv. Azerb. fil. AN SSSR, 3 
(1945), 85, Figure 11. 

Subshrub 12-25 cm high, with a short woody branching caudex; stems many, ascend- 
ing at base, curved or almost straight; leafy shoots and petioles rather densely covered 
with short spreading slightly crisp hairs; internodes usually elongate; leaves medium to 
large, 7-23 mm long, 5-20 mm wide, triangular or broadly ovate, with truncate or subcor- 
date base, usually acuminate, rather deeply incised-crenate or dentate with 4-7 long (ca. 
3 mm) very uneven sliglitly spreading usually obtuse teeth on each margin, dark g ■> en, 
rather sparsely appressed-hairy above, softly white- or gray-tomentose beneath, with nerves 
not usually concealed by tomentum, more or less distinctly netted-veined, the lowermost 
with petioles to 1.2 cm long, the upper short-petioled to subsessile; inflorescence 2.5- 
5 cm long, dense; bracts broadly ovate, hardly as long as wide, the lower ca. 1 .5 cm long, 
1.2 cm wide, convex, short-attenuate to subacute at apex, the lowermost few-toothed, the 

162 upper entire, sparsely covered with very fine short curved hairs, punctate-glandular, spar- 
ingly long-hairy on the margins, greenish or mostly dark purple, netted-nerved; calyx 
rather densely short-hairy and glandular; corolla medium or fairly large, to 3 cm long, 
yellow, covered outside with long hairs and short-stipitate glands. April-July. (Plate IX, 
Figure 2.) 

Stony slopes, steppes. — Caucasus: S. Transc. Endemic. Described from Karabakh, 
between Chai Tumos and Kyagrizami. Type in Leningrad. 

Note. Akin to S. orientalis; slightly differing in habit and chiefly distinguished by 
vesture of leaf undersurface. 

76. S. grossheimiana Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 405. - 
S. prilipkoana Grossh. in sched. p. p. (nee alibi). — S. orientalis var. chamaedryfolia 
etvar. pinnatifida N. Pop. in Mat. fl. Kavk. IV,3 (1916) 73 et 79. p.p. non Rchb. 



108 



Perennial, 6-18 cm high; stems densely covered with long spreading hairs; internodes 
generally shorter than in S. platystegia Juz., not exceeding leaves; leaves 0.5-2 cm long, 
0.4-1.5 cm wide, orbicular or broadly triangular-ovate, obtuse or subobtuse, very deeply 
(more than half the half-blade width) incised-crenate with 4-7 oblong or broadly linear 
teeth on each margin, gray-green, densely appressed-hairy above; petioles to 1 .2 cm long; 
inflorescence 2.5-5 cm long, in fruit to 7 cm; bracts more compacted than in S. platy- 
stegia Juz. pale grayish-green, sometimes slightly purple, more densely covered with slight- 
ly longer curved hairs, the inconspicuous minutely stipitate or sessile glands almost con- 
cealed; lower bracts 1-1.5 cm long, 1-1.4 cm wide; corolla 2.5-3 cm long, ca. 5 mm wide 
at throat. Otherwise, similar to S. platystegia Juz. May— June. 

Dry, mostly stony slopes. — Caucasus: Tal. Endemic. Described from near Lerik. 
Type in Leningrad. 

77. S. artvinensis Grossh. in Izv. Azerb. fil. AN SSSR, 10 (1944) 37. - S. orientalis 
var. pinnatifida N. Pop. in Mat. fl. Kavk. IV, 3 (1916) 79, p.p. non Rchb. — Ic: Grossg., 
op. cit. 3 (1945) 85, Figure 16. 

Subshrub 12-25 cm high, caudex long-procumbent, strongly branching, often flexuous; 
stems usually ascending, curved or slightly flexuous; internodes mostly short; leaves 0.6- 
2.4 cm long, 0.5-1 .8 cm wide, broadly ovate to suborbicular, obtuse, very deeply (to 1/3- 
2/5 the width of blade) incised-dentate, with 3-8 triangular or digitate often slightly re- 
163 curved obtuse teeth to 5 mm long on each margin, grayish-green, densely appressed-hairy 
above, white-tomentose beneath, the midrib and lateral veins prominent, not concealed by 
tomentum; petioles to 1-1.5 cm long; inflorescence 2.5-5 cm long, loose; bracts not com- 
pacted, mostly entire, densely patent-hairy and stipitate-glandular, pale green or often purple- 
tinged, with prominent longitudinal nerves, the lower to 1 .3 cm long, 1 cm wide; corolla 
2.5-3.5 cm long, rather densely hairy and stipitate-glandular outside. Otherwise, like S. 
platystegia Juz. which it resembles particularly in the vesture of stems and of leaf under- 
side. April-June, again in August. 

Taluses, stony bluffs and slopes. — Caucasus: S. Transc? Gen. distr.: Arm.-Kurd. 
Described from Artvin district. Type in Baku (or Tbilisi?). 

78. S. k^rjaginii Grossh. in Izv. Azerb. fil. AN SSSR, 3 (1945) 81. - S. improvisa 
Grossh. Opred. rast. Kavk. (1949) 327. - S. rhomboidalis Grossh. in Bot. mat. gerb. 
Bot. inst. AN SSSR, XIII (1950) 19, p. p. - Ic: Grossg., op. cit. (1945) 85, Figure 10. 

Subshrub, 8-30 cm high; caudex short, strongly branching; stems many, ascending, 
erect or shghtly curved, simple or somewhat branching, sparsely or rather densely covered 
with short curved hairs; leaves 0.7-2.5 cm long, 0.5-2 cm wide, triangular-ovate, truncate 
or obtusely angled at base, obtuse to subacute at apex, coarsely incised-dentate with 5-7 
obliquely triangular obtuse teeth on each margin (the teeth not exceeding 1/3, rarely 
reaching 1/2 of half-blade width), green above, slightly rugose, finely appressed-hairy, 
densely covered beneath with white or gray appressed tomentum, the midrib and lateral 
veins often prominent; petioles tomentose-hairy, to 1.5 cm long; inflorescence 3-4cm long, 
in fruit to 10cm, rather dense; bractsmedium to fairly large, 0.8- 1.4 cm long, 0.5-1.1 cm 
wide, rhombic-ovate, with 2-4 triangular large teeth on each margin or the upper entire, 
diffusely covered with short hairs, with rather long spreading hairs on the margins, slightly 



109 



stipitate-glandular, mostly pale green, sometimes turning lilac; corolla 2.5-3 cm long, 
4-5 mm wide, short-hairy and stipitate-glandular outside, bright yellow, the lower Up 
darker (brownish); nutlets ca. 1.5 mm long, angled, densely covered with appressed white 
hairs. April-June. (Plate IX, Figure 3.) 

Dry stony and rocky slopes, pebbles in the middle and lower mountain belts. - Cauca- 

164 sus: S.Transc. Gen.distr.: Iran (?). Described from Aznabyurt (Nakhichevan ASSR, 
Nakhichevan district). Type in Baku, paratypes and topotypes in Leningrad. 

Note. The synonymy cited indicates how vague the author himself was concerning 
this species. It should also be noted that Grossgeim referred the more mesophytic speci- 
mens of S. karjaginii to S. sevanensis Sosn. (in herbarium of the Botanical Institute 
of the Academy of Sciences of the USSR). 

79. S. rhomboidalis Grossh. Opred. rast. Kavk. (1949) 328 (rossice); Grossg. in Bot. 
mat. gerb. Bot. inst. ANSSSR, XIII (1950) 19, sensu restricto. 

Perennial, 12-22 cm high, similar on the whole to S. karjaginii but leaves (especially 
the upper) mostly elongate; bracts smaller, 6-1 1 mm long, 3-8 mm wide, acute or acumi- 
nate, entire or the lower often with few small teeth on the margin, green, diffusely tomen- 
tulose, the tomentum intermixed with few short straight hairs, these more abundant on 
the margins; inflorescence 3-8 cm long (apparently elongating after flowering); corolla 
2-2.5 cm long, with slender tube. May-July. 

Pebble beds. - Caucasus: S. Transc. (Nakliichevan ASSR). Endemic. Described from 
Norashen district, estuary of Kabakhly-Chai River near Dize. Type in Leningrad. 

Note. This is apparently an "interserial" hybrid, intermediate between S. karjaginii 
Grossh. and S. araxensis Grossh. We accepted as its type one of Grossgeim 's "isotypes" 
(i.e. dupUcates of type) because it appeared to us that the specimen he recorded as the 
type was none otlier than S. karjaginii Grossh. which occurs in the location indicated 
together with S. rhomboidalis. As far as S. araxensis Grossh. is concerned, it was not 
collected here and, generally speaking, S. rhomboidalis appears to be a completely fixed 
form (we refer to it specimens preserved in the Botanical Institute of the Academy of 
Sciences of the USSR that were collected by T. Heideman from an entirely different 
locahty - near the village of Bichenakhj. 

80. S. prilipkoana Grossh. in Izv. Azerb. fil. AN SSSR, 3 (1945) 82. - S. orientalis 
var. pinnatifida N. Pop. in Mat. fl. Kavk. IV, 3 (1916) 79, p.p. non Rchb. - Ic: Grossg., 
op. cit. 85, Figure 13. 

Perennial, 8-25 cm high; caudex procumbent, flexuous, branching; stems many, as- 
cending, simple or branching, curved or flexuous, densely covered (like petioles) with long 
spreading crisp hairs, often turning lilac; leaves 0.5-1 .5 (2) cm long, 0.4-1 .4 cm wide, ovate 

165 to broad-ovate, truncate or arched-incurved at base, obtuse to subacute, deeply (to 1/3- 
2/5 width of blade) pinnatifid, with 4-6 nearly horizontally spreading or commonly 
obliquely antrorse, oblong or linear, obtuse teeth or lobules to 6 mm long at each side, 
grayish-green, densely covered with fine crisp hairs above, grayish with dense soft spreading 
tomentum beneath, the veins distinct but not prominent; inflorescence 3-4 cm long, lax 
or sometimes initially dense; bracts 1-1.5 cm long, 0.7-1 cm wide, ovate to broad-ovate, 
short-acuminate, entire, the lower often with a few teeth, rather densely soft-hairy, slightly 



110 



glandular, pale green or sometimes lilac; corolla 2.4-3 cm long, 4-5 mm wide at throat, 
bright yellow, soft-hairy and glandular outside. May— August. 

Dry stony and rocky slopes. - Caucasus: Tal. Endemic. Described from Kosmodal'- 
yan in Zuvand. Type in Baku. 

Note. The species is close to the Iranian S. mucida Stapf, but differs in having mostly 
entire bracts. 

81. S. darriensis Grossh. in Izv. Azerb. fil. AN SSSR, 3 (1945) 83. - Ic: Grossg., op. 
cit. 85, Figure 17. 

Perennial, 15-30 cm high; stems many, ascending; leaves 1-2 cm long, ovate, cut near- 
ly to base into 4-5 Unear obtuse globes ca. 2 mm wide, green, finely pubescent above, gray, 
finely crisp-hairy beneath; bracts 1-1.5 cm long, 5-8 mm wide, ovate-rhombic, acute, with 
2-3 rather large triangular acute teeth on each margin, rather densely puberulent; corolla 
2.5-3 cm long, 4-5 mm wide at throat, yellow. July (?). 

Rocky places in the lower mountain belt. — Caucasus: S. Transc. Gen. distr.: N. Iran? 
Described from Darri-Dagh Mountains. Type in Baku. 

Note. We have not seen specimens of this species and we have borrowed its very brief 
description from the work of Grossgeim. 

82. S. sosnovskyi Takht. in Zam. po sist. i geogr. rast. izd. Gruz. fil. AN SSSR, 9(1940) 
24. — S. orientalis a. pinnatifida Rchb. Ic. pi. crit. I (1823) 10 (non S. pinnatifida 
A. Hamilt.); N. Pop. in Mat. fl. Kavk. IV, 3, 79, p.p. - Ic: Rchb. 1. c. tab. VIII, fig. 16; 
Grossg. in Izv. Azerb. fil. AN SSSR, 3 (1945) 85, Figure 14. 

Perennial, low-growing, producing small tufts; caudex procumbent, woody, branching; 
flowering and sterile shoots numerous, ascending, leafy, 5-15 cm long, covered with short 
spreading crisp hairs; leaves 5-25 mm long, 2.5-12 mm wide, elliptic or oblong, usually 
166 truncate at base, deeply (2/5-3/7 the width of blade, sometimes nearly to midrib) pinnati- 
sect into 4-7 horizontal or slightly antrorse linear or narrowly Unear obtuse teeth or seg- 
ments on each margin, sometimes with one long tooth, green, usually with scattered short 
hairs above, grayish or whitish beneath with copious short appressed or spreading tomen- 
tum, the veins prominent or often indistinct; petioles usually long, slender, patent-tomen- 
tose, 0.3-2 cm long; inflorescence loosely spicate, 2.5-5 cm long; bracts 8-13 mm long, 
3-6 mm wide, narrowly ovate or ovate-lanceolate, tapering to base and sessile, obtuse or 
acute, the lower often with 1-2 teeth per margin, rather densely covered with long spread- 
ing hairs and scattered short-stipitate glands; calyx densely patent-hairy and glandular; 
corolla 2.5-3 cm long, ca. 7 mm wide in throat, yellow, hairy and stipitate-glandular out- 
side. August. (Plate IX, Figure 4.) 

Steppes and steppe slopes, stony mountain meadows, rocks, taluses. — Caucasus: E. 
Transc. (Borzhomi and Akhaltsikhe), S. Transc. (western part of Armenian SSR). Gen. 
distr.: Arm.-Kurd. Described from Amass district, Kaikuli. Type in Erevan. 

83. S. amphichlora Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 411. 

Subshrub, 5-22 cm high; caudex low, woody, branching; stems leafy, ascending, spar- 
ingly branching, slightly flexuous, pale, woody in lower part, sparsely covered with short 
curved hairs interspersed with short-stipitate glands; leaves 0.5-2 cm long, 0.5-1 cm wide, 



111 



ovate, with truncate or obtusely angled base, obtuse at apex, pale green on both sides, 
sparsely or beneath more densely short-hairy with intermixed (sometimes abundant) short- 
stipitate glands, tire pinnatiseet-dentate margins with obliquely antrorse teeth or segments, 
these luilf of half-blade width or usually slightly longer, oblong or broadly linear, usually 
rounded at the ends; petioles patent-hairy and stipitate-glandular, 0.2-1.5 cm long; intlo- 
rescence 2.5-5 cm long; bracts 0.6-1 .4cm long, 0.3-0.8 Cm wide, ovate to ovate-lanceolate, 
obtuse or sometimes acuminate, entire or the lowermost with few acute teeth, sparsely 
covered with long hairs and stipitate glands, pale green or slightly colored; corolla 2-3 cm 
long, 4-6 mm wide at throat, yellow, often with a brown spot at the center of lower lip, 
covered outside with short hairs and stipitate glands. July. 
167 Dry mountain slopes and pastures (2400-2800 m). Caucasus: S. Transc. (Aragats 
Mountain). Described from Aragats Mountain, Dali-Chai River. Type in Leningrad. 

Note. On establishing this species we submitted the notion that it derived from a 
crossing of S. sosnowskyi Takht. with a member of the series Sevanenses Grossh. from 
the next subsection. It is possible that S. amphichlora derives directly from S. sos- 
nowskyi Takht. 

84. S. araratica Grossh. in Izv. Azerb. fil. AN SSSR, 3 ( W45) 84. S. orientalis var. 
pinnatifida N. Pop. in Mat. fl. Kavk. IV, 3(1916) 79, p. p. non Rchb. - Ic: Grossg., 
op. cit. 85, Figure 18. 

Perennial, 7-10 cm high, with procumbent, rooting rhizome; stems ascending, sparsely 
covered with short spreading hairs, pale green; leaves small, 4-15 mm long, 2-7 mm wide, 
broadly ovate or oblong-ovate, usually truncate at base, obtuse or acute, very finely ap- 
pressed-hairy above, sparsely gray-tomentose beneath, very deeply pinnatifid-dentate with 
4-7 teeth on each margin, these long, narrow, obtuse oblong or linear, apically rounded, 
often to middle of half-blade and commonly 1/3 tlie width of blade; petioles slender, to 
1.2 cm long, nearly the length of blade, sparsely patent-tomentose; inflorescence dense, 
ovate, 2.5-3.5 cm long; lowermost bracts large, 1-1. 5 cm long, 5-7 mm wide, oblong-ovate, 
acuminate, in upper part with 3-5 triangular teeth at each side; middle bracts smaller, 0.6- 
1 cm long, 3-4 (5) mm wide, entire or rarely with 1-2 teeth above; all bracts short-patent- 
hairy and stipitate-glandular; corolla ca. 2.5 cm long, 5-6 mm wide at throat, yellow, pu- 
bescent and short-stipitate-glandular outside. 

Caucasus: S. Transc? Gen.distr.: N.Iran. Described from Great Ararat peak near 
Sardar-Bulak (from Khodz'ko specimens). Type in Leningrad. 

Note. It is possible that tliis species, provisionally included in "Flora of the USSR," 
does not occur within the Soviet Union. 



Series 2. Adenostegiae Juz. - True subshrubs, with shoots strongly lignified in lower 
part; vesture of bracts consisting mainly of large long-stipitatc glands clearly visible even 
under low magnification. 

Note. The species of this series are very closely related to each other, while their affin- 
168 ity to species of the preceding series is much less evident; they may in fact be of different 
derivation. Whereas the Eu-platystegiae are clearly connected phylogenetically with the 
Oxystegiae, the series Adenostegiae may have, in our view, originated directly from species 



112 



of the series Multicaules, or, rather, from its progenitor, as evidenced by the proximity of 
verticillasters in the inflorescence (for example, a species such as S. gontscharovii Juz. 
which is very close to S. ju/.epc/ukii Gontsch.). Should our assumption be confirmed 
by further investigation, the taxonornic position of the scries Aflcnostcgiae would fiJive to 
be changed. 

85. S. adenostegia Hriq. in Bot. Tidskr. 28 (1908) 233. 

Perennial; rcn^t robust, woody, oblique or vertical, branching; sterns rnfjstly numerous, 
12-32 cm high, ascending or erect, branching at base, usually simple above, subvirgate, 
usually cinereous, with copious short partly crisp hairs, often slightly suffused with lilac 
in lower part; leaves rather small, 0.6-2.5 (3) cm long, 0.3-2 cm wide, ovate or the upper 
almost triangular, at base truncate, subcordate or slightly tapering to petiole and (jbtusoly 
angled (the upper obtuse or often acute at base), deeply and often rather regularly crenate 
with 5-10 crenations 1-2 mm long on each margin, rugose, green above with scattered 
short fine hairs or often grayish with sparse very fine appressed tc>mentum, appressed- 
white- or cinereous-tc^mentose beneath, with very prominent midrib and lateral veins, 
these spreading flabellately from base; petioles dilated, slightly winged, tomentose, 0.1- 
1 cm long, half the length of blade; inflorescence 4-1 cm long, loose; bracts to 1 .3 cm 
long, 1 .5 cm wide, broadly ovate, shorl-apiculate, sometimes (especially the lr;wer) sligl-itly 
crenulate, mostly prominently netted-nerved, sparsely covered with distinct long-stipitate 
glands intermixed with long simple hairs, usually pale green; calyx at anthesis small, 3- 
4 mm long, glandular-hairy, viscous; corolla large, to 3-3.5 cm long, glandular-hairy out- 
side, yellow; lower lip usually with purple lateral lobes. May July. (Plate X, Figure 3.) 

Stony and clayey mcjuntain slopes and outcr(jps, dry riverbeds and streams, stony 
placers and pebbles along banks of rivers. Centr. Asia: T. Sh.(Susamyr Range), Pam.-Al. 
(Alai Range, Turkestan Range, Zeravshan). Endemic. Described from the southern slope 
of Alai Range near Gul'chi. Type in Copenhagen, isotype (Paulsen, Ncj. 3y/>) in Leningrad. 

169 86. S.bucharica Ju/.. in iiot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 406. 

Perennial; stems often few, 12-28 cm long, ascending at base and very often branching 
above, covered with short spreading curved or crisp hairs, often turning lilac, sometimes 
only at base; leaves medium to large, 0.6-3 cm long, 0.5-2.5 cm wide, broadly ovate, at 
base usually shallowly cordate, rounded or obtuse at apex, rather shallowly and coarsely 
crenate with 4-8 rounded crenations on each margin, green or often grayish-green, very 
finely appressed-tomentose above, densely white- or gray-tomentose beneath, the veins 
mostly not prominent but not completely concealed by tomentum, hence network of 
veins usually distinct; inflorescence 2.5-4 cm long at first, considerably elongating after 
flowering and reaching 10cm; bracts to 12 mm long, 10 mm wide, covered with long 
eglandulose hairs and often less numerous stipitate glands, subacuminate or subacute at 
apex, pale green, often turning lilac; corolla medium-sized, 1 .5-2.5 cm long, bright yellow, 
with brownish lower lip; nutlets to 2 mm long, whitish, densely pubescent. Otherwise, 
similar to S. adenostegia Briq. May- June. 

Clayey and stony mountain slopes, granite (jutcrops, riverbanks. (Jentr. Asia; Pam.- 
Al. (within borders of Tadzhikistan). Endemic. Described from Chul'bair Mountains near 
Ak-Su village and from Babatag Range (northeastern tip), north of Dzhambulak village. 
Type and paratype in Ix-ningrad. 



113 



87. S. gontscharovii Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 407. 
Perennial; stems few, 10-35 cm long, ascending, rather sparingly and shortly pubescent, 

more or less lilac; leaves 0.7-2.4 cm long, 0.5-1 .6 cm wide, broadly ovate or nearly rhom- 
bic, with rounded or obtusely angled to rectangular base, usually subacute, incised-crenate 
with (3) 4-7 crenations to 3 mm long on each margin, green with scattered hairs and im- 
pressed veins above, grayish beneath with sparse slightly spreading tomentum, the violet 
coloration of the leaf undersurface showing more or less clearly through the tomentum 
(as in S. haematochlora Juz.), the veins not or slightly prominent; petioles hairy, to 
1 cm long; inflorescence 3-6 cm long, loose; bracts 0.8-1 .3 cm long, 0.5-0.9 cm wide, 
short- or long-acuminate, lower bracts finely dentate, those above them with isolated 

170 small teeth, rather copiously but not evenly covered (mainly along margins and midrib) 
with simple, long, spreading hairs and numerous large stipitate and sessile glands; corolla 
2-3 cm long, completely yellow or in rare cases the upper lip and its appendages red. 
Otherwise, like S. adenostegia Briq. and S. bucharica Juz. May— June. 

Stony mountain slopes. — Centr. Asia: Pam.-Al. (S. Tadzhikistan). Endemic. Describ- 
ed from Kulyab region, near Arzanchi village. Type in Leningrad. 

88. S. haematochlora Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV(195 1)408, s.str. 
Subshrub with sturdy woody flexuous caudex; stems rather numerous, 15-35 cm long, 

woody in lower part, procumbent or ascending, often rooting at nodes and strongly branch- 
ing, producing many ascending or suberect shoots, these sparsely covered with short curved 
or sUghtly crisp hairs, often lilac; leaves medium to fairly large, 0.7-4 cm long, 0.4-3 cm 
wide, broadly ovate or ovate-rhombic, with rounded or usually angled (broadly cuneate) 
base, tapering to petiole, obtuse to subacute at apex, incised-crenate-dentate with 4-10 
rather large uneven obtuse or subacute or sometimes acute teeth on each margin, green 
above, glabrous or covered with very fine faintly discernible sparse or rather dense hairs, 
usually sparsely short-tomentose beneath, often lilac between the veins, these densely pu- 
bescent, yellowish, very distinct against the background of anthocyanin coloring and to- 
mentum; petioles usually well developed, even in upper leaves, 0.2-1 .5 cm long, pubescent; 
inflorescence 2-4 cm long, in fruit to 10 cm; bracts dark green, the lower being 1-1.6 cm 
long and 0.6-1 cm wide, ovate or rather broadly ovate, gradually or (especially the lower) 
abruptly long-acuminate, often terminating in a slightly curved point, usually with sparse 
simple long hairs and numerous stipitate glands only on the margins and nerves, otherwise 
sparingly pubescent and glandular; calyx at anthesis ca. 2 mm long, hairy and glandular; 
corolla 1.5-2.5 cm long, yellow, often with purple or dark spot on lower lip, covered out- 
side with short-stipitate glands; nutlets ca. 1.5 mm long, whitish, densely pubescent. 
May -July. 

Stony, mostly grassy mountain slopes. — Centr. Asia: T. Sh. (western part, mainly 
along valleys of right-hand tributaries of Chatkal River). Endemic. Described from Aktash 

171 forest district near Tashkent (in southern Kazakhstan) and from Bol'shoi Chimgan. Type 
and paratypes in Leningrad. 

Note. A unique plant, collected by Z.A. Minkvits in the Ugam River valley, is allied to 
S. haematochlora, but differs from it in the smallish leaves and, in particular, the narrow- 
er, ovate-lanceolate, gradually acuminate, pale green bracts, covered with very short, ap- 
pressed and curved hairs interspersed with small, very short-stipitate or subsessile glands, 



114 



and in the flowers up to 2 cm long. We suspect that this plant is a hybrid (S. haemato- 
chlora s.l. X S. microdasys Juz.) and we have named it S. minkwitziae Juz. (p. 346). 

89. S. tschimganica Juz. sp. nov. in Addenda XIX, 346. — S. haematochlora Juz. in 
Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 408, p. p. 

Perennial; close to preceding species but differing in the following characters: plant 
lower, 7-25 cm high; leaves smaller and narrower, to 2 cm long, 1 .4 cm wide, ovate or 
narrowly rhombic, cuneate at base, with 3-7 teeth on each margin, very sparingly short- 
tomentose beneath (some leaves subglabrous); petioles shorter; inflorescence 2-3 cm long, 
up to 4 cm in fruit, loose; bracts narrowly ovate, gradually tapering to a mostly straight 
point, very sparingly pubescent between nerves or subglabrous. Otherwise, like S. haema- 
tochlora Juz. May— June. (Plate X, Figure 4.) 

Stony, dry and grassy mountain slopes. - Centr. Asia: T. Sh. (western part: Chimgan 
and right bank of Central and Upper Angren). Endemic. Described from Bol'shoi Chim- 
gan. Type in Leningrad. 

90. S. urticifolia Juz. et Vved. in Addenda XIX, (p. 346). 

Perennial; caudex and bases of stems woody; annotinous shoots suberect, pale green or 
pale violet, covered with fine curved hairs; leaves 1-3.2 cm long, 0.6-1 .8 cm wide, ovate, 
with broadly cuneate base, acute at apex, coarsely incised-crenate-dentate with 5-7 oblique- 
ly antrorse teeth on each margin, green on both sides, glabrous above, faintly tomentulose 
or sometimes glabrescent beneath, without a trace of anthocyanin coloration; petioles 1- 
8 mm long, slightly tomentose, eglandulose; inflorescence ca. 4 cm long, in fruit to 7 cm, 
rather dense; bracts pale green, rather small, usually to 1 cm long and wide, broadly ovate 
or suborbicular, shortly and abruptly acuminate, the lower often remotely dentate, copi- 
172 ously covered with long spreading simple hairs and much shorter stipitate glands; corolla 
yellow, short-stipitate-glandular outside, with broad tube. June. 

Slopes of hills, on clayey gravelly soil. — Centr. Asia: T. Sh. (western part: foothills of 
Fergana Range). Endemic. Described from the vicinity of Cava village. Type in Tashkent. 

91. S. iskanderi Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 410. 
Subshrub; stems4)rocumbent, woody, 12-30 cm long, branching above, producing low 

antrorse leafy tomentose shoots; leaves small, 4-12 mm long, 2-10 mm wide, ovate to sub- 
orbicular-ovate, with 2-5 rather deeply incised unequal often acute teeth on each margin, 
gray, tomentose above with impressed veins, gray- or white-tomentose beneath, the veins not 
prominent, sparingly pubescent and hence distinctly visible; petioles 1-8 mm long; inflo- 
rescence 2-4 cm long; bracts small, to 1 cm long, 6 mm wide, rhombic-elliptic, obtuse or 
acute, more or less violet, with prominent longitudinal nerves; corolla 2.2-3 cm long, yel- 
low, the upper lip with purple lobes. Otherwise, like S. adenostegia Briq.; it probably 
represents a high-mountain race of that species. May— July. 

Dry stony slopes (in Central Asian juniper zone). — Centr. Asia: Pam.-Al. (basin of 
Zeravshan). Endemic. Described from Iskander Lake (Iskander-Kul'). Type in Leningrad. 

92. S. pycnoclada Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (195 1) 41 1 . 
Perennial; very similar to preceding species but the whole plant more compact, with 



115 



(173) 




PLATE X. 1 - Scutellaria transiliensis Juz., summit of stem, bract; 2 - S. sieversii Bge., general 
aspect, flower; 3 - S. adenostegia Briq., general aspect, bract, fruiting calyx; 4 - S. tschimganica 
Juz., summit of stem, corolla; 5 — S. pycnoclada Juz., general aspect, bract. 



116 



crowded shoots; leaves smaller and more regular, less deeply toothed, often with very 
prominent veins on the underside; bracts slightly wider and more obtuse, often colorless; 
smaller flowers 1.5-2.5 cm long, plain yellow. June— July. (Plate X, Figure 5.) 

Mountain slopes, dry riverbanks, bottom of gorges. — Centr. Asia: T. Sh. (western 
part). Endemic. Described from upper reaches of Chatkal River (in southern Kirghizia). 
Type in Leningrad. 

175 93. S. angrenica Juz. et Vved. in Addenda XIX, 347. 

Perennial; similar to S. pycnoclada Juz., but shoots more erect, with longer and 
looser (sometimes spreading) hairs; leaves narrower, with narrower cuneate base, acute 
at apex, more densely pubescent above, the margins not revolute, with more deeply in- 
cised acute obUquely antrorse teeth; bracts narrower and more acute; corolla small, ca. 
2mm long, stipitate-glandular outside. Otherwise, like S. pycnoclada Juz. June. 

Stony and gravelly slopes, dry stony-sandy hillocks, quackgrass steppes. — Centr. Asia: 
W. T. Sh. (basin of Angren River, especially along its left bank). Described from the 
northern slope of Kuramin Range, Lyailak-Sai. Type in Tashkent. 



Subsection 4. Alpinae Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 363. - 
Shrubs, subshrubs or perennial herbs; leaves green or grayish-green on both sides, rather 
sparingly or densely covered with simple straight or sUghtly crisp hairs (sometimes with 
intermixed stipitate glands), but always without true tomentum; inflorescence spikelike, 
uninterrupted; bracts broad, ovate or eUiptic, scarious or sometimes sub herbaceous; 
corolla-tube usually rather short and wide, abruptly dilated at throat; lower lip of corolla 
mostly rather short, as long as or shorter than the upper lip. 



Series 1. Cordifrondes Juz. — Small shrubs; vesture of stems and leaves including 
long-stipitate glandular hairs; lower hp of corolla not hairy above. 

94. S. cordifrons Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 363. - S. al- 
pinavar. cordifolia Rgl. inTr. Bot. sada, VI (1879) 365. - S. cordifolia O. Fedtsch. 
in 0. and B. Fedch. Perech. rast. Turk. V (1913) 158, non Benth. nee Miihlenb. 

Low, erect or sometimes nearly prostrate shrub 12-40 cm high, with branching flexuous 
nodose brownish-gray stem; leafy branches with long internodes, densely covered with 
horizontally spreading long coarse simple and glandular hairs; leaves 0.7-2.7 cm long, 0.6- 
3 cm wide, cordate or cordate-reniform, with broadly cordate or truncate base, subobtuse 
to obtuse, often rounded at apex, unevenly and coarsely crenate with 4-10 teeth (to 1- 
176 3 mm) on each margin, yellowish-green on both sides, covered sparsely or rather densely 
above, usually densely beneath, with fairly simple stiff long hairs and large stipitate glands, 
rugose above, prominently veined beneath; petioles 0.3-1 .4 cm long, densely patent-hairy 
and glandular, approximately half as long as blade; upper leaves subsessile; inflorescence 
2-6 cm long, at first rather dense, later loose; bracts to 1-1.4 cm long, 0.6-1.2 cm wide, 
subherbaceous, short-elliptic, slightly pointed at both ends, with few or solitary teeth or 
(the upper) entire, nearly flat, green, rather densely covered, especially on nerves and 



117 



margins, with long simple and glandular hairs; calyx to 3 mm long at anthesis, densely 
patent-hairy and stipitate-glandular; corolla small, 1.5-2.5 cm long, with a broad tube, 
densely stipitate-glandular outside, yellow, the tip and lobes of upper lip purple and the 
lower lip with a purple spot (sometimes completely purple); nutlets 1-1.5 mm long, angled, 
gray, covered with very fine stellate hairs. July— August. (Plate XI, Figure 6.) 

Rocks and stony placers in subalpine and alpine mountain belts, sides of ravines. — 
Centr. Asia: W. T. Sh. Endemic. Described from Aktag-Tau Range (Korol'kov collec- 
tions). Type in Leningrad. 



Series 2. Adsurgentes Juz. — Like the preceding series, but vesture of stems and 
leaves without glandular hairs or with rather short-stipitate glands; lower lip of corolla 
pubescent above. 

95. S. adsurgens M. Pop. in Bot. mat. gerb. Bot. sada, V, 10 (1924) 154. 

Shrub, 20-40 cm high; stem erect or often procumbent or ascending, branching from 
base or slightly above base; old shoots stout, straight or flexuous, woody, glabrous, with 
brown bark, with remnants of previous year's petioles at nodes; young shoots 5-10 cm 
long, leafy or floriferous, densely covered with short spreading slightly crisp hairs; inter- 
nodes of medium length; leaves 0.8-2 cm long, 0.1-1.5 cm wide, ovate or oblong-ovate, 
broadly cuneate or truncate or subcordate at base, acuminate, coarsely and subacutely 
dentate with (3) 4-6 teeth (1) 1.5-3 mm high on each margin, green on both sides, sub- 
glabrous or sparsely covered above, more densely beneath, with rather short curved hairs 
interspersed with very small stipitate glands, the veins rather strongly impressed above, 
177 very prominent beneath; petioles 0.5-1 cm long, slender, often densely hairy; inflores- 
cence 1-3.5 cm long, to 6 mm in fruit; bracts loosely disposed, eUiptic or oblong-elliptic, 
pointed at both ends, except for the lowermost usually entire, almost flat, green, rather 
densely covered with simple long hairs and rather long stipitate glands, prominently nerved, 
the lower ca. 1 cm long, 4 mm wide; calyx to 3 mm long at anthesis, hairy and glandular, 
with large scutellum; corolla small, 12-16 mm long, densely covered outside with stipitate 
glands, yellow; lower lip purple-red with a yellow spot at middle, pubescent above (some- 
times only at throat); nutlets ca. 1 mm long, cinereous with stellate hairs. 

Mountains, at the upper limit of Central Asian juniper zone. - Centr. Asia: T. Sh. 
(Kara-Tau, Talass Ala-Tau, Ugam mountains). Endemic. Described from Ugam Mountains 
(near Tashkent), Kovak-Ata survey mark. Type (M. Popov, No. 1282) in Tashkent. 

Note. The most typical specimens of this plant occur in the Kara-Tau and Talass Ala- 
Tau mountains. Before we became acquainted with the description and the type of S. ad- 
surgens M. Pop., we had named the Kara-Tau plant in schedis S. dendroides Juz., and 
this name is still to be found in some herbaria. It was with some doubt that we identi- 
fied the Tashkent plant described by Popov with the Kara-Tau (and Talass) specimens, 
although it gravitates somewhat toward the next species, described from Gissar Range but 
apparently also widespread in West Tien Shan. 

Economic importance. A high quality dyestuff (aqueous extraction imparts a golden- 
yellow color to wool and silk). 



118 



96. S. phyllostachya Juz. in Bot. mat. gerb. Bot inst. AN SSSR, XIV (1951) 364. - 
Ic: Yuz., op. cit. 365, Figure 1. 

Shrub; 12-35 cm high, with ascending or procumbent branching stem; old shoots 
slender or fairly stout, mostly flexuous, woody, with gray or brown bark, nodose; young 
shoots leafy or floriferous, patent-hairy, usually short-stipitate-glandular; internodes me- 
dium to very long; leaves medium-sized, 3.5-20 mm long, 4-15 mm wide, broadly ovate 
to orbicular, subcordate or truncate, rarely broadly cuneate at base, obtuse or rounded at 
178 apex, coarsely and unevenly crenate with 2-5 (7) obtuse teeth 0.5-2 (3) mm high on each 
margin, green on both sides and covered, more densely beneath, with short crisp hairs 
interspersed with short-stipitate glands; petioles 0.2-1 .7 cm half as long to as long as blade, 
patent-hairy and stipitate-glandular; inflorescence 2-5 cm long, loose; bracts to 1 .5 cm 
long, 1 cm wide, broadly ovate or elliptic, obtuse to subacute, green, foUaceous, the lower- 
most pair usually resembling cauline leaves and distinguished only by the very short petioles 
and reduced crenations, these gradually disappearing toward summit though sporadically 
occurring even in upper bracts, the margins and midrib beset with long hairs; calyx ca. 
3 mm long, sparsely long-hairy and short-glandular; corolla rather large, 2-3 cm long, 
sparsely long-hairy and short-glandular; corolla rather large, 2-3 cm long, yellow or pink- 
ish, sparingly pubescent and densely glandular outside; upper lip slightly longer than the 
lower; nutlets to 1 .5 mm long, angled and broadly ovoid, gray. June— July. 

Subalpine mountain belt. - Centr. Asia: T. Sh. (western part), Pam.-Al. (Tadzhikistan). 
Endemic. Described from the watershed between Unou and Lyuchob on Gissar Range. 
Type in Leningrad. 



Series 3. Sevanenses Grossh. in Izv. Azerb. fil. AN SSSR, 3 (1943) 83. - Like the 
preceding series but structurally the corolla approaches the type characteristic of the sub- 
section Orientales Juz.; lower lip glabrous. 

97. S. sevanensis Sosn. in Grossg. Fl. Kavk. Ill (1932) 286. - Ic: Grossg. in Izv. 
Azerb. fil. AN SSSR, 3 (1945) 85, Figure 15. 

Low-growing, prostrate or ascending shrub 8-25 cm high, the rough branching trunk 
covered with brown bark and producing numerous leafy elongate fertile and short sterile 
green shoots or branches, these sparsely covered with short curved hairs interspersed with 
short-stipitate glands; leaves 0.7-2 cm long, 0.4-1.5 cm wide, ovate, obtusish or obtusely 
angled at base, obtuse at apex, with 4-6 semi-elliptic or oblong obtuse teeth on each mar- 
gin, green on both sides, very finely sparsely or- densely pubescent and glandular but never 
white-tomentose beneath; petioles slender, 0.2-1 cm long, pubescent and glandular; inflo- 
rescence 3-5.5 cm long, dense at first, becoming lax; lower bracts rather large, 1-1.6 cm 
long, 0.7-1 cm wide, rhombic-ovate, attenuate at both ends, acutish or short-acuminate at 
179 apex, entire, with prominent longitudinal nerves, covered with short spreading hairs and 
short-stipitate glands, green, often violet-tinged at apex; calyx ca. 2 mm long at anthesis, 
densely stipitate-glandular; corolla 2-2.7 cm long, pale yellow, with broad tube to 7 mm 
broad at throat, relatively short upper lip and large lower lip, covered outside with short 
spreading hairs and glands. June— August. 

Meadows and pastures, rocks and stony placesin upper (alpine and subalpine) mountain 



119 



belts. - Caucasus: S. Transc. (near Lake Sevan, Zangezur, Nakhichevan ASSR). Endemic. 
Apparently described from the vicinity of Tazakend (formerly Novobayazetskii district). 
Type (or paratype?) in Leningrad. 

Note. We believe that this species is very closely allied to S. virens Boiss. et Ky., 
described from Lesser Armenia; thus Grossgeim's proposed name for the series in which 
the species is included cannot be regarded as a good choice. 



Series 4. Supinae Juz. - Perennial herbs, more or less mesophytic in habit; leaves 
rather sparingly pubescent, and numerous (commonly 7-1 1 per margin, rarely less) mostly 
very distinct teeth; bracts rather loosely disposed. 



98. S. verna Bess. Prim. fl. Galic. II (1809) 43; Yuz. in Tr. N.-d. inst. bot. IV, 206. - 
S. alpina j3. lupulina Ldb. Fl. Ross. Ill (1847-1849) 394, p. p. - Exs.: PI. pol. exs. 
No. 259 (nom. S. lupulina L.). 

Subshrub, with woody crooked root; stems few, 10-30 cm high, ascending at base, 
generally curved or slightly flexuous, simple or often branching, usually pale violet, except 
for lower part densely covered with fme white reclinate hairs, at summit and in inflores- 
cence densely stipitate-glandular and patent-hairy; leaves medium-sized, 1.2-2.3 cm long, 
0.5-1.5 cm wide, narrowly ovate or usually ovate, approximately twice as long as wide, 
usually truncate, rarely rounded or slightly tapering at base, mostly obtuse at apex, cover- 
ed above with minute hardly discernible spreading hairs, copiously glandular-punctate be- 
neath, with prominent, very finely pubescent veins, evenly and regularly crenate-dentate 
with 5-7 obtuse rather high and slightly spreading teeth on each margin, the outer margin 
of each tooth much longer than the inner; petioles of lower leaves to 1 cm long, of median 
ca. 5 cm, those of upper leaves much shorter (leaves subsessile); petiole and margin of 
blade at base with fine and rather long, horizontally spreading or recurved hairs (seeming- 
ly ciliate); inflorescence ca. 2 cm long at first, later ca. 9 cm, with flowers in loosely dis- 
180 posed verticillasters; bracts early deciduous, broad-ovate or ovate, the lower 1-2 cm long, 
0.8-1.2 cm wide, gradually tapering at apex to a short or longish point, almost scarious, 
pale green or very often suffused with lilac, glandular, long-ciUate at margin, the nerves 
slightly prominent; calyx at an thesis ca. 2 mm long, densely hairy and glandular; corolla 
small or often large, 1-2.5 cm long, yellow, densely hairy and glandular outside. July- 
August. 

Limestone outcrops. — European part: M. Dnp. (western part), U. Dns. Gen. distr.: 
? Centr.Eur. (Hungary). Described from Zaleshchiki on the Dniester. Type (or isotype) 
in Leningrad. 

Note. The identity of the Hungarian (Transylvanian) plant with S. verna Bess, is 
doubtful and in any case needs verification. Its flowers are apparently larger than those 
of the true S. verna from Podolia and its bracts do not fall quite so early. It was publish- 
ed: Dorfler, Herb. norm. No. 5154 (under the name S. supina). 

99. S. creticola Juz. in Tr. N.-d. inst. bot. IV (1941) 205. - S. alpina jS. lupulina 
Schmalh. Fl. II(1897)327,p.p. 

Subshrub, with woody flexuous and nodose root; stems many, 10-25 (30) cm high, 
at base usually ascending or rarely suberect, usually curved or slightly flexuous, simple or 

120 



often short-branching, covered (except at base) with fine white retrorse curved hairs, in 
inflorescence densely stipitate-glandular and short-patent-hairy; leaves glabrous above, 
densely glandular-punctate beneath, very finely pubescent on the prominent veins, medi- 
um-sized, 1-3 cm long, 0.3-1.2 cm wide, ovate or usually narrowly ovate or even ovate- 
lanceolate, approximately twice as long as wide, mostly truncate, rarely rounded or slight- 
ly tapering at base, unevenly and shallowly toothed, acute and entire at apex, the upper- 
most leaves often nearly entire, with 5-7 teeth on each margin, in lower leaves the'teeth often 
crenately rounded at apex, in others nearly rectangular, asymmetrical, the outer margin 
much longer than inner and nearly parallel to leaf margin and hence the teeth very low; 
petioles short, in lower leaves to 7 mm long, in middle leaves 2-4 mm, the vesture as on 
stems; inflorescence ca. 2 cm long, dense, later to 6 cm long, the flowers in loosely dis- 
posed verticillasters, lower bracts 1-1.7 cm long, 0.4-1 cm wide, ovate or ovate-lanceolate, 

181 gradually tapering to a short or fairly long point, almost scarious, pale green or very often 
lilac-tinged, glandular, with long-ciliate margins, the nerves slightly prominent; calyx ca. 
2 mm long at anthesis, densely hairy and glandular; corolla rather small, 1.5-2 cm long, 
yellow, densely hairy and glandular outside. June— August. 

Chalk outcrops and slopes. - European part: V.-Don (southwestern part) and L. Don 
(western part). Endemic. Described from the vicinity of former Svyatogorskii monastery 
and Derkul'skaya steppe. Type and para types in Leningrad. 

100. S. chitrovoi Juz. in Spisokrast. gerb. fl. SSSR, XI (1949) 147. - Exs.: GRF, 
No. 3472. 

Subshrub, with woody flexuous rhizome; stems few, 25-56 cm long, ascending at base, 
woody in lower part, strongly flexuous, covered in inflorescence with profuse stipitate 
glands and short (though longer than glands) straight spreading hairs; branches suberect, 
elongate, flexuous, violet (like stems) and covered over nearly the whole length with fine 
white recUnate curved hairs; leaves on main stems large, 1.8-3.5 cm long, 0.7-2 cm wide, 
broadly ovate, 1 1/2-2 times as long as wide, subtruncate or very often subcordate, rarely 
rounded at base, obtuse or rarely subacute at apex, unevenly dentate, with 6-1 1 rather 
high obtuse or rarely acute teeth; upper cauUne and ramal leaves narrower, ovate or 
lanceolate, acutish with fewer shorter acute teeth, distally entire, subglabrous above, 
densely glandular-punctate beneath, the veins prominent, very finely hairy; lower leaves 
long-petioled (ca. 1 cm), other leaves with shorter petioles and vesture like that of branches; 
inflorescences terminal, at first 3-4 cm long, dense, later elongating to more than 10 cm, 
with loosely disposed whorls; bracts small, the lower 0.8-1 .2 cm long, 5-8 mm wide, all 
broadly ovate or elUptic, abruptly attenuate to a very short point or subobtuse, almost 
scarious, pale green or sometimes apically slightly violet, glandular and hairy, with long- 
ciUate margins, the nerves somewhat prominent; calyx small, 2-2.5 mm long, densely 
hairy and glandular; corolla rather large (1.5) 1.8-2.5 cm long, densely hairy and glandu- 
lar outside, yellow, the upper lip and lateral lobes violet. May— June. 

Meadows, calcareous taluses and debris. - European part: V.-Don (western part). 
Endemic. Described from El'tsa (bank of Sosna River). Type in Leningrad. 

182 101 . S. cisvolgensis Juz. in Spisok rast. gerb. fl. SSSR, XI (1945) 149. 

Subshrub, with vertical sparingly branching slightly flexuous root; stems few to fairly 



121 



numerous, 15-20 cm long, arched-ascending at base, branching only at base, violet, in 
lower part glabrous, with fine white antrorse hairs above, in inflorescence densely patent- 
hairy and glandular; leaves medium-sized, 1.5-3 cm long, (0.6) 0.9-1.5 cm wide, oblong- 
ovate, l-lVz times as long as wide, usually rounded at base, obtuse at apex or the upper 
subacute, shallowly crenate-dentate with 3-7 strongly tapering obtuse or acutish teeth on 
each margin, with scattered hairs above, glandular-punctate beneath, very finely hairy 
along the veins; petioles to 1 .2 cm long, covered with hairs of varying length; upper leaves 
with petioles 1-2 mm long; inflorescence 2-3 cm long, dense, later sUghtly elongating; 
bracts rather short, the lower 1.3-1.5 cm long, 0.8-1 cm wide, broadly ovate, gradually 
attenuate to a well developed point, almost scarious, pale green, slightly hairy and glandular, 
with cihate margin; calyx 3-4 mm long at an thesis, densely hairy and glandular; corolla 
rather small, to 2 cm long, yellow, hairy and glandular outside. June. 

Steppes. - European part: V.-Don (eastern part). Endemic. Described from Lavin- 
skaya steppe (Ul'yanov Region), between Kanadya and Lavo. Type in Leningrad. 

Note. A little known species that is apparently becoming extinct; needs further 
investigation. 

102. S. oxyphylla Juz. in Spisok rast. gerb. fl. SSSR, XI (1945) 149. - S. alpina jS. 
lupulina Ldb.Fl. Ross. 111(1849)394, p.p.; Shmal'g. Fl. II, 327, p.p. 

Subshrub; root and caudex woody, flexuous, tuberculate; stems few or many, 15- 
40 cm high, ascending at base, usually slightly curved or flexuous, simple or often branch- 
ing, mostly violet-tinged, covered, except in lower woody part, with rather copious fine 
white reclinate curved hairs, in inflorescence densely stipitate-glandular and patent-hairy; 
leaves medium-sized to fairly large, 1 .3-4 cm long, 0.5-1 .7 cm wide, ovate, 2 V3-2 V2 fimes 
as long as wide, obtusish or rounded or sometimes broadly cuneate at base, subacute or 
upper leaves acute, attenuate at apex, irregularly serrate-dentate or crenate-serrate with 
183 5-8 rather large high obtuse or often acute teeth on each margin, the outer margin of 
teeth straight or mostly slightly curved, twice as long as their inner margin, distally entire, 
glabrous or sparsely and finely appressed-hairy above, glandular-punctate beneath, glabrous 
or very finely hairy on the veins, the petioles to 1 .2 cm long (lower leaves); uppermost 
leaves short-petioled; vesture of petioles on the margins as on stems, elsewhere as on veins 
on the underside of leaves; inflorescence ca. 3-5 cm long, rather dense, later to 9 cm long, 
with flowers in loosely disposed verticillasters; lower bracts (1) 1.5-2 cm long, (5) 6- 
8 mm wide, oval or elongate-ovate, pointed at apex, somewhat scarious, pale green or 
suffused with lilac, sparsely glandular-hairy or subglabrous, with long-ciliate and stipitate- 
glandular margin, the veins somewhat prominent; calyx at anthesis ca. 3 mm long, densely 
hairy and glandular; corolla 2.5-3.5 cm long, plain yellow or often with 3 violet spots (at 
the tip of upper Hp and on lateral lobes), hairy and glandular outside. June -August. 
(Plate XI, Figure 1.) 

Steppes, stony slopes, outcrops, bluffs, pebbles. — European part: Transv., V.-Kama; 
West Siberia: U. Tob. Endemic. Described from Bashkiria, Mesyagutov canton, near 
Duvan and Arkhaul. Type and paratypes in Leningrad. 

103. S. supina L. Sp. pi. (1753) 598, p. p. (excl. pi. e "Tataria" et syn. Tournef.); 
KryL Fl. Zap. Sib. IX, 2302. - S. lupulina L. Sp. pi. II (1763) 835. - S. alpina i^. 



ni 



lupulina Benth. in DC. Prodr. XII (1848) 412; Ldb. Fl. Ross. Ill, 394, p. p. - Exs.: 
E. Smirnov, PI. alt. exs. No. 69. 

Subshrub, with woody oblique or creeping rhizome; stems rather numerous, 10-45 cm 
long, ascending or sometimes suberect, simple or branching, covered with fine subappres- 
sed usually reclinate hairs and longer divergent or horizontally spreading hairs (the latter 
often very copious, especially in upper part of stem), pale yellowish-green or fairly purple; 
leaves 1-4 cm long, 0.6-2 cm wide, oblong-ovate or ovate, obtuse, only the upper some- 
times subacute, shallowly or rather deeply crenate-dentate with 4-7 teeth on each margin, 
the upper surface sparsely or rather copiously hairy, the hairs stiff and mostly curved, 
often thickened at base, the lower surface glandular-punctate and often with same vesture 
as the upper side or with long or short spreading hairs confined to veins; petioles of lower 

184 leaves approximately half the length of blade, with vesture as on stems; upper leaves sub- 
sessile; inflorescence short, compact, slightly elongating in fruit, 2.5-4 cm long; bracts 
large, the lower 1 .5-2 cm long, 0.5-1 .2 cm wide, ovate or rarely oblong-ovate, obtuse to 
subacute, pale green or lilac-tinged, rather sparsely long-hairy, with intermixed shorter 
glandular hairs or sometimes subglabrous, the margins long-cilia te ; calyx ca. 2 mm long, 
glandular-hairy; corolla usually large, 2.2-3.5 cm long, yellow, glandular-hairy outside; 
nutlets ca. 1 .5 mm long, triangular-ovoid, covered with short stellate hairs. End of May — 
August. 

Stony mountain slopes or meadows, ascends to the alpine belt. — West Siberia: Irt.? 
Alt.; East Siberia: Ang. -Say. (Minusinsk district); Centr. Asia: Dzu.-Tarb. Gen.distr.: 
W. Mong. Described from Siberia. Type in London. 

Note. When writing about the occurrence of S. supina L. in Siberia and "Tataria," 
Linnaeus probably had in mind S. oxyphylla Juz. as well as the Siberian plant. Fixing 
the name S. supina L. for the Siberian (Altai) plant was proposed by us (cp.: Spisok 
rast. gerb. fl. SSSR, XI (1945) 149 (?)). 

104. S. altaica Fisch. ex Sweet Brit. Flow. Card. (1823-1825) 45; Spreng. Suppl. 
Syst. veg. (1828) 16; Kryl. Fl. Zap. Sib. IX, 2302. -S.lupuhna ^. violacea Bge. in 
Ldb. Fl. Alt. II (1830) 394. - S. alpina Ldb. Fl. Ross. Ill (1849) 394 (excl. var. (J) et 
auct. plur. non L. — Ic: Sweet, 1. c. tab. 45. 

Perennial; rhizome rather stout, oblique or creeping, woody; stems few, 10-40 cm 
long, ascending or suberect, simple or branching, covered with short reclinate hairs and, 
sometimes only in upper part, long spreading hairs; leaves 1.2-4 cm long, 0.5-2.5 cm 
wide, ovate or oblong-ovate, rounded or subcordate or sometimes slightly tapering at base, 
subobtuse to subacute, crenate-dentate with 4-9 obtuse or acute teeth on each margin, 
sparsely covered on both sides (mainly on veins beneath) with short or fairly long hairs or 
sometimes subglabrous; petioles of lower leaves approximately 1/3 as long as blade, with 
vesture like that on stems; upper leaves subsessile; inflorescence at first 2.5-4 cm long, to 
10 cm after flowering; bracts medium-sized, the lower 14-20 mm long, 5-1 2 mm wide, 
ovate, obtuse or acute, scarious or rather firm, usually violet, sparsely long-hairy or sub- 

185 glabrous, with ciUate margins; calyx ca. 2 mm long, glandular-hairy; corolla 20-25 mm 
long, smaller than in S. supina, glandular-hairy outside, violet, with paler lower lip. 
June— August. 

Stony mountain slopes or meadows, steppes or sometimes alpine meadows, forests, 



123 



pebble-beds. - West Siberia: Ob. (Tomi River valley, Kuznetskii Ala-Tau), Alt.; East 
Siberia: Ang.-Say. (basin of Abakan River). Endemic. Described from Altai (from Mor- 
dovkin specimens, probably from Ridder mine). Chirotype (Fischer) in Leningrad. 

105. S. mongoiica Sobolevsk. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 49. 

Perennial; rhizome thick, nearly vertical; stems few, 16-25 cm long, erect, green, cov- 
ered v^ith fine reclinate appressed hairs interspersed with long fine spreading hairs; leaves 
1-1.5 cm long, 0.5-1 cm wide, oblong-ovate, cuneately tapering at base, glabrous on both 
sides, the petioles 0.5-1 cm long, covered with fine white spreading hairs; inflorescence 
dense; flowers 1-2 in axils of bracts, crowded, on short pubescent pedicels; calyx 2- 

3 mm long, densely glandular-hairy; corolla large, 2.5-3 cm long, dark blue, pubescent, 
upper lip slightly shorter than the lower. June. 

Sandy terraces and river valleys. - East Siberia: Ang.-Say. (Tuva ASSR). Endemic? 
Described from Kaakhem River valley near Bel'bya. Type in Tomsk. 

Note. We have not seen specimens of this plant and we rely entirely upon the author's 
description. 

106. S. irregularis Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 369. 
Perennial; root vertical, woody; rhizome decumbent, branching or strongly branching; 

stems several to fairly numerous, 8-16 cm long, ascending, usually simple, covered with 
short or mostly medium white reclinate or, in upper part, horizontally spreading hairs 
usually somewhat lilac-tinged; leaves ca. 2 cm long, 1-2 cm wide, ovate or mostly oblong- 
ovate to narrowly subrhombic, cuneately tapering at base, usually subacute, deeply incised- 
dentate with 2-4 teeth on each margin, the teeth irregular, tapering, semiovate or frequently 
oblong, obtuse, antrorse or divaricate; both sides sparsely or rather densely short- 
spreading-hairy; petioles to 1 cm long; inflorescence lax, 3-6 cm long; bracts to 1.2- 
186 1 .5 cm long, 8-10 mm wide, ovate, acute, rather densely long-villous with admixture of 
very short stipitate glands, more or less suffused with lilac; corolla rather small, ca. 2- 
2.5 cm long, yellow, hairy and glandular outside. Otherwise, similar to S. supina. 
June— July. 

Meadow and stony slopes in the subalpine meadow zone. — Centr. Asia: Dzu.-Tarb. 
(Saur Range). Endemic. Described from Upper Chegan-Obo, Taya-Bulak River. Type 
in Leningrad. 

Note. There is no doubt that this species is very closely related to the true S. supina 
L., yet distinctly leaning toward the Tien Shan S. oligodonta Juz. which, together with 
a few other rather closely aUied forms, has been segregated as the series Oligodontae Juz. 
(see below). 

107. S. knorringiae Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 366. 
Subshrub, with woody creeping caudex; stems few, 10-20 cm long, ascending at base; 

shoots covered with soft spreading or recHnate hairs, rather densely so in their upper part; 
internodes medium long; leaves large, 1 .5-3.5 cm long, 1 .2-2.5 cm wide, broadly ovate, 
cordate at base, obtuse at apex, crenate-dentate with 6-10 subequal obtuse teeth ca. 
1.5 mm high on each margin, green on both sides, covered with numerous small stipitate 
glands and long simple hairs, these more numerous on the margin and on veins beneath, 



124 



very few elsewhere ; petioles short; inflorescence in flower 2.5-4.5 cm long; bracts loose- 
ly imbricated, ca. 1 .3 cm long, 0.7 cm wide, broadly elliptic or ovate, entire or the lower 
generally with few teeth, green or slightly suffused with lilac, rather densely glandular and 
hairy; calyx at anthesis small, ca. 2.5 mm long, densely stipitate-glandular and much less 
profusely long-hairy; corolla large, to 3 cm long, copiously glandular outside, yellow or 
upper lip slightly violet-tinged. July. 

Mountain slopes and ravines. —Centr. Asia: T.Sh. (western part). Endemic. Described 
from Chotkal Range, ravine near Aflatun River, and from Kul'dambes Mountains near 
Sary-Chilek Lake. Type and paratype in Leningrad. 

108. S.xanthosiphon Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 367. 

Subshrub; rhizome and roots woody; stems few (2-5), 15-25 cm long, suberect from 
somewhat ascending base, woody in lower part, slightly curved or flexuous, simple, pale 
187 green, covered all over with numerous short as well as scattered long spreading hairs; leaves 
rather large, 1-3 cm long, 0.5-2 cm wide, ovate, approximately twice as long as wide, slight- 
ly tapering or broadly cuneate at base, obtuse at apex, green on both sides although covered 
with scattered (more numerous beneath) fine hairs; margins rather shallowly and unevenly 
dentate with 6-9 teeth on each margin, the lower teeth sometimes acute, others obtuse, 
inequilateral, with outer margin twice the length of the inner, obtuse, mostly round-tipped; 
veins prominent beneath, rather densely covered with fine hairs; lower leaves with rather 
long petioles (to 1 .5 cm), others short-petioled; vesture of petioles like that of stems, but 
the hairs longer and often more profuse; inflorescence 3.5-6 cm long, loose; bracts 1.5- 
2 cm long, 0.6-1.3 cm wide, ovate or elliptic, acute but scarcely acuminate, foHaceous, 
green, covered, more densely on the margin, with long simple soft hairs, the nerves rather 
prominent; calyx at anthesis ca. 3 mm long, rather densely hairy; corolla large, ca. 3 cm 
long, very finely hairy and glandular outside, bicolored, the tube bright yellow, the lips 
purple; lower Up medium-sized. June. 

Subalpine meadows. - Centr. Asia: T.Sh. Endemic. Described from Dzhelalabad 
region, Kugartskii sovkhoz, on the road to Urumbash. Type in Leningrad. 



Series 5. Oligodontae Juz. — Like the preceding series, but leaves with fewer (usually 
not more than 5), less developed, often flattened or obsolescent teeth. High-mountain 
forms, usually smaller than members of series Supinae Juz. 

109. S. oligodonta Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 370. - S. 
alpina var. prostrata Trautv. Enum. pi. Songar, a Schrenk collect. (1866) 134. — S. al- 
pina var. minor Rupr. in Osten-Sacken et Rupr. Sert. tiansch. in Mem. Ac. Sc. Petersb. 
VII ser. t. XIV, No. 4 (1869) 66. - S. alpina var. elegans Meinsh. in sched. - S. alpina 
var. bedelensis Krassn. in sched. (nomen) and in Spisok rast., sobr.v vost.Tyan'-Shane 
letom 1886g. (1887) 94 (diagn. sine nomine varietatis!). 

Subshrub, with vertical root and stoutish or rather slender, branching, procumbent 
woody rhizome; stems numerous, slender or sometimes fairly stout, ascending or some- 
188 times suberect, flexuous, curved, simple or sometimes strongly branching, 6-20 cm long; 
leafy shoots with long (especially in upper part) or short spreading hairs intermixed with 



125 



stipitate glands, often lilac-tinged (especially in lower part), the internodes short or long; 
leaves small or medium, 0.4-2.4 cm long, 0.2-1 .4 cm wide, ovate, widest below middle, 
rounded or, in upper leaves, angled at base, apically obtuse, rather deeply crenate-dentate 
with 1-5 teeth to 3 mm high on each margin, occasionally some leaves entire, green on 
both sides, sparsely or (especially beneath) rather densely covered with hairs and stipitate 
glands of varying size; petioles with short spreading hairs, 1.5-6 mm long; inflorescence 
3-5.5 cm long, dense; bracts ovate-elliptic or broadly ovate, the lower bracts 1-1.5 cm 
long, 0.5-1.2 cm wide, subobtuse to subacute, entire or the lower few-toothed, subherba- 
ceous, rather densely covered all over with long spreading hairs and stipitate glands, green 
or often deeply lilac; calyx at anthesis 2-3 mm long, hairy and copiously glandular; corol- 
la medium to fairly large, 2.5-3.5 cm long, densely hairy and stipitate-glandular outside, 
yellow, the upper lip and its lobes violet, the lower lip with violet markings. June— July. 
(Plate XI, Figure 2.) 

Meadow slopes, alpine meadows, placers. — Centr. Asia: Dzu.-Tarb. (Dzungaria Ala- 
Tau),T.Sh. Gen. distr.: Dzu.-Kash. Described from Aulie-Sai ravine (Dzungaria Ala- 
Tau). Type in Leningrad. 

1 10. S. paulsenii Briq. in Bot. Tidskr. 28 (1908) 233. - S. alpina var. prostrata f 
pamirica 0. Fedtsch. Flora Pamira (1903) 171. — S. filicaulis auct. plur. non Rgl. — 
Ic. (phot.): Briq. I.e. 234 (fig. 1). 

Subshrub, with woody rather slender rhizome; stems numerous, mostly rather slender, 
procumbent or ascending, flexuous, not or sHghtly branching, 6-15 cm long; shoots with 
very short appressed or somewhat spreading hairs or subglabrous, the internodes short or 
sometimes elongate; leaves small, 0.6-1 .4 cm long, 0.4-0.9 cm wide, ovate, widest below 
middle, rounded-cuneate at base, obtuse or rounded at apex, very shallowly crenate with 
1-4 teeth ca. 0.5 mm high on each side (some leaves nearly entire at times), green on both 
sides, diffusely puberulent, sometimes only on the veins beneath and on margins, or gla- 
189 brate all over, finely granular-glandular beneath; petioles short (1-2.5 mm) or leaves sub- 
sessile; inflorescences to 5 cm long, rather dense; bracts to 1.2cm long, 1 cm wide, elliptic, 
subherbaceous, entire or with isolated teeth, loosely long-villous along the margin, with 
scattered long hairs elsewhere, mostly turning deep lilac, occasionally pale green; calyx at 
anthesis small, to 2-3 mm long, loosely short-villous; corolla rather large, ca. 2.5 cm long, 
with short hairs and short stipitate glands outside, bicolored, pale yellow, the tip and lateral 
lobes of upper Up violet; nutlets unknown. June-August. 

Alpine meadows, mountain slopes, rocks, banks and dry beds of mountain streams, at 
altitudes of 3500-3700 m. Centr. Asia: Pam.-Al. (Alai valley, Pamir). Endemic. Describ- 
ed from Bordo-Aba and Kara-Su River. Type in Copenhagen, paratype in Leningrad. 

Note. As far as known, this plant was collected for the first time in 1878 by Kushake- 
vich, Skornyakov and Ashurbaev. 

111. S. filicaulis Rgl. in Opis. nov. vid. v putesh. v Turkest. A. P. Fedchenko, 18, III 
(1882)69. 

Perennial; caudex (or rhizome) rather slender, procumbent, flexuous, nodose, gray, 
branching; stems numerous, slender, subfiliform, procumbent, branching, 4-10 cm long, 
with very short spreading hairs (bloomlike pubescence visible only with a magnifying glass). 



126 



often lilac-tinged; leaves 0.3-1 .4 cm long, 0.2-1 cm wide, short-elliptic, ovate or obovate, 
rounded or slightly tapering at base, obtuse, entire or with few (1-3) shallow and asym- 
metrical crenations on each margin, green on both sides or suffused with lilac beneath, 
diffusely covered all over above, mainly along veins and margins beneath, with very fine 
short scarcely discernible hairs or subglabrous, the undersurface finely granular-glandular; 
petioles 1-10 mm long, slender, very sparingly pubescent; inflorescence 2.5-3.5 cm long, 
few-flowered (mostly 4-6 flowers); bracts 0.4-1 cm long, 2-5.5 mm wide, subherbaceous, 
elliptic, acute, entire, diffusely puberulent and with fine short stipitate glands, often lilac- 
tinged; calyx ca. 2 mm long, rather densely pubescent; corolla ca. 2.5 cm (sometimes to 
3 cm) long, short-hairy and glandular outside, with rather slender yellowish tube, the upper 
lip violet-blue, the lower Up with a spot or markings of same color; nutlets ca. 1 mm long, 
short-hairy. July -August. (Plate XI, Figure 3.) 
190 Dry beds of streams and taluses in the alpine mountain belt, near glaciers and snow at 
2500-3 100 m. — Centr. Asia: Pam.-Al. (western tip of Alai Range, Turkestan Range). 
Endemic. Described from Kavuk. Type in Tashkent, isotype in Leningrad. 

112. S. kugarti Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 371. 
Perennial; rhizome slender; stems numerous, procumbent, strongly flexuous, 6-12 cm 

long; leafy shoots softly patent-hairy and glandular, pale green or scarcely colored; leaves 
0.4-2.2 cm long, 0.2-1 cm wide, ovate or obovate, tapering at base, obtuse to subacute, 
deeply incised-crenate or crenate-dentate with 1-3 teeth on each margin (in upper leaves 
these to 5 mm high), grayish-green on both sides, densely hairy and glandular, short-petiol- 
ed; inflorescence 3-4 cm long; bracts to 1 .5 cm long, 0.8 cm wide, green, herbaceous, 
scarcely differing from cauUne leaves in consistency, shape, dentation and vesture; calyx 
3 mm long, rather sparsely hairy and glandular; corolla ca. 2.5 cm long, hairy and glandu- 
lar outside, purple-lilac, the relatively slender tube pale. July— August. 

Mountain slopes in the alpine zone. - Centr. Asia: T. Sh. (Fergana Pass). Endemic. 
Described from the Kugart region. Type in Leningrad. 

113. S. popovii Vved. sp. nov. in Addenda XIX, 347. - S. physocalyx M. Pop. in 
Sched. ad Herb. As. Med. Fasc. IX (1926) No. 202, nee aliorum. - Exs.: HFAM, No. 202. 

Perennial; flowering stems 5-8 cm long, in habit recalling most of aU S. filicaulis Rgl., 
but in vesture of stems, leaves and bracts, and in color of corolla similar to S. oligodonta 
Juz.; distinguishable from the latter by smaller, suborbicular to rounded-reniform leaves 
approximately as long as or even shorter than wide (3-9 mm by 3.5-1 1 mm), subcordate 
at base, the teeth more numerous (to 6 on each margin), smaller, rounded, both surfaces 
grayish-green, densely covered with stiffish simple hairs and rather long-stipitate glands; 
petioles 2-10 mm long; inflorescence 2.5-3.5 cm long, dense; bracts large, to 1.3 cm long, 
193 1 cm wide, broadly elUptic or oblong, attenuate at both ends, obtuse or mostly subacute 
at apex, densely covered with long hairs and long-stipitate glands, dark violet; corolla 2- 
2.5 cm long, with large lips and broad tube, yellow, with purple-tipped upper lip and pur- 
ple markings on lower Up. August. 

Loose stony placers in the alpine mountain belt. — Centr. Asia: T. Sh. (Kirghiz Ala- 
Tau). Endemic. Described from Dzhaman-Ichke Mountain (Ak-Su River). Type m 
Tashkent, isotype in Leningrad and other places. 



127 




PLATE XI. 1 - Scutellaria oxyphylla Juz., general aspect, fruiting calyx, its upper and lower parts, 
nutlet; 2 - S. oligodonta Juz., general aspect; 3 - S. filicaulis Rgl., general aspect; 4 -S. ocellata 
Juz., flower; 5 - S. karatavica Juz., summit of stem, flower; 6 - S. cordifrons Juz., general aspect. 



128 



Note. This plant was accepted in error by Popov as S. physocalyx and even distributed 
as such with the inclusion of mature calyces of that species taken from Alai specimens. 
There is no need to demonstrate that the similarity between S. popovii and S. physo- 
calyx is purely superficial. In our view, S. popovii is closely allied to S. oligodonta 
Juz., being apparently an intermediate form between the latter and S. talassica Juz. 

1 14. S. talassica Juz. in Bot. mat. gerb. Bot. inst. ANSSSR, XIV (1951) 392. 

Perennial; rhizome slender, branching; stems ca. 4 cm long, slender, flexuous, gray, 
with short spreading crisp hairs, the nodes approximate; leaves small, 3-7 mm long, 2- 
6 mm wide, ovate-rhombic, rectangular at base, incised-crenate-dentate with 2-4 obtuse 
semielliptic teeth on each margin, grayish-green or whitish-green on both sides, with short 
imperfect tomentum, the veins impressed above, prominent beneath; petioles 3-6 mm 
long, rather densely pubescent; inflorescence terminal, short, loose; flowers few; bracts 
6-8 mm long, broadly lanceolate, acute to subobtuse, few-toothed or the upper almost 
entire, densely covered with long spreading hairs interspersed with stipitate glands, pale 
violet in upper part; pedicels ca. 1 mm long; calyx 2-3 mm long, densely covered (like 
pedicels) with almost straight spreading hairs of medium length; corolla disproportionate- 
ly large (for the size of the plant and its leaves), 2.2-2.5 cm long, 6 mm wide at throat, 
with lips 6-7 mm long, apparently pink, the lower Hp with a yellow spot. August. 

Mountain slopes and ravines. - Centr. Asia: T. Sh. (Talass Ala-Tau). Endemic. De- 
scribed from Sarym-Sak ravine. Type in Leningrad. 

Note. A most distinctive, possibly hybrid form which, in some features, approaches 
certain Pulchellae Juz. of the preceding subsection. 



Series 6. Subcordatae Juz. — Subshrubs or perennial herbs, resembling species of 
194 the two preceding series (recalling Oligodontae in size and Supinae in leaf dentation), but 
vesture well developed on both leaf surfaces, consisting of somewhat crisp hairs but not 
forming a real tomentum; leaves with more or less distinctly cordate base, the strongly 
developed lower lateral veins flabellately spreading from near the base. 

115. S. linczevskii Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 373. - 
S. cordifolia var. tomentosa Lincz. in sched. 

Subshrub 8-15 cm high; root vertical, woody, caudex procumbent, strongly branching, 
flexuous, gray-brown, woody, radicant; leafy shoots numerous, densely disposed, short, 
straight or curved, not colored or faintly suffused with lilac, very densely and continuous- 
ly villous, the hairs soft, horizontally spreading, slightly crisp, of medium length (stipitate 
glands, as also on leaves and petioles, wanting or very few); leaves 0.5-1 .5 cm long, 0.3- 
1 cm wide, ovate or broadly ovate, with cordate (especially in lower leaves) or truncate 
base, obtuse or rounded at apex, evenly finely and shallowly crenate with 4-8 rounded 
teeth on each margin, green or grayish-green on both sides, densely covered with spread- 
ing, slightly crisp or nearly erect hairs of medium length, veins flabellately spreading, 
usually impressed above, prominent beneath and densely patent-hairy; petioles patent- 
villous, 1-8 mm long, or upper leaves subsessile; inflorescence 2-3.5 cm long, rather dense; 
bracts to 1-1.5 cm long, 5-9 mm wide, ovate-elliptic, tapering at base, acuminate at apex. 



129 



almost flat, with rather prominent longitudinal nerves, suffused with violet, rather densely 
covered with long spreading hairs interspersed with long-stipitate glands; calyx at an thesis 
ca. 3 mm long, villous; corolla rather small, 1.5-2.5 cm long, with broad tube, rather dense- 
ly hairy and glandular outside, yellow, the lower lip and the tip and lateral lobes of upper 
lip sometimes colored; nutlets so far unknown. June— August. 

Stony slopes in subalpine zone, alpine grassy areas. — Centr. Asia: T. Sh. (western part, 
Talass Ala-Tau in particular). Endemic. Described from upper reaches of Kshi-Kainda 
River. Type in Leningrad. 

116. S. subcordata Juz. sp. nov. in Addenda XIX, 348. 
195 Subshrub 8-20 cm high, similar in habit to S. linczevskii Juz.; leafy shoots less dense- 
ly covered with relatively short curved or crisp hairs, sometimes almost tomentose, usually 
not anthocyanin-colored; leaves 1-2.5 cm long, 0.4-2.2 (2.6) cm wide, ovate to suborbi- 
cular, at least the lower subcordate, others mostly obtuse or broadly cuneate at base, ob- 
tuse or rounded at apex, sometimes acute, coarsely crenate-dentate with 5-8 unequal ob- 
tuse or rounded teeth or crenations on each margin, both sides green, with scattered short 
hairs, these crisp beneath, straighter and almost accumbent above; lower lateral veins most- 
ly arcuately divergent from base, moderately impressed above and prominent beneath; 
petioles 1 .5-12 mm long, softly crisp-hairy; inflorescence 3-4.5 cm long, loose; bracts 
1-1 .5 cm long, 0.4-0.8 cm wide, ovate, obtuse, almost flat, with inconspicuous longitudinal 
nerves, rather sparsely covered with long spreading hairs interspersed with stipitate glands, 
green, subherbaceous; calyx at anthesis ca. 2.5 mm long, with long soft hairs; corolla 2.2- 
2.7 cm long, rather sparingly long-stipitate-glandular outside, with rather narrow tube, 
yellow with dark spot on lower lip. July. 

Rocks and gravelly places in subalpine and alpine mountain belts. — Centr. Asia: T. Sh. 
(western part, Kirghiz Ala-Tau at its western tip). Endemic. Described from Taldy-Bulak 
(opposite Akyr-Tyube). Type in Tashkent. 



Series 7. Subcaespitosae Juz. — Like the preceding series, but leaves broadly cuneate 
at base; lower lateral veins weakly developed, hence venation essentially pinnate. 

1 17. S. subcaespitosa Pavl. in Tr. Sredneaz. Gos. univ. ser. VIII-C (botany), 19 
(1935)33. 

Perennial; caudices numerous, congested, procumbent, flexuous, cespitose, persistent, 
woody radicant; annotinous stems ascending, rarely erect, low, 5-8 cm long, rather dense- 
ly patent-hairy, remotely glandular above; leaves 1 1-20 mm long, 10-15 mm wide, ovate, 
broadly cuneate at base, obtuse at apex, crenate-dentate with 5-7 large obtuse teeth on 
each margin, covered on both sides, especially on the veins beneath, with short spreading 
hairs; petioles patent-hairy, as long as or slightly shorter than blade; inflorescence 2-5 cm 
196 long, rather dense, 3-10-flowered; bracts oblong-ovate, scarious, finally almost flat, entire 
or with few teeth, sparsely short-stipitate-glandular and long-hairy, pale green or often 
slightly lilac-tinged ; calyx small, short-glandular, sparsely hairy; scutellum large, rounded, 
glabrous, netted-veined; corolla large, 2-3 cm long with long narrow tube, yellow, the 
lower lip deeply colored and usually shorter than the upper; nutlets ca. 1 mm long, cine- 
reous, with short stellate hairs. June. 



130 



Stony slopes. - Centr. Asia: T. Sh. (eastern part of Kara-Tau mountain system). 
Endemic. Described from Mashat-Tau mountains near Tyul'kubass railway station. Type 
in Moscow. 

Economic importance. A dye (water extract dyes fabric pale lemon yellow). 

1 18. S. lanipes Juz. sp. nov. in Addenda XIX, 349. 

Perennial; very similar to S. subcaespitosaPavl.; differing in growth habit and ves- 
ture; caudex ramifications rather few, elongate, hence plant not cespitose; annotinous 
shoots (stems) suberect, ca. 10 cm long, very densely long-patent-hairy, almost villous; 
leaves ovate or broad-ovate, with obtuse or slightly cordate base, grayish-green on both 
sides with very profuse slightly crisp hairs, beneath almost (imperfectly) tomentose; pe- 
tioles and young leaves densely long-villous-lanate-hairy ; bracts commonly to 1 cm long, 
ovate-elliptic or rather broadly ovate, acute, almost villous; corolla unknown. Otherwise, 
Uke S. subcaespitosa Pavl. June. 

Mountain slopes, stony substratum. — Centr. Asia: T. Sh. (western part: Ichkele-Tau 
mountains). Endemic. Described from Ichkele-Tau mountains. Type in Tashkent. 

Note. A little known species, described from insufficient and unsatisfactory material. 

119. S. toguztoraviensis Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 375. 
Perennial; rhizome woody, rather stout, pale brown, nodose, branching; leafy stems 

7-13 cm long, few, ascending or suberect, densely covered with long horizontally spreading 
hairs, not colored; leaves 0.6-1 .7 cm long, 0.5-1 .2 cm wide, usually longer than internodes, 
197 ovate, with obtusely angled base, obtuse at apex, the margin with 4-7 angled teeth of medi- 
um height, the upper surface darkish green or grayish-green, densely covered with long ap- 
pressed hairs, the lower surface grayish-green, rather densely patent-hairy, the hairs shorter, 
partly slightly crisp, partly straight; petioles densely long-patent-hairy, to 1cm long; inflo- 
rescence 3-4.5 cm long (in full flower), rather dense; bracts to 1.7 cm long, 1 cm wide, 
elUptic, short-attenuate at both ends, obtuse at apex, almost flat, rather densely covered 
with simple longish spreading hairs and numerous short-stipitate glands, slightly lilac-tinged; 
calyx small, long-hairy and densely stipitate-glandular; corolla large, 2.5-3 cm long, bright 
yellow, copiously covered outside with short-stipitate glands, the broad tube slightly 
curved, the lower Up large, often exceeding the upper, the tip and lateral lobes of upper 
lip sometimes purple; nutlets unknown. June. 

Mountain slopes. - Centr. Asia: T. Sh. (central). Endemic. Described from natural 
boundary Toguz-Torau, near Oikain mountain pass. Type in Leningrad. 

120. S. aiexeenkoi Juz. sp. nov. in Addenda XIX, 349. 

Subshrub, with rather sturdy, strongly branching caudex; stems numerous, stout, as- 
cending and arcuately or even geniculately curved, mostly simple, 8-20 cm long; shoots 
densely covered with horizontally spreading short and long hairs, the latter very copious, 
soft and slightly flexuous; internodes short or medium, leaves rather large, to 4 cm long, 
to 2.5 cm wide, ovate or broadly ovate, with truncate or sometimes subcordate base, ob- 
tuse at apex, shallowly and finely crenate-dentate with 5-8 teeth on each margin, green on 
both sides, more or less densely short-hairy; petioles 0.5-1.2 cm long; inflorescence ca. 
4-5, sometimes to 8 cm long, rather dense; bracts to 1 .8 cm long, to 1 cm wide, oblong- 
ovate, persistent, subherbaceous, entire or with soUtary teeth, rather densely covered with 



131 



long and short spreading hairs intermixed with stipitate glands, pale green; calyx at anthe- 
sis small, to 3 mm long, villous-hairy and glandular, in fruit to 1 cm long including the sub- 
glabrous scutellum; corolla large, ca. 3 cm long, short-hairy and short-stipitate-glandular 
outside, with rather slender tube, essentially yellow, the tip and lateral lobes of upper lip 
198 purple, the lower hp with violet marking (often two spots); nutlets angled-ovoid, ca. 
1 .5 mm long, bluish, densely covered all over with stellate hairs. June. 

Grassy mountain slopes. — Centr. Asia: Pam.-Al. (foothills of Alai Range). Endemic. 
Described from Lyangar along Taldyk River and from Shakhimardan. Type in Leningrad, 
paratype in Tashkent. 

Note. The type of this species is bereft of flowers and hence their description is based 
on a much better specimen from Shakhimardan. In our interpretation, S. alexeenkoi 
links S. xanthosiphon Juz., the primary species of the entire series Subcaespitosae, with 
S. ocellata Juz., and for this reason we have included it in this series. The vesture of its 
leaves, however, differs slightly as compared with other species of the series. 

121. S. ocellata Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 376. 

Subshrub, 12-25 cm high, with woody branching long-creeping caudex; stems numer- 
ous, flexuous, ascending at base and rather densely covered, like petioles, with long soft 
crisp spreading hairs, the internodes often elongate; leaves medium to fairly large, 12- 
25 mm long, 7-17 mm wide, ovate, mostly truncate at base, coarsely and irregularly cre- 
nate-dentate with 4-6 large and rather high straight obtuse teeth (to 4 mm long) on each 
margin, green on both sides, densely covered (especially on the veins beneath) with long 
simple crisp hairs; petioles 6-18 mm long; inflorescence 5-6 cm long, usually loose;bracts 
ovate to broad-ovate, the lower ca. 1 5 mm long, 7 mm wide, aU entire or the lower toothed, 
flat, subacute to subobtuse, rather densely covered with long hairs interspersed with few 
short stipitate glands, green or mostly suffused with purple, loosely imbricate; calyx at 
anthesis medium-sized, ca. 3 mm long, densely hairy, corolla large, to 3.3 cm long, remote- 
ly glandular outside, yellow, the upper lip purple-tipped, the lower brownish with 2 large 
round dark brown spots. May. (Plate XI, Figure 4.) 

Habitat unknown. — Centr. Asia: Pam.-Al. (Zeravshan). Endemic. Described from 
Naufin-Kul' Lake. Type in Leningrad. 



Series 8. Karatavicae Juz. — In the shape and venation of the leaves, the plant recalls 
species of the series Subcordatae, but is much more xerophytic; leaves subcoriaceous but 
very brittle when dry; bracts closely contiguous. 

122. S. karatavica Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 372. 
199 Perennial, with long slender flexuous branching rhizome; stems procumbent, ascending, 
slightly flexuous or curved, 10-30 cm long, branching from base, densely short-villous, 
scarcely colored or more or less lilac-tinged; leaves 0.8-2 cm long, 0.6-2 cm wide, stiff, 
very broadly ovate or suborbicular, subcordate or rounded at base, obtuse at apex, with 
2-6 large rounded crenations on each margin, grayish-green on both sides, densely covered 
with short spreading velutinous hairs, midrib and lateral veins impressed above, prominent 
beneath; petioles thickish, to 7mm long; inflorescence 2.5-3.5 cm long, very dense; 



132 



bracts to 1.5 cm long, 1 cm wide, broadly ovate, tapering at base, long-acuminate at apex, 
subherbaceous, firm, distally lilac-tinged, densely covered, especially along margins, with 
long simple white hairs, the parallel nerves very prominent; corolla ca. 1 .5 cm long, yellow, 
sparsely hairy outside, with short tube, short hood and large lower lip; nutlets unknown. 
June. (Plate XI, Figure 5.) 

Mountain slopes. — Centr. Asia: T. Sh. (Kara Tau Range). Endemic. Described from 
Berk-Kara ravine. Type in Leningrad. 

Note. This interesting species differs so strikingly from all the other species of the sub- 
section that we were able to segregate it as a series. Apparently it is phylogenetically 
closely allied to species of the series Subcordatae Juz. 



Subgenus 2. Cystaspis Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1952) 413 
(pro sect.). — Perennials, recalling in habit some species of the section Lupulinaria A. 
Hamilt. of the preceding subgenus (especially those of the subsection Alpinae and the 
affined S. oligodonta Juz. and others). They differ notably in that the scutellum on 
the upper (caducous) part of the calyx is replaced by a vesicular-sacciform, membranous 
appendage that expands markedly in fruit. This appendage encloses the nutlets and un- 
doubtedly aids in their dispersal (apparently by wind and perhaps also by water of 
streams). 

A Central Asian group. 

Note. Only two species, S. physocalyx Rgl. et Schmalh. and S. jodudiana B. 
Fedtsch., were distinguished within this group prior to our investigation. However, the 
attempts by different authorities (and collectors) to account for all the Central Asian ma- 
terial of the Cystaspis group were unsuccessful since it was virtually impossible to assign 
some of these forms to one or the other species. We have attempted a more thorough sub- 
200 division of the group, stressing in greater detail the characteristics distinguishing these spe- 
cies. But the material at our disposal has been insufficient, and the whole group is in need 
of further study. 

All the species of the subgenus Cystaspis so far known apparently form a single series 
which can be named Physocalyces Juz. (its characteristics concur with those of the sub- 
genus). 

1. Stems and leaves densely villous with long hairs .... 127. S. poly tricha Juz. 
+ Vesture different 2. 

2. Corolla rather large, usually 1 .8-2.5 cm long, its predominant color yellow or whit- 
ish; appendage of upper calyx Up rather large, to 1 1 mm in diameter 3. 

+ Corolla usually smaller, 2-1 .5 (2) cm long, its predominant color lilac; appendage of 
upper calyx lip usually not more than 8 mm in diameter 6. 

3. Nearly all bracts with few large marginal teeth; flowers rather small (to 1 .8 cm long) 
130. S. schachristanica Juz. 

+ Bracts usually entire, only the lowermost sometimes toothed at base; flowers 

larger 4. 

4. Plants with vertical root and decumbent stems 10-45 cm high; inflorescence rather 
large, 2.5-6 cm long 5. 



133 



+ Plants with long slender branching rhizome and stems 3.5-10 cm high; inflorescence 
ca. 2 cm long 125. S. pamirica Juz. 

5. Leaves broadly ovate to suborbicular, with truncate or subcordate base; corolla yel- 
low, with violet upper lip 123. S. physocalyx Rgl. et Schmalh. 

+ Leaves generally ovate to broad-ovate, tapering at base; corolla whitish, the tip and 
lateral lobes of upper lip purple . . . 124. S.anitaeJuz. 

6. Bracts rather narrow, ovate or narrowly rhombic, usually sparsely short-hairy, only 

at margins densely long-ciliate; corolla relatively large, 1.5-2 cm long 

126. S. microphysa Juz. 

+ Bracts broad-ovate or eUiptic, villous all over with long spreading hairs; corolla 

small, 1.2-1.5 mm long 7. 

7. Leaves with usually 3 teeth on each margin; corolla violet-purple, with pale tube . 
128. S.jodudianaB.Fedtsch. 

201 + Leaves with usually 5 (often to 8) teeth on each margin; upper lip of corolla and its 
appendages very dark purple, lower lip with a yellow spot at base, the tube yellow. 
129. S.darvasica Juz. 



123. S. physocalyx Rgl. et Schmalh. in E. Kegel', Opis. nov. vid v putesh. v Turkest. 
A. P. Fedchenko, 18, III (1882) 68. - Exs.: HFAM, No. 202 (solum calyces fructiferi, 
neque ipsa planta!). 

Perennial; root woody, erect, branching; stems decumbent, flexuous, 10-45 cm long, 
woody in lower part, sparsely covered with short spreading hairs, more or less suffused 
with lilac; leaves 0.6-1.7 cm long, 0.5-1 .5 cm wide, broadly ovate, triangular or some- 
times suborbicular, widest at truncate or sometimes subcordate base, obtuse or acute at 
apex, incised-crenate or sometimes nearly crenate-lobate, with 2-6 large irregular obtuse 
crenae on each margin, grayish-lanate on both sides with rather short hairs, these fewer, 
appressed above, more copious and divergent beneath, on the veins patent; upper surface 
rugose, with impressed veins, the lower with somewhat prominent veins; petioles 2.5- 
10mm long, with short spreading hairs; inflorescence 2.5-6 cm long, dense, subcapitate; 
bracts 0.8-1 .5 cm long, 0.5-1 .2 cm wide, much longer than calyx, broadly ovate, eUiptic or 
the upper oblong-eUiptic, short- or long-acuminate at apex, the lower crenate at base, the 
upper entire, green or usually lilac, densely covered on the back with simple crisp hairs 
of medium length, the margin densely patent-hairy; calyx at anthesis ca. 3 mm long, 
densely villous with long white hairs, its upper part inflated into an ovoid-globular calcei- 
form sac, in fruit ca. 1 cm in diameter, much larger than the lower part, turning lilac; 
corolla 2-2.5 cm long, broad at throat, half as long again as bracts, apparently yellow, with 
violet helmet, hairy and sparsely short-stipitate-glandular; nutlets ca. 2 mm long, ovoid, 
dark brown or nearly black, sparsely or remotely stellate-hairy. July-August. 

Gravelly slopes, pebbles (ca. 2480 m altitude). - Centr. Asia: Pam.-Al. (Alai Range 
and Alai valley). Endemic. Described from Alai Range (from 0. Fedchenko collections). 
Type in Tashkent, isotype in Leningrad. 

124. S. anitae Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 413. 

202 Perennial, stems 6-25 cm long, very strongly flexuous, sparingly pubescent, turning 



134 



(203) 




PLATE XII. 1 - Scutellaria jodudiana Fedtsch., general aspect, leaves, bracts, corolla; 2 - S. ani- 
tae Juz., summit of stem, lower bract, flowers, fruiting calyx, nutlet; 3 - S. schachristanica Juz., 
cauline leaf, bracts, flower; 4 - S. pamirica Juz., general aspect, cauline leaf, bracts, flower, fruiting 
calyx. 



135 



lilac; leaves 4-lOmm long, 2.5-7 mm wide, ovate or broad-ovate, with 1-4 shallow crenae 
on each margin; petioles to 7 mm long; inflorescence 2-4 cm long; bracts to 1 cm long, 
3-6 mm wide, oblong-ovate, tapering at base, gradually long-acuminate at apex, densely 
covered all over with simple long straight spreading hairs, usually lilac; sacciform appen- 
dage on upper part of calyx 6-11 mm in diameter, pale green or sHghtly lilac-tinged, patent- 
hairy; corolla 2-2.3 cm long, white, partially purple-nerved, the tip and lateral lobes of 
upper lip purple, the lower lip (sometimes also purple) with a yellow rim or spot; nutlets 
ca. 2 mm long, blackish, densely white-punctate with stellate hairs. Other characters as in 
S. physocalyx Rgl. et Schmalh. August. (Plate XII, Figure 2.) 

Gravelly deluvial slopes, stony placers in alpine mountain belt. — Centr. Asia: Pam.-Al. 
(Gissar Range). Endemic. Described from southern slope of Gissar Range, basin of Sarai- 
Mion River, upper reaches of Kafandar River. Type in Leningrad. 

125. S. pamirica Juz. in Bot. mat. gerb. Bot. inst. ANSSSR, XIV (1951) 417. 
Perennial, with slender woody root; rhizome long, slender, branching, flexuous; stems 

low, slender, leafy, 3.5-10 cm long, with short internodes, densely covered with short 
spreading white hairs, more or less suffused with lilac; leaves in few pairs, 2.5-8 mm long, 
2-6.5 mm wide, ovate, with truncate or obtusely angled base, subobtuse, with 2-4 small 
superficial crenae on each margin, grayish on both sides, with dense imperfect tomentum; 
petioles densely short-hairy, to 7 mm long; inflorescence ca. 2 cm long; lower bracts ca. 
1 cm long, 7 mm wide, ovate-rhombic, tapering at both ends, subacute, densely short- 
hairy, with long-white-ciliate margins, turning lilac; calyx rather sparingly pubescent, the 
appendage of its upper part ca. 7 mm in diameter in fruit, slightly lilac; corolla large for 
205 size of plant, ca. 2 cm long, apparently yellow with lilac tip and lobes of upper lip, strongly 
pubescent outside; nutlets ca. 1.5 mm in diameter, blackish, sparsely covered with white 
stellate hairs. July-August. (Plate XII, Figure 4.) 

Stony placers. — Centr. Asia: Pam.-Al. (Shugnan, W. Pamir). Endemic. Described 
from Yamg district and elevations on Ak-Tailakh. Type and paratype in Leningrad. 

126. S. microphysa Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 414. - 
Ic: Yuz., op. cit. 415, Figure 4. 

Perennial; stems 7-30 cm long, slightly flexuous or subgeniculately curved, sparsely or 
densely pubescent, greenish or frequently lilac; leaves 2.5-10 mm long, 1 .5-10 mm wide, 
broadly ovate to rhombic, shallowly or even superficially crenate with 2-6 crenae on each 
margin, grayish on both sides, with contiguous tomentum; petioles to 5 mm long; inflo- 
rescence 2.5-4.5 cm long; bracts small, 7-10 mm long, 3-6 mm wide, ovate to narrowly 
rhombic, tapering at base, gradually acuminate at apex, densely or usually sparsely short- 
crisp-hairy, with densely long-white-ciliate margins, usually dark lilac; sacciform appendage 
on upper part of calyx rather small, 6-8 mm in diameter, lilac, densely patent-hairy; corol- 
la smallish, 1 .5-2 cm long, ca. 6-7 mm in diameter at throat, dark purple, with white tube; 
nutlets ca. 1.5 mm long, very finely and sparsely stellate-hairy. Otherwise resembling 
S. physocalyx Rgl. et Schmalh. and S. anitae Juz. July-August. 

Stony taluses in alpine zone. — Centr. Asia: Pam.-Al. (Zeravshan valley). Endemic. 
Described from Langlif. Type in Leningrad. 



136 



127. S. polytricha Juz. in Bot. mat. gerb. Bot. inst. ANSSSR, XIV (1951) 416. 
Perennial, with woody root; rhizome long, procumbent, strongly branching, flexuous; 

leafy stems 6-1 8 cm long, slender, ascending, very densely long-patent-hairy, greenish or 
faintly lilac; leaves 3-10 mm long, 2.5-10 mm wide, semiorbicular-triangular, with trun- 
cate base, obtuse or rounded at apex, with 3-5 rounded crenae on each margin, yellowish- 
grayish on both sides, densely villous-tomentose, with spreading hairs, rugose; petioles 
densely covered with long horizontally spreading hairs, to 6 mm long; inflorescence ca. 
2.5-4 cm long, dense; lowermost bracts ca. 1 cm long, 6 mm wide; all bracts ovate, long- 
acuminate, scarious, with prominent longitudinal nerves, pale green or faintly lilac, densely 
covered all over with long white hairs; lower bracts few-toothed; appendage on upper 

206 part of calyx 8-10 mm in diameter in fruit, pale green, densely long-villous; corolla to 
2 cm long, with dull pink upper Up and yellow throat (according to collector's note), 
densely hairy and glandular outside; nutlets ca. 1.5 mm long, minutely punctate, densely 
covered all over with very fine hairs. August-September. 

Stony places (limestone). — Centr. Asia: Pam.-Al. (Tadzhikistan). Endemic. Described 
from Gendedere ravine (between Bal'dzhuan and Karategin). Type in Leningrad. 

128. S. jodudiana B. Fedtsch. in 0. and B. Fedch. Perech. rast. Turk. V (19 13) 158. - 
S. phy socalyx var. jodudiana M. Pop. in sched. 

Perennial with woody root and strongly branching rhizome; stems rather numerous, 
procumbent or ascending, slender, often filiform, herbaceous, slightly flexuous, 5-20 cm 
long, covered with short fine horizontally spreading white hairs, often faintly lilac; leaves 
3.5-17 mm long, 3-15 mm wide, shorter or longer than internodes, semiorbicular to broad- 
ly ovate, with subcordate or truncate base, rounded or obtuse at apex, shallowly crenate- 
dentate with 1-4 (usually 3) large irregular rounded crenae on each margin, grayish-green 
on both sides, with short slightly crisp white hairs, these appressed above, more or less 
spreading beneath, the upper surface slightly rugose, the veins not or slightly prominent 
beneath; petioles covered with short spreading hairs, to 1.4 cm long; inflorescence a 
4-angled spike, 2.5-5 cm long, many-flowered, dense or rather lax, at the ends of some of 
the branches; bracts 9-12 mm long, 3.5-9 mm wide, greatly exceeding calyx, broadly 
ovate to oblong-elliptic, tapering toward base, short-acuminate at apex, the lowermost 
subfoHaceous, others scarious, entire, white-villous with long spreading hairs, usually lilac- 
tinged, very rarely pale green; calyx ca. 3 mm long at anthesis, villous with long white 
hairs, the vesicular appendage on its upper part reaching 6-9 mm in diameter, strongly 
accrescent after flowering, often suffused with lilac, the longitudinal nerves prominent, 
palmately divergent; corolla small, 1 .2-1 .5 cm long, 3-5 mm wide at throat, often scarce- 
ly exceeding bracts, violet-purple, with whitish or yellowish tube; nutlets puncticulate, 
with rather scattered fine stellate hairs, sometimes subglabrous. June -August. (Plate XII, 
Figure 1.) 

Taluses, stony slopes, rock crevices in high-mountain belt. — Centr. Asia: Pam.-Al. 

207 (S. Tadzhikistan). Endemic. Described from Roshan, lodudi gorge on territory of Kala-i- 
Vamar. Type in Leningrad. 

129. S. darvasica Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951)418. 
Perennial; similarto S. jodudiana B. Fedtsch., but stems somewhat stouter and tougher. 



137 



ascending or suberect; leaves 4-10 mm long and wide, with 4-8 (usually 5) smaller crenae 
on each margin, more densely pubescent, grayish, firmer, strongly rugose, with very pro- 
minent veins beneath; petioles to 6 mm long; inflorescence 2-5 cm long; bracts and calyx 
very densely villous; vesicular appendage on upper part of calyx ca. 5-8 mm across in fruit; 
corolla ca. 1 .5 cm long, with yellow tube, the upper lip and its appendages very dark purple 
(almost black), the lower lip purple with a yellow spot at base. In all other characters like 
S. jodudiana B. Fedtsch. July-August. 

Alpine mountain belt. — Centr. Asia: Pam.-Al. (Tadzhikistan). Described from north- 
ern slope of Darvaz Range, Gul'bed valley. Type in Leningrad. 

130. S. schachristanica Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 419. 

Perennial; like S. darvasica Juz., but leaves more deeply incised-crenate, the grayish- 
green lower surface with more spreading hairs; bracts thinner, long-acuminate, pale green 
or faintly colored only in upper part, nearly all bracts with large marginal teeth, the longi- 
tudinal nerves prominent in dry state; corolla slightly larger, to 1 .8 cm long, yellow, with 
lilac-tipped upper lip. July-August. (Plate XII, Figure 3.) 

Mountain slopes (upper limit of Central Asian juniper zone). — Centr. Asia: Pam.-Al. 
(Turkestan Range). Endemic. Described from northern slope of Turkestan Range, Sha- 
khristan mountain pass. Type in Leningrad. 



Subgenus 3. Anaspis [Reching. fil. in Notizbl. Bot. Gart. u. Mus. Berlin-Dahlem, 15 
(1941) 630-632, pro gen.] Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 420, 
pro sect. — Flowers in one-sided (dorsiventral) racemose inflorescence; cauHne leaves 
gradually passing into bracts, these leaflike but much smaller, entire, short-petioled or 
subsessile; calyx accrescent, scarious or coriaceous, the upper part unappendaged but with 
distinct bulge at the middle, the lower part without a bulge, both parts falling away after 
208 ripening of fruit. Brittle, herbaceous or suffruticose plants, with thick leaves, varying in 
vesture; usually growing on precipitous rocks and in their crevices. 

A Central Asian group. 

Note. We have not segregated this group into a genus, as proposed by Rechinger, be- 
cause we are convinced of its close alHance with the subgenus Euscutellaria (and in 
particular with the section Stachymacris A. Hamilt.). The presence of a rudiment of 
scutellum on the upper lip of the calyx in species of the Anaspis group provides support- 
ing evidence. 

All the known species of this subgenus apparently vicariate with each other to form a 
specific series although, as regards the character of the inflorescence, they may be divided 
into two groups, the leading representatives being S. hissarica B. Fedtsch. and S. schu- 
gnanica B. Fedtsch., respectively. 

1 . Flowers in distant verticillasters forming very lax elongate inflorescences; both lips 
of (mature) calyx much longer than wide 2. 

+ Flowers in closely approximate verticillasters forming abbreviated inflorescences; 

lips of calyx as long as or slightly longer than wide 5. 

2. Whole plant glabrous 131. S. fedtschenkoi Bornm. 



138 



+ At least calyx hairy all over 3. 

3. Whole plant densely covered with medium or short, spreading, straight and crisp 

hairs, seemingly velutinous (especially lewder side of leaves) 

133. S. velutina Juz. et Vved. 

+ Plants with different vesture 4. 

4. Stems glabrous or with a very diffuse bloomlike coating of minute, almost papilli- 
form hairs 132. S. hissarica B. Fedtsch. 

+ Stems densely pubescent all over 5. 

5. Pubescence bloomlike, consisting of minute papilliform hairs 

134. S. nevskii Juz. et Vved. 

+ Pubescence containing papilliform as well as long simple hairs 

135. S.heterotricha Juz. et Vved. 

6. Stems glabrous or with scattered hairs 7. 

+ Stems, like the whole plant, densely hairy 8. 

7. Leaves sparsely or densely covered on both sides with short fine hairs 

136. S. schugnanica B. Fedtsch. 

+ Leaves densely covered on both sides with appressed gladiate somewhat crisp hairs 
of medium length 137. S. lipskyi Juz. 

8. Plant villous with long multicellular somewhat crisp hairs interspersed with short- 
stipitate glands 138. S. villosissima Gontsch. 

+ Plant velutinous with short or medium hairs . . . .139. S. holosericea Gontsch. 

131. S. fedtschenkoi Bornm. in Allgem. Bot. Zeitschr. Nos. 1-2 (1914) 8 (nomen); 
Beih z. Bot. Centralbl. XXXVI, 2 (1918) 60 (descr.). - S. hissarica var. fedtschenkoi 
M. Pop. in Bot. mat. gerb. Bot. sada, V, 10 (1924) 154, cum var. glabrescens M. Pop. 
1. c. — S. hissarica var. glabra M. Pop. in sched. — Anaspis fedtschenkoi Reching. fil. 
in Notizbl. Bot. Gart. u. Mus. Berlin-Dahlem, 15 (1941) 630-632 (non vidi). - Ic: Rech- 
ing. fil. 1. c. 

Perennial; caudex woody, branching; stems many, (30) 35-45 (50) cm high, slender, 
almost virgate, stiff (stems of previous year often persisting), obscurely 4-angled to sub- 
terete, completely glabrous, glaucescent, leafy, divaricately branching from middle, the 
branches slender, elongate, flexuous or pendulous, small-leaved, passing into inflorescence; 
lower cauline leaves 3-4 pairs, 2-3.5 cm long, 2-3 cm wide, approximately half the length 
of internodes (the latter 4-6 cm long), ovate, with shallowly cordate base, obtuse, with 2 
or 3 large obtuse teeth on each margin (the very large terminal tooth often lobeHke), or 
sometimes entire, glabrous, green, dull on both sides, the undersurface with a bloomlike 
coating of small scattered sessile glands; petioles glabrous, 5-18 mm long; upper cauline 
leaves (above insertion of first branches) entire; all leaves thick, flat, veins not prominent; 
leaves of lateral branches much smaller, orbicular, ovate or oblong-ovate to lanceolate, 
entire, subsessile, very distant, gradually passing into bracts; flowers in very loose one- 
sided few-flowered inflorescence consisting of 4-6 distant verticillasters, on pedicels ca. 
1 mm long, horizontally spreading, as long as or shorter than internodes of inflorescence; 
calyx 4.5 mm long, 3-3.5 mm wide at flowering, campanulate, generally sparingly pubes- 
cent only at margin, otherwise with scattered sessile glands, in fruit 6.5 mm long, 4.5 mm 
wide, with short broad arched or truncate Ups; corolla 12 mm long, more than twice the 



139 



length of calyx, blue, the straight tube gradually expanding into a short 2-lipped limb. 
August. 

In crevices of shady precipitous rocks. — Centr. Asia: Pam.-Al. (southern slope of 
210 western part of Gissar Range, Baisun province). Endemic. Described from a ravine be- 
tween Derbend and Akrabat. Type in Weimar, isotype in Leningrad. 

132. S. hissarica B. Fedtsch. in O. and B. Fedch. Perech. rast. Turk. V (1913) 159. - 
S. hissarica var. typica M. Pop. in Bot. mat. gerb. Bot. sada, V(1924) 153. — Anaspis 
hissarica Juz. hoc loco (nom. alternat.). 

Perennial; caudex woody, strongly branching at summit, stems numerous, 30-50 cm 
long, rather slender, obscurely 4-angled, almost virgate, green, glabrous or sometimes with 
barely discernible bloom of minute almost papilliform hairs, only in inflorescence distinct- 
ly covered with short horizontally spreading hairs intermixed with stipitate glands, simple 
or in upper part branching, the suberect branches varying in length; cauline leaves 5-6 
pairs, 1.5-5.5 cm long, 1-4.5 cm wide, 1/3 to 1/2 as long as internodes (these 6-11 cm 
long), ovate, broadly ovate or nearly triangular-ovate, with sHghtly cordate base, obtuse; 
lower leaves sometimes subentire, mostly with 1-5 teeth on each margin, these large, low 
and broad, arcuate, crenate; middle and lower leaves of lateral shoots usually obscurely 
crenate, rounded at apex, short-petioled; upper leaves entire; all leaves thick, glabrous on 
both sides or covered on the veins beneath with scattered, minute subpapilliform spreading 
hairs, the veins prominent beneath; petioles 0.6-1 .5 cm long; flowers in very loose one- 
sided, few-flowered inflorescence of 6-12 remote verticillasters (pairs of flowers), short- 
pediceled; bracts small, 2-5 mm long, in fruit to 14 mm, 1/3 to 1/2 the length of calyx, 
broad-ovate or ovate, entire, obtuse, usually densely covered with very fine short appres- 
sed hairs, long-ciHate on margins, short-petioled or sessile; calyx at anthesis 3-4 mm long, 
2.5-3 mm wide, in fruit to 8 mm long, 6 mm wide, densely covered with appressed or 
spreading fine hairs, often with some long stiff hairs and sessile glands, the lips very short, 
arcuate; corolla 1.7-2.2 cm long, more than twice as long as calyx, glandular and hairy 
outside, with slightly curved tube gradually expanding into a short 2-lipped limb, dark 
lilac, the lower lip pale; nutlets papillate-rugose. May-July. (Plate XIII, Figure 1.) 

Crevices of precipitous rocks. — Centr. Asia: Pam.-Al. (Gissar). Endemic. Described 
from gorge of Karatag River near Khakimi. Type in Leningrad. 



211 



133. S. veiutina Juz. et Vved. in Addenda XIX, 350.— Anaspis velutina Juz. hoc 
loco (nom. alternat.). 

Perennial, very close to S. hissarica B. Fedtsch., but readily distinguished from the 
latter by the characteristic copious vesture of its parts, consisting of soft short spreading, 
both straight and crisp hairs, especially dense whitish-velutinous on underside of leaves; 
inflorescence erect or slightly curved, elongate; verticillasters distant, 0.5-2.5 cm apart; 
calyx with vesture of two components: shorter hairs as on other parts of the plant and 
less profuse long hairs. Otherwise, like S. hissarica B. Fedtsch. June. 

Apparently on rocks (habitat not recorded by collector). — Centr. Asia: Pam.-Al. 
(Gissar Range). Endemic. Described from Varzob River valley. Type (collected by 
V. Titov in 1930) in Tashkent. 

Note. In its vesture, this plant vividly recalls S. holosericea Gontsch. (see below) 



140 



from which, however, it differs strikingly in the remote verticillasters of its inflorescence, 
and this feature indicates kinship to S. hissarica B. Fedtsch. rather than S. schugnani- 
ca B. Fedtsch. 

134. S. nevskii Juz. et Vved. in Bot. mat. gerb. Bot. inst. ANSSSR, XIV (1951) 424. - 
S. hissarica Nevski in Tr. Bot. inst. AN SSSR, I, 4 (1937) 322, non B. Fedtsch. s. str. - 
Anaspis nevskii Juz. hoc loco (nom. altern.). 

Perennial, with woody branching caudex; stems few, 12-25 (40) cm long, subterete, 
glaucescent, with a dense bloomUke coating of minute papilHform hairs, these more pro- 
fuse in inflorescence where color of stem usually turns dark violet; cauline leaves 3-4 
pairs, 1-3.5 cm long, 0.5-2 cm wide, much shorter than internodes (usually 1/3 as long), 
ovate or broad-ovate, with truncate or rounded base, obtuse at apex, obscurely crenulate 
or almost entire, with hairs as on stems but scattered, denser on lower surface or leaves 
sometimes glabrous, with papilliform hairs confined to midrib and petiole and with scat- 
tered small sessile glands only beneath; petioles 0.4-1 .6 cm long; flowers in lax one-sided 
inflorescence consisting of 3-6 pairs of flowers; bracts 2-5 mm long, 1-3 mm wide, nar- 
rowly ovate, entire, acute, with bloomlike vesture as on vegetative parts; calyx ca. 6 mm 
long, 2.5-3 mm wide, nearly black-lilac, densely covered with minute papilliform hairs 
interspersed with small sessile glands; corolla 2-2.4 cm long, four times as long as calyx, 
212 with nearly straight or slightly curved tube, 3.5-4 mm wide at throat, with Hps to 3 mm 
long, apparently violet, covered with short spreading hairs and short stipitate glands; nut- 
lets ca. 1 mm long, blackish-brown, finely tuberculate. Otherwise, similar to S. hissarica 
B. Fedtsch. June. 

Walls of ravines in the juniper forest belt. — Centr. Asia: Pam.-Al. (Kugitang Range in 
Turkmen SSR). Endemic. Described from northwestern slope of Kugitang Range oppo- 
site Kugitang village. Type in Leningrad. 

135. S. heterotricha Juz. et Vved. in Bot. mat. gerb. Bot. inst. ANSSSR, XIV (1951) 
427. — Anaspis heterotricha Juz. hoc loco (nom. alternat.). 

Perennial, 8-12 cm high, very similar to S. nevskii Juz. et Vved., but, in addition to 
minute hairs as in S. nevskii, with long simple spreading hairs, more copiously on axis of 
inflorescence, bracts and calyces, the upper side of leaves with scattered hairs, the lower 
side densely velutinous; lower leaves sometimes with subcordate base, in lower part with 
3 large obtuse teeth (larger toward apex) on each margin, resembUng leaves of S. fed- 
tschenkoi Bornm.; inflorescence at onset of flowering 2.5-4 cm long, of 4-5 pairs of 
flowers; calyx ca. 5 mm long; corolla ca. 1.8 cm long. Otherwise, like S. nevskii Juz. 
et Vved. June. 

Rocks. — Centr. Asia: Pam.-Al. (Tadzhikistan, Shirabad district). Endemic. Described 
from eastern slopes of Kugitang near Khodzhanka. Type in Leningrad. 

136. S. schugnanica B. Fedtsch. in 0. and B. Fedch. Perech. rast. Turk. V (1913) 159. 
— S. hissarica ssp. schugnanica M. Pop. in Bot. mat. gerb. Bot. Sada, V, 10 (1924) 
154. — Anaspis schugnanica Juz. hoc loco (nom. alternat.). 

Low subshrub; caudex stout, woody, strongly branching in upper part; stems numer- 
ous, crowded, 8-20 cm long, forming small tufts together with stems persisting from 



141 



previous year; stems 10-20 cm long, ascending or suberect, curved or slightly flexuous, 
obscurely 4-angled, brittle, with rather dense whitish bloomlike cover of minute hairs; 
leaves in 3-4 pairs, usually exceeding the shortish internodes, 0.6-4 cm long, 0.5-3.5 cm 
wide, broad-ovate, subcordate or truncate at base, obtuse or nearly rectangular at apex, 
coarsely crenate-dentate with 3-5 shallow obtuse teeth on each margin, sparsely or dense- 

213 ly covered on both sides with short fine hairs, thickish, firm, the veins scarcely prominent 
beneath; petioles spreading, 0.5-3 cm long, with vesture as on stems; inflorescence short- 
racemose, 3-6 cm long, one-sided, with internodes 3-5 mm long, and axis covered with 
short and longer horizontally spreading hairs interspersed with stipitate glands; bracts 
elliptic-rhombic, 2-4 (7) mm long, patent-hairy, with solitary glands, tapering to short 
petiole, this as long as or slightly longer than pedicel; calyx ca. 3 mm long, 2.5 mm wide, 
in fruit 5.5 mm long, with hairs as on axis of inflorescence but longer and more copious; 
corolla 2-2.3 cm long, many times the length of calyx, ca. 3 mm wide at throat, pink- 
violet, puberulent outside, with lips ca. 3 mm long. June-July. 

Rocks, crevices of bluffs. — Centr. Asia: Pam.-Al. (Shugnan, Darvaz). Endemic. De- 
scribed from Pyandzh River valley between Khorog and Dasht and from Kergovat. Type 
and paratypes in Leningrad. 

137. S. iipskyi Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 420. - S. his- 
sarica ssp. schugnanica var. pubescens M. Pop. in Bot. mat. gerb. Bot. sada, V (1924) 
154. — Anaspis Iipskyi Juz. hoc loco (nom. alternat.). 

Perennial, low or medium-sized; caudex woody, branching at summit; stems rather 
numerous, erect, simple, 7-17 (25) cm long, slender, subterete, brittle, glabrous or covered 
in inflorescence with scattered short spreading hairs or with more profuse horizontally 
spreading hairs of medium length and numerous small short-stipitate glands, encircled at 
base with remnants of previous year's stems; leaves 2-4 pairs, 0.8-3 (3.5) cm long, 0.6- 
2.7 (3) cm wide, as long as or slightly longer than internodes, broadly ovate or nearly 
triangular-ovate, with subcordate base, obtuse at apex, fairly thin, shallowly crenate- 
dentate with 1-3 (4) large obtuse teeth on each margin, densely covered on both sides 
with appressed gladiate somewhat crisp hairs, the veins not prominent beneath; petioles 
slender, 0.3-1 .5 (2) cm long, covered like stems, but more densely, with short and medium 
hairs; inflorescence 2-6 (8) cm long, consisting of 2-8 (10) approximate verticillasters; 
bracts rather large, 4-15 mm long, 2-10 mm wide, ovate or broad-ovate, obtuse, entire, 
with vesture like that of cauline leaves, short-petioled, at an thesis as long as or longer than 

214 calyx, in fruit twice as long; pedicels short; calyx ca. 4 mm long, 3 mm wide, in fruit 

5 mm long and wide, rather densely covered with long spreading hairs and short-stipitate 
glands; corolla ca. 2 cm long, several times the length of calyx, 3-4 mm wide at throat, 
sparingly pubescent and glandular outside, with slightly curved tube. June. 

Steep walls of ravines. - Centr. Asia: Pam.-Al. (Yakkabag). Endemic. Described from 
Mushketov ravine. Type in Leningrad. 

138. S. villosissima Gontsch. ex Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV(1951) 
421.— S. hissarica ssp. schugnanica var. villosa M. Pop. in Bot. mat. gerb. Bot. sada, 
V(1924) 154. — S. schugnanica var. villosa M. Pop. in sched. — Anaspis villosissi- 
ma Juz. hoc loco (nom. alternat.). 



142 



(215) , 




PLATE XIII. 1 - Scutellaria hissarica B. Fedtsch., general aspect, bract, flower, fruiting calyx, nut- 
let; 2 - S. baldshuanica Nevski, general aspect, leaf, fragment of inflorescence, flower, fruiting calyx, 
its parts; 3 - S. guttata Nevski, general aspect, leaf, bracts, flower, fragment of fruiting inflorescence, 
fruiting calyx. 



143 



Perennial, of medium size; caudex woody, branching; stems rather numerous, erect, 
simple or slightly branching, 17-30 cm long, stout, obscurely 4-angled, brittle, covered 
with long spreading somewhat crisp white hairs intermixed with rather numerous small 
short-stipitate glands, in inflorescence almost villous with more copious and longer hairs 
and gland stipes; leaves in few pairs, 1-3.5 cm long, 0.8-3.2 cm wide, equaling or shorter 
than internodes, broadly ovate, with rather deeply cordate base, obtuse at apex, rather 
deeply and coarsely, unevenly dentate with (0) 2-5 teeth on each margin, densely covered 
on both sides with long spreading multicellular slightly crisp hairs interspersed with short- 
stipitate glands; petioles slender, 0.8-1 .8 cm long, with hairs as on stems and on inflores- 
cence axis; inflorescence (2) 3.5-5 cm long, consisting of (2)3-6 subapproximate verticil- 
lasters; bracts 0.5-1 .2 cm long, 3-7 mm wide, ovate or narrowly ovate, usually acute, with 
hairs as on leaves; petioles 2.5-6 mm long; calyx ca. 5 mm long, 3 mm wide, in fruit ca. 
6 mm long, 4 mm wide, densely covered with long spreading hairs and stipitate glands; 
corolla ca. 1 .8 cm long, with nearly straight tube, ca. 4 mm wide at throat. June. 

Rocks. - Centr. Asia: Pam.-Al. (Guzar). Endemic. Described from upper reaches of 
Sangardak River, northern slope of Bel'-Auty Mountain. Type in Leningrad. 

^^^ 139. S. holosericea Gontsch. ex Juz. in Bot. mat. gerb. Hot. inst. ANSSSR, XIV 
(1951)423. — Anaspis holosericea Juz. hoc loco (nom. alternat.). - Ic: Yuz., op. 
cit. 425, Figure 5. 

Subshrub of medium size; caudex woody, 1-1.5 cm in diameter, branching at summit; 
stems few, 25-45 cm long, branching nearly from base or at summit; branches elongate, 
declined, curved or pendent, terminating like main stem in inflorescence, white with al- 
most velutinous spreading somewhat crisp hairs of medium length; leaves 1 .5-4 cm long, 
1.2-3.2 cm wide, together with petiole about equaling internodes, broad-ovate or nearly 
triangular-ovate, with shallowly cordate base, obtuse at apex, irregularly or rather regular- 
ly, coarsely and rather deeply crenate-dentate with 4-8 apically rounded teeth on each 
margin, whitish-green, rather densely velutinous on both sides, on upper surface with stiff- 
ish nearly straight and softer somewhat crisp hairs, on the lower surface hairs straight, 
horizontally spreading along the veins, somewhat crisp elsewhere; petioles 0.5-2 cm long, 
horizontally spreading, with hairs as on stems; inflorescence 5-12 cm long, branching or 
compound, the branches rather long, in the axils of usually entire upper cauline leaves, 
the summit of inflorescence and the branches secund, often nodding, the component in- 
florescences 3-7 cm long, of 4-12 approximate verticillasters; bracts 0.3-1 cm long, 1.5- 
5 mm wide, ovate, acute, entire, densely white-hairy; petioles to 1 mm long, equaling or 
slightly exceeding calyx, the latter ca. 2.5 mm long, 2 mm wide, in fruit ca. 5 mm long and 
wide, densely covered with rather long spreading hairs and short-stipitate glands; corolla 
rather small, to 1.5 cm long, densely covered outside with fine crisp hairs, the tube very 
slender, to 1 mm in diameter, ca. 3 mm wide at throat, the hps ca. 2 mm long. To September. 

Rocks (granite). - Centr. Asia: Pam.-Al. (western part). Endemic. Described from 
Shakhrut River valley, natural boundary Darai-Nikhan. Type in Leningrad. 



Subgenus 4. Apeltanthus Nevski ex Pavlov in Tr. Sredneaz. Gos. univ. ser. VIlI-c, ed. 
19 (1935) 35 (nomen nudum genericum); Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, 



144 



XIV (1951) 427 (pro sect.). — Inflorescence round or quadrangular in cross section (not 
dorsiventral) ; bracts herbaceous, similar to cauline leaves but sessile, smaller and narrow- 

218 er; calyx very slightly accrescent, concealed by bracts, the hps subequal, conchoid, the 
upper without appendage or distinct protuberance, persistent. Brittle usually rather 
densely patent-hairy and glandular, thin-leaved herbaceous plants, growing on precipitous 
outcrops and rocks. A Central Asian group. 

Note. A unique group that may be phylogenetically related to the subgenus (or genus) 
Anaspis, but in any case could be formally connected with it. However, with fractional 
interpretation of the generic category, it would be better to separate it from Anaspis as 
an independent genus, on the Unes proposed by Nevskii, inasmuch as it is possible to con- 
ceive of the group Apeltanthus as having evolved quite independently of Anaspis 
(most likely from the section Lupulinaria). 

1. Flowers rather small, ca. 0.7-2 cm long, the tube to 1 cm long 2. 

+ Flowers larger, to 2.5-3 cm long, the tube to 2-2.5 cm long 5. 

2. Inflorescences only terminal; corolla 1 .2-2 cm long 3. 

+ In addition to larger, terminal inflorescences, smaller lateral inflorescences produced 

in axils of upper cauline leaves or bracts; flowers smaller, not more than 1 cm long 
4. 

3. Leaves entire or nearly so, rarely with few flat crenations; flowers ca. 2 cm long. . 
142. S. poecilanthaNevski. 

+ Leaves (at least some) with some well developed, large teeth; flowers 1 .2-1 .5 cm 

long 143. S. macrodontaNevski. 

4. Stems and leaves densely covered with minute hairs visible only under high magni- 
fication 140. S. andrachnoides Vved. 

+ Stems and leaves covered with long spreading hairs and short-stipitate glands . . . 
141. S. baldshuanica Nevski. 

5. Basic color of flowers yellow 6. 

+ Flowers pink 148. S. ieptosiphon Nevski. 

6. Leaves densely covered on both sides with spreading soft white hairs 7. 

+ Stems and leaves covered on both sides with spreading hairs and copious glands . 8. 

7. Lower lip of corolla not spotted 144. S. orbicularis Bge. 

+ Lower Hp of corolla with red spots 147. S. rubromaculata Juz. et Vved. 

219 8. Lower lip of corolla not spotted 145. S. immacuiata Nevski. 

+ Lower Up of corolla with dark violet spots 146. S. guttata Nevski. 



Series 1. Baldshuanicae Juz. — Inflorescences of two kinds: terminal and lateral, the 
latter much smaller; corolla-tube short. 

140. S. andrachnoides Vved. in Addenda XIX, 350. - Anaspis andrachnoides vel 
Apeltanthus andrachnoides Juz. hoc loco (nom. alternat.). 

Small subshrub; caudex woody, 0.5-1 cm long, branching at summit; stems several to 
many, ascending or erect, slender, brittle, simple, 4-12 cm long, straight or curved, glau- 
cous with a dense bloomlike cover of minute hairs; cauline leaves commonly 2 pairs (not 



145 



counting leaves subtending inflorescence), 0.5-1.5 cm long, 0.3-1.3 cm wide, broadly 
ovate, with subcordate or rounded base, obtuse, entire or sometimes with 1-2 very in- 
distinct teeth on each margin, glaucescent on both sides with a dense cover of minute 
hairs distinguishable only under high magnification, firm, coriaceous; petioles 0.5- 
2mm long; terminal inflorescences ca. 2 cm long, ovoid-oblong, dense; lateral inflores- 
cences, often developing in the axils of terminal (floral) leaves, much shorter (ca. 1 cm), 
short-ovoid; bracts 0.5-1 cm long, 2.5-6 mm wide, broad- to narrow-ovate, almost scari- 
ous, pale green, sparsely covered with long flne simple hairs and small short-stipitate 
glands, differing markedly from cauline and floral leaves in consistency, color and vesture; 
calyx sparingly short-hairy and glandular, very slightly enlarging in fruit, ca. 2 mm long, 
subchartaceous; corolla small, ca. 1 cm long, sparsely short-hairy and finely stipitate- 
glandular. Fl. to September. 

Rocks. - Centr. Asia: T. Sh. (E. Fergana). Endemic. Described from Kyzyl-Dzhar, 
Naryn River, natural boundary Ine-Sai. Type in Tashkent, isotype in Leningrad. 

Note. This species differs sharply from the other members of the subgenus in its ves- 
ture which approaches that of some members of the preceding subgenus (Anaspis); on 
the whole, it would seem to hnk the subgenera (if not converging with the species of 
220 Anaspis). However, in consideration of the character of inflorescence- it must undoubted- 
ly be placed in the subgenus Apeltanthus. 

141 . S. baldshuanica Nevski ex Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (195 1) 
432. — Anaspis baldshuanica Juz. vel Apeltanthus baldshuanicus Juz. hoc loco 
(nom. alternat.). 

Subshrub; caudex woody, 5-7 mm thick, with flexuous subimplexed nodose ramifica- 
tions; flowering stems 2.5-7 cm long, ascending, curved or flexuous, slender, brittle, 
densely covered with long spreading white simple hairs and short-stipitate glands, produc- 
ing 2-several pairs of leaves and branches in some of the axils, these branches mostly short, 
leafy (usually with 1 pair of leaves), terminating like main stem in small inflorescences, 
sheathed by upper pair of leaves; leaves 0.3-1.2 cm long, 0.4-1 cm wide, broad-ovate or 
suborbicular, rounded or retuse at apex, entire or very often with few (1-3) large crena- 
tions on each margin, with 5-7 veins strongly impressed above and very prominent be- 
neath, almost plicate, sparsely or densely covered above with spreading hairs and short- 
stipitate glands, rather densely covered beneath with long spreading hairs and numerous 
short-stipitate glands, abruptly tapering to petiole, this short, with long spreading hairs and 
scattered stipitate glands; inflorescences ca. 1 .5 cm long, 8 mm wide (the lateral ca. 8 mm 
long), ovoid, dense, few-flowered; bracts much smaller than upper leaves, to 8 mm long, 
4 mm wide, narrowly ovate to ovate-lanceolate, tapering at base, sessile, subobtuse to sub- 
acute, differing markedly from cauhne leaves in size and shape but with similar vesture; 
calyx ca. 1 mm long at anthesis, the upper segment densely white-hairy, the lower glabrous; 
corolla small, ca. 7 mm long, pale lilac or whitish, with lilac upper lip; the tube short, 
0.75 mm in diameter, not exserted from bracts, to 2 mm wide at throat, the upper lip ca. 
2 mm long, the lower ca. 3 mm. July-August. (Plate XIII, Figure 2.) 

Slopes, crevices of rocks and crags, limestone outcrops, conglomerate bluffs, red sands. 
- Centr. Asia: Pam.-Al. (Gissar). Endemic. Described from Nargubakh, eastern slope of 
Rengan plateau between Kafirnigan and Yavan. Type in Leningrad. 



146 



Series 2. S. poec/7c«f/2fle Juz. — Inflorescences all alike, terminal. Corolla-tube 
short and wide. 

^2^ 142. S. poecilantha Nevski ex Juz. in Bot. mat. gerb. Bot. inst. ANSSSR, XIV (1951) 
431. — Anaspis poecilantha Juz. vel. Apeltanthus poecilanthus Juz. hoc loco 
(nom. altemat.). — Ic: Yuz., op. cit. 432, Figure 6. 

Subshrub; caudex woody, ca. 1 cm thick, with numerous ramifications, these assurgent 
or spreading, covered with yellowish bark, distally bearing squamiform villous-hairy basal 
leaves; stems 5-15 cm long, slender, brittle, covered with horizontally spreading long fme 
soft hairs and short glands, commonly developing 2-3 pairs of leaves; leaves 0.6-2 cm 
long and wide, broad-ovate to orbicular, rounded at apex, entire or nearly so (sometimes 
deeply notched at one of the margins, rarely with 2-3 flat crenae), thin, sparsely or rather 
densely covered on both sides, more copiously beneath, with long fine spreading hairs and 
subsessile glands, cuneate at base, the slender petiole patent-hairy, 0.5-1.2 cm long, the 
veins beneath somewhat prominent at flowering time; inflorescences at ends of stems, at 
onset of flowering ca. 1.5 cm long (without flowers), in fruit to 3 cm, ovoid-spicate, dense; 
lowermost bracts to 2.5 cm long, 1 .2 cm wide, resembling cauUne leaves but narrower, 
others smaller, ovate or narrowly ovate, often (especially the upper) subacute, tne longitu- 
dinal nerves on lower surface very prominent in fruit; calyx in flower to 2 mm long, in 
fruit ca. 3 mm long, densely patent-hairy; corolla ca. 2 cm long, mainly white, the tube 
to 1 cm long (largely concealed by bracts), to 2 mm thick; lips large, the upper 8 mm 
long, pale or purple-tinged, the lower ca. 6 mm long, mottled with dark purple, interrupt- 
ed stripes and spots. July. 

Rocks. — Centr. Asia: Pam.-Al. (basin of Upper Zeravshan). Endemic. Described from 
Yagnob River north of Anzob. Type in Leningrad. 

143. S. macrodonta Nevski ex Juz. sp. nov. in Addenda XIX, 351. - Anaspis macro- 
donta vel Apeltanthus macrodontus Juz. hoc loco (nom. alternat.). 

Perennial; similar to the preceding species, but leaves often bearing to 3 large obtuse or 
rounded teeth on each margin; bracts broader and more acute than in S. poecilantha; 
flowers exceptionally small, ca. 1 cm long. Otherwise, like S. poecilantha Nevski. 
June-July. 
222 Granite rocks. — Centr. Asia: Pam.-Al. (Gissar Range). Endemic. Described from 

Sardai-Mion River valley south of Vistap. Type (B. A. Fedchenko, No. 506) in Leningrad. 



Series 3. Orbiculares Juz. — Inflorescences alike, terminal; corolla-tube long and 
slender. 

144. S. orbicularis Bge. in Mem. pres. Ac. Sc. Petersb. div. sav. VII (1854) 436, s. str. - 
S, sericophylla Nevski in sched. — Anaspis orbicularis Juz. vel Apeltanthus orbi- 
cularis Juz. hoc loco (nom. alternat.). 

Subshrub; caudex woody, twisted, ca. 1 (to 2) cm thick, strongly branching from base; 
branches divaricate, prostrate, mostly flexuous, branching in turn and forming small tufts; 
green shoots 2-9 cm long, slender, brittle, the vegetative densely leafy, the fertile with 



147 



1-2 distant pairs of leaves, usually densely covered with spreading white soft hairs; leaves 
4-15 mm long, 3-15 mm wide, broad-ovate or obovate to orbicular, rounded-obtuse at 
apex, entire or sometimes with 1-3 shallow notches on each margin, densely covered on 
both sides with long soft appressed hairs, grayish-villous to whitish-sericeous, abruptly or 
gradually attenuate at base, usually with 7 veins (including midvein), these usually depres- 
sed above, prominent beneath; petiole about as long as blade (3-15 cm); leaves of sterile 
shoots slightly smaller, at first very densely silky lanate-villous; inflorescences at ends of 
fertile shoots, spicate-capitate, subglobose, 0.8-2 cm long (not including flowers), finally 
elongating to 2.5 cm; bracts tightly crowded, herbaceous, the lowermost to 1.2 cm long, 
1 cm wide, similar in every respect to cauline leaves but subsessile, the others gradually 
decreasing in size, thicker, orbicular-obovate to ovate, tapering at base, acute at apex; 
flowers 12-15; calyx sessile, at anthesis ca. 2 mm long, the upper segment rounded at 
apex, with an obsolescent dorsal protuberance but even in fruit unappendaged, densely 
covered with long silky hairs, the lower segment slightly shorter than the upper, very 
slightly emarginate, with very fine silky hairs, in fruit neither segment increasing signifi- 
cantly in size, ca. 3 mm long, coriaceous, falling away separately; corolla 1.7-2.8 cm long, 
stra\. J ^..ow, slightly glandular and pubescent outside, with long upcurved tube and large 
limb, to 5 mm wide at throat; nutlets brown, dull, glabrous. May-September. 
223 Rock crevices, outcrops (usually precipitous). — Centr. Asia: Pam.-Al. Endemic. 
Described from Zeravshan. Type in Leningrad. 

145. S. immaculata Nevski ex Juz. in Bot. mat. gerb. Bot. inst. ANSSSR, XIV (1951) 
428. — S. orientalis var. sewerzowii Herder in Bull. Soc. Nat. Mosc. XLI, II (1868) 
387. - S. suberosa Nevski in sched. - S. gymnocarpa Nevski in sched. - Anaspis 
immaculata Juz. vel Apeltanthus immaculatus Juz. hoc loco (nom. alternat.). 

Subshrub; caudex stout, woody, usually erect, twisted; ramifications numerous, short 
or long, with thick yellowish-brownish corky bark, their upper part and the lower part of 
stems densely covered with very crowded small squamiform obovate densely long-white- 
villous lower leaves; flowering shoots 3-15 cm long, slender, brittle, densely covered with 
long spreading white hairs, bearing 2-3 pairs of leaves; leaves 0.6-2 cm long, 0.5-1.7 cm 
wide, orbicular-ovate, rounded or angled at apex, entire or nearly so, grayish-green, with 
rather profuse fine spreading white hairs and numerous (especially beneath) subsessile 
glands, thinly coriaceous, mostly with 5 prominent veins beneath, abruptly cuneately 
tapering to petiole 6-10 mm long; inflorescence 1-2 cm long (without corollas), dense, 
capitate, in fruit to 5 cm long, ovoid to oblong-ovoid; bracts like cauline leaves but slight- 
ly narrower, acute, sessile, persistent in fruit, the lower 1 .2-1 .5 cm long, 7-9 mm wide; 
calyx to 2 mm long at anthesis, later ca. 2.5 mm long, the upper segment sparingly pubes- 
cent or subglabrous, the lower with few sessile glands; corolla 2.2 cm long, pale yellow, 
the tube rather long, 1 .5-2 mm in diameter, to 5 mm at throat, the upper lip ca. 7 mm 
long, the lower to 8 mm, not spotted; nutlets to 1.5 mm long, brown, sparsely and finely 
papillose-tuberculate. May -August. 

Rock crevices, precipitous stony ravine walls. — Centr. Asia: T. Sh. (western part), 
Pam.-Al. (foothills). Endemic. Described from Chatkal north of BrichmuUa and from 
Mashat (S. Kazakhstan). Type and paratypes in Leningrad. 

Note. Nevskii designated specimens of S. immaculata from Mogol-Tau mountains as 
S. gymnocarpa and specimens from the foothills of Pamir-Alai as S. suberosa. 

148 



146. S. guttata Nevski ex Juz. in Bot. mat. gerb. Bot. inst. ANSSSR, XIV (1951) 

429. - Anaspis guttata Juz. vel Apeltanthus guttata Juz. hoc loco (nom. alternat.). 

224 Subshrub ; caudex woody, long, flexuous, loosely branching in upper part, its branches 
gray-brown, nodose, often bearing in upper part numerous orbicular squamiform villous- 
hairy basal leaves; fertile shoots (stems) 5-10 cm long, usually ascending, simple or few- 
branched, slender, brittle, covered with horizontally spreading long fine simple hairs and 
very copious short and long glandular hairs; cauline leaves few, 0.6-1 .5 cm long, 0.5- 

1.2 cm wide, broadly ovate or suborbicular, generally rounded at apex, entire or nearly so, 
dull green, sparsely covered on both sides with long spreading hairs and numerous stipitate 
glands, with glandular hairs of varying length, often abruptly tapering to petiole 2-7 mm 
long; inflorescence at onset of flowering ca. 2 cm long (not including flowers), dense, 
ovoid-capitate, after flowering reaching up to 8 cm, narrowly cylindrical or columnar, 
some closely approximate; bracts narrower than cauUne leaves, ovate, usually acute and 
slightly revolute at apex, the lower ca. 1 cm long, 7 mm wide; calyx ca. 1 .5 mm long at 
anthesis, with densely stipitate-glandular upper and glabrous lower segment; corolla to 
2.5 cm long, bright yellow; tube 1.75-2 mm across, ca. 4 mm across at throat; upper lip 
ca. 5 mm long; lower lip ca. 6 mm long, with 9-18 small round dark violet spots, these 
somewhat randomly though fairly evenly disposed, sometimes confluent in pairs. June- 
September. (Plate XIII, Figure 3.) 

Sandstone outcrops. - Centr. Asia: Pam.-Al. (Tadzhik SSR, basin of Tupalang River). 
Endemic. Described from Gissar, Duob and Zambat. Type and paratype in Leningrad. 

147. S. rubromaculata Juz. et Vved. in Bot. mat. gerb. Bot. inst. ANSSSR, XIV (1951) 

430. - Anaspis rubromaculata Juz. vel Apeltanthus rubromaculatus Juz. hoc 
loco (nom. alternat.). 

Perennial; apparently a very stenotypic species, of somewhat composite character; its 
vesture resembles closely that of S. orbicularis Bge., to which it is most closely allied, 
while the red (not dark violet) spots on the lower lip of the corolla are of the same size and 
shape as in S. guttata Nevski. May. 

Habitat unknown. - Centr. Asia: Pam.-Al. (Zeravshan). Endemic. Described from 
Zeravshan valley, Naufin-Kul' Lake. Type in Leningrad. 

225 148. S. leptosiphon Nevski in Tr. Bot. inst. AN SSSR, I, 4 (1937) 324. - S. propin- 
qua Nevski in sched. - S. microcalycina Nevski in sched. - Anapsis leptosiphon 
Juz. vel Apeltanthus leptosiphon Juz. hoc loco (nom. alternat.). - Ic: Nevskii, op. 
cit.. Figure 13 (p. 323). 

Cespitose subshrub; caudex twisted, usually decumbent, woody, strongly branching 
from base, the branches covered with yellowish-gray corky bark and often bearing basal 
leaves in upper part, these squamiform, orbicular-obovate, ca. 3 mm long, tapering toward 
base; flowering shoots mostly ascending, 3-8 (15) cm long, slender, with 1-3 pairs of 
leaves; leaves 8-17 mm long, 7.5-15 mm wide, orbicular-ovate, entire or very rarely ob- 
scurely crenate-dentate, covered, more copiously beneath, with fine white spreading hairs 
and stipitate glands, abruptly tapering to petiole, this 3-6 (10) mm long; inflorescence 
ovoid-capitate, 1-2 cm long (not including corollas); bracts similar to cauline leaves but 
sessile, ovate, subacute; pedicels 0.5-1 mm long; calyx at anthesis to 2 mm long, after 



149 



flowering 2.5-3 mm long, both segments rounded at apex, the upper with low protuber- 
ance and long wliite hairs, the lower slightly shorter, scarcely glandular at base; corolla 
3 cm long, pale pink, the tube very long, much exserted, curved upward, glandular-hairy 
outside, 1-1.25 mm thick, the upper Up mitriform, short-winged laterally, glandular on the 
back, ca. 5 mm long, the lower lip ca. 6 mm long, suborbicular, with a purple-lilac spot at 
middle and numerous small violet dots. June -July. 

Limestone rocks. - Centr. Asia: Pam.-Al. Endemic. Described from foothills of Kugi- 
tang Range near Kugitang village. Type in Leningrad. 



Subfamily 3. LAVANDULOIDEAE Briq. in Bull. Herb. Boiss. II (1894) 715; id. in 
Pflanzenfam. IV, 3a (1895) 207 et 227. - Calyx 13-15-nerved, 2-lipped, of the ^/j or V4 
type; corolla 2-hpped of the type ^3; stamens 4, included and curved downward; anthers 
with 2 spreading cells fusing at apex; ovary 4-parted; nutlets with spinobasal attachment; 
disk lobes liguihform, opposite ovary cells and nutlets; embryo straight, with straight 
short radicle. 

226 Genus 1245.* Lavandula** L. 

L.Syst.ed. 2 (1740)25. 

Calyx tubular, 13-15-nerved, 2-lipped, the upper Up 1-toothed, the lower with 4 equal 
teeth; corolla blue or violet, with exserted tube, 2-lipped, the upper lip of 2, the lower lip 
of 3 orbicular, more or less flat lobes; stamens 4, not exserted from corolla, the lower 
longer than the upper; style 2-cleft above; nutlets with smooth shining surface, yellowish- 
brown. Subshrubs or small shrubs, rarely perennial herbs, with entire, linear or lanceolate- 
linear, obtuse, sometimes pinnatifid leaves. 

More than 20 species distributed from the Canary Islands to India and Pakistan are 
represented in the genus. 

1. L. spicaL. Sp. pi. (1753) 572 var. j3. exclud. - L. vulgaris var. a. Lam. Fl. Franc. 
11(1778)403. - L. spica L. a. angustifolia L. fil. in Amoen. Acad. 10(1790) 58. - 
L. officinalis Chaix in Vill. Hist. pi. Dauph. I (1786) 355. -L. fragrans Salisb. Prodr. 
(1796) 78. - L. vera DC. in Lam. et DC. Fl. Franc. V (1815) 398. - L. spicata Eaton, 
Man. Bot. N. Amer. (1817) 65. - Ic: Komarov, Sbor, sushka i razved. lekarstv. rast., 
3rd. ed., Plate 26 (1917); Rchb. Ic. Fl. Germ. XVIII, tab. 1227. 

Subshrub 20-1 00 cm high; branches numerous, ascending or erect, strongly branching; 
leaves oblong-linear, entire, obtuse, revolute, gray-tomentose when young, 2-6 cm long, 
2-6 mm wide; flowers in 6-10-flowered verticillasters at ends of stems, forming an inter- 
rupted spiciform inflorescence; bracts rhombic-ovate, acuminate, short, 3-5 mm long; 

* Treatment by B.K. Shishkin. 

** From the Latin lavare - to wash, referring to the use of lavender in the bath by the ancient Greeks 
and Romans. 



150 



calyx 4-5 mm long, rather densely covered with white hairs, with 5 obtuse, very short 
teeth; corolla 1 cm long, violet, twice the length of calyx. 

Cultivated for its valuable essential oil in the Crimea, Caucasus and Central Asia. Gen, 
distr.: W. Mediterranean region. Described from S. Europe. Type in London. 

Note. Another species of Lavandula - L. latifolia (L.) Vill., distinguished by its 
227 broader leaves and linear bracts, is also cultivated for its essential oil which is known as 
oil of spike. It is similar to oil of lavender but of somewhat inferior quality. 

Economic importance. Lavender was used for its perfume by the ancient Romans. 
The essential oil is contained in epidermal cells of the leaves, in glandular hairs and, in 
appreciable amount, in the calyx. It is obtained by steam distillation of inflorescences 
and vegetative parts of the plant. An extract is also obtained by applying solvents (for 
example, benzene) to the inflorescence. Freshly collected plants yield 0.8% oil and dried 
plants up to 1.5%. The most valuable component is linalyl acetate. The distilled oil has 
a pleasant aroma; it is slightly bitter in taste and faintly yellowish-green in color. Laven- 
der oil is used in the production of perfumes, soap and other cosmetics. It is one of the 
principal constituents of eau de Cologne. 



Subfamily 4. DREPANOCARYOIDEAE* Pojark. subfam. nova in Addenda XIX, 
351 . — Calyx 1 8-20-nerved, nearly straight at throat, obliquely inflated at base; corolla 
2-lipped of the % type; stamens 4; anthers diverging at an obtuse angle of over 1 80° and 
thus the locules projecting upward; ovary 4-partite, with style attached to base of lobes. 
Nutlets with basiventral attachment, falcately curved along longitudinal axis so that the 
ventral side appears concave and the dorsal side strongly convex, semi-globular; each nut- 
let attached by a large surface of contact (occupying its entire ventral side) to the outer 
concave side of one of 4 large lunate apical appendages of the disk, these alternating with 
the liguliform disk-lobes (disk-lobes finally scarcely visible). Embryo slightly curved, 
with fleshy cotyledons and radicle turned downward and attached at base of cotyledons. 

Type of subfamily — genus Drepanocaryum Pojark. 

Note. The characters which sharply distinguish the genus Drepanocaryum from 
other Labiatae are as follows: 1) basiventral attachment of nutlets, 2) shape of the nut- 
lets: curvature along the longitudinal axis and surface of attachment occupying the whole 
ventral side, 3) presence of 4 large disk appendages (4-lobed gynophore) to which the 
nutlets are attached; the latter feature belongs to the category of characters which under- 
lie the division of the Labiatae family into subfamilies. On this basis the genus Drepano- 
caryum is segregated into a subfamily. 

228 Genus 1246. Drepanocaryum** Pojark. 

Gen. nov. in Addenda XIX, 352. 

Calyx erect, gibbous at base, without a ring of hairs inside, 18-20-nerved, nearly straight 
at throat, the lower teeth longer than the upper; corolla-tube glabrous inside; upper lip of 

* Treatment by A.I. Poyaikova. 
** From the Greek drepanon, sickle and caryon, seed (in this case, the nutlet). 

151 



(229) 




PLATE XIV. 1 - Drepenocaryum sewerzowii (Rgl.) Pojark., a) flower unfolded, b) corolla un- 
folded, c) stamen, d) fruiting calyx in rear view, e) same in side view, f) early stage of fruiting (note 
disk with appendages (gynophore) enclosing nutlets), g) advanced stage of fruiting (2 nutlets removed), 
h) mature nutlets in bottom and side view, i) gynophore in side view (after shedding of nutlets), k) the 
same in bottom view (note disk), 1) embryo; 2 - Agastache rugosa (Fisch. et Mey.) O. Ktze., flower 
unfolded; 3 - Schizonepeta multifida (L.) Briq., flower unfolded. 



152 



corolla 2-lobed; lower lip with erect lateral lobes, the flat cuneate middle lobe directed 
upward; stamens arcuately ascending under upper lip of coroUa, the upper longer than 
the lower, all fertile; anther cells not confluent; style with 2 unequal lobes; nutlets 
coarsely tuberculate. Annual plants, with flowers in remote cymes. Type and only re- 
presentative — D. sewerzowii (Rgl.) Pojark. 

1. D. sewerzowii (Rgl.) Pojark. comb. nov. — Nepeta sewerzowii Rgl. in Tr. Bot. 
sada, VI (1879) 360; Aitch. in Trans. Linn. Soc. 2 ser. Ill, 147; Lipsk. in Tr. Bot. sada, 
XVIII, 101; 0. and B. Fedch. Perech. rast. Turk. V, 147. - Ic: Aitch. 1. c. tab. 41. 

Annual; root slender, short; stems 6-60 cm long, erect or ascending, slender, often 
flexuous, usually simple, rarely with 1-2 pairs of leafy fertile branches in lower part, more 
or less densely covered with hairs, these very small (magnifying glass!), spreading, simple, 
2 (3)-cellular, thickened proximally, sometimes interspersed with coarser and longer hairs; 
leaves thin (chartaceous when dry), bright green, sparsely, rarely densely, covered on both 
sides with simple coarse jointed hairs, these appressed above, more or less spreading be- 
neath, and sparsely punctate with soon deciduous yellow glands, ovate or cordate or the 
upper often narrower, oblong-ovate or ovate-elliptic, obtuse, rarely acute, the lower and 
middle leaves to 4 cm long, 3.5 cm wide, upper leaves often nearly half as long, the margin 
with large obtuse or acute teeth; petioles slender, patent-hairy or downy, in lower leaves 
23 1 equaling or longer than blade, decreasing in size toward summit, the uppermost 1 .5-2 mm 
long; cymes in axils of all leaves (except sometimes the lower with axillary branches); 
leaves exceeding peduncles, sometimes barely as long; peduncles slender, 4.5-6 cm long; 
upper cymes 3-8-flowered, usually dense, the others 8-20-flowered, loosely dichotomous- 
ly branching; floral leaves linear-subulate, 2-3 mm long; bracts slightly smaller, shorter 
than pedicels, these 1 .5-3 mm long, in case of bifurcation to 5-10 mm long; all axial parts 
of inflorescence densely covered with very short simple spreading hairs, sometimes with 
admixture of capitate glands; calyx 6-7.3 mm long, green or violet at an thesis, nearly 
tubular, distinctly gibbous at base, in fruit broader, ovoid, strongly inflated, membranous, 
with thickish nerves, short-patent-hairy, the teeth half as long as tube, the 3 upper lanceo- 
late-triangular, 2-2.3 mm long, the two lower longer, 2.2-2.5 mm, narrower and long- 
acuminate, all terminating in a hard but not spiny point and lined inside with long appress- 
ed hairs; corolla (blue or lilac) 6-6.7 mm long, included in calyx to limb or even higher, 
hairy outside; tube 3.7 mm long, gradually dilated into narrowly infundibular neck, 1.3- 
1.7 mm long; upper lip 0.8-4 mm long, deeply cut into semiorbicular-ovate obtuse lobes; 
lower lip twice as long as the upper, the lateral lobes about equaling the lobes of upper lip, 
0.7-0.8 mm long and as wide, the middle lobe 1 .4-2 mm long and wide, cuneate, notched 
at apex, flat, entire; nutlets 0.7-0.8 mm long, 1-1.2 mm wide, subglobose above, concave 
at lower end, black, coarsely tuberculate. Fl. second half of April -first half of July; fr. 
May -July (August). (Plate XIV, Figure 1.) 

Lower and middle mountain belt, 700-2000 m, ravines, stony and gravelly slopes in the 
shade of rocks and trees. - Centr. Asia: Mtn. Turkm. (Greater Balkhan Range, Kopet- 
Dagh), Kyz. K. (Aktau mountains), Pam.-Al. (all mountains, apparently, except for 
Shugnan and Pamir), T. Sh. (mountains of Kara-Tau and W. T. Sh.). Gen. distr.: Iran 
(Badgyz mountains in Afghanistan and Baluchistan). Described from Mogol-Tau moun- 
tains. Type (lectotype) in Leningrad. 



153 



Subfamily 5. STACHYDOIDEAE Briq. in Pflanzenfam. IV, 3a (1895) 307 et 329. - 
Calyx varying in shape; corolla 2-lipped, of the %type, rarely almost regular, the tube 
cylindrical or rarely campanulate; stamens 4, rarely 2, ascending or turned forward; 
ovary 4-partite, lobes of disk alternate with lobes of ovary; nutlets ob ovoid, with small 
basal surface of attachment; embryo with short straight radicle. 



232 Tribe 1. MARRUBIEAE Endl. Gen. (1838) 627; Benth. in DC. Prodr. X, 407; Briq. 
in Pflanzenfam. IV, 3a, 207. — Calyx tubular or campanulate, with prominent nerves, 
5-10-toothed; corolla-tube included in calyx, rarely exserted; stamens and style included 
in corolla-tube. 



Genus 1247.* Thuspeinantha** Durand. 

Ind. Gen. Phan. (1888) 703. - Tapeinanthus Boiss. Diagn. Ser. 1, 12 (1848) 68; Boiss. ex Benth. in 
DC. Prodr. XII, 446; non Herb. (1837). 

Calyx broadly campanulate, 10-nerved, slightly 2-Upped, nearly regular, herbaceous, 
with very short broadly triangular teeth, two of which slightly longer and less deeply cut, 
glabrous at throat, inflated in fruit; corolla-tube long, slender, exserted from calyx, slight- 
ly dilated at throat, without hairy ring inside; upper lip 2-lobed, the lobes small, obtuse, 
oblong-lanceolate, flat; lower hp 3-lobed, the lobes flat, the middle one broader; stamens 
4, not exserted from throat of corolla, attenuate or ascending, nearly parallel, the lower 
slightly longer; filaments short; anthers with coalescent cells, almost unilocular, oblong, 
attached to filaments at middle; style bipartite, the truncate upper lobe sheathed at base 
by the lower lobe; nutlets large, oblong, slightly attenuate at base, obtuse at apex, keeled 
on the ventral side. Small annual herbs, with opposite petiolate leaves. 

Two species of the genus are confined to desert habitats in Central Asia, Iran and 
Afghanistan. 

1. T.persica (Boiss.) Briq. in Pflanzenfam. IV, 3a (1897) 229; 0. and B. Fedch.Perech. 
rast. Turk. V (1913) 162; Fedch. Rast. Turk. (1915) 672. - Tapeinanthus persicus 
Boiss. Diagn. Ser. 1, 12(1848)63; Boiss. ex Benth. in DC. Prodr. XII, 436; Boiss. Fl. or. 
IV, 679. - Chamaesphacos persicus (Boiss.) Aitch. et Hemsl. in Transact. Linn. Soc. 
Ser. II, III (1888-1894) 97. - Ic: Aitch. et Hemsl. 1. c. tab. XLII, fig. 1-6. 

A small glandular-viscid annual, dull green, simple or branching from base; stems 
erect; leaves 1-veined, obtuse, entire or obscurely dentate, 1.5-2.5 cm long, the lower 
oblong, ca. 2.5 cm long, 5-7 mm wide, longer than the upper, lanceolate to oblong-lanceo- 
233 late, all leaves petiolate, the lower with longer petioles; flowers paired on short peduncles 
in axils of upper leaves; calyx 6-7 mm long, with short glandular hairs around the teeth 
inside and outside, the teeth broadly triangular, acute, at anthesis 1/6 as long as the tube, 

* Treatment by A.G. Borisova. 
** From the Greek tuspein and anthos, flower. 



154 



234 



in fruit larger, the whole calyx enlarging sacciformly, with open throat; corolla 12-15 mm 
long, white or pink, with long tube, dilated at throat; nutlets over 3 mm long, 1.5 mm 
wide, broadening and obtuse at apex, with sessile glands at base, with fine hairs on both 
faces, convex on the outside, flat and keeled on ventral side. Flowers March-April, fruit 
April-June. 

Sands in deserts, red sandstone, pebbly places. — Centr. Asia: Ar.-Casp., Kara K., Kyz. 
K. Gen. distr.: Iran. Described from Iran. Type in Geneva. 



Genus 1248.* Marrubium** L. 

L. Sp. pi. (1753) 582; N. Pop. in Protok. Obshch. estestv. Yur'evsk. univ. XXIII, 4 (1916). 

Calyx tubular, 5-10-nerved, 5-10-toothed, the teeth equal or the alternate shorter, 
erect or declined and re flexed; corolla 2-lipped, the tube shorter than calyx, rarely ex- 
serted; upper lip of corolla erect, nearly flat, narrow, notched or 2-lobed; lower lip de- 
clined, 3-lobed, with the middle lobe broader, often notched, the lateral lobes oblong; 
stamens 4, included in corolla-tube, the upper shorter than the lower; anther cells divari- 
cate, apically coalescent and dehiscent by a common slit; lobes of style short, obtuse; 
nutlets trigonous-ovoid or oblong, rounded at apex. Perennial, very rarely annual, fre- 
quently white-tomentose-lanate plants, with stellate fasciculate multicellular hairs; in- 
florescence of small flowers in usually dense axillary verticillasters, with subulate or 
setaceous bracts. 

Upward of 40 species distributed in Europe, N. Africa and extratropical Asia. 

Economic importance. All species of the genus provide a source of tanning agents and 
dyes (with ferrous salts they yield an excellent black color). Some of the species are nec- 
tariferous and some have medicinal value (see Note toM. vulgare L.). 

1 . Calyx (6) 8-10-toothed, the auxiliary (commissural) teeth as long as or slightly 
shorter than the principal teeth 2. 

+ Calyx 5-toothed (very rarely 6-7 and then very small) 6. 

2. Calyx-teeth erect 3. 

+ ' Calyx-teeth recurved 4. 

3. Calyx-teeth subulate-lanceolate; corolla white, twice as long as calyx; leaves oblong- 
ovate, entire at base, crenate-serrate above 4. M. praecox Janka. 

+ Calyx-teeth subulate; corolla purple, slightly exserted from calyx; leaves semiorbi- 
cular, crenate 3. M. woronowii N. Pop. 

4. Bracts curved, 1/3-1/2 as long as calyx; corolla whitish-yellowish, scarcely exserted 
from calyx-tube 5. M. parviflorum Fisch. et May. 

* Treatment by O.E. Knorring. 

** Name used by many Latin authors for various species of the genus and also for some other Labiatae. 
It appears to derive from the Hebrew words mar, bitter and rob, much (i.e. "very bitter"), but 
Linnaeus formed it from "Maria-Urbs" (town on Lake Fitin in Latium). 



155 



235 



+ Bracts as long as or longer than calyx; corolla pink, white or yellow, somewhat 

exserted from calyx-tube 5. 

5. Calyx-teeth divergent, hamate; corolla white, slightly longer than calyx; nutlets 
oblong-ellipsoid 1 . M. vulgare L. 

+ Calyx-teeth slightly curved or straight, not hamate; corolla pink or pale yellow, half 
as long again as calyx; nutlets obovoid 2. M. alternidens Rech. 

6. Calyx-teeth erect 7. 

+ Calyx-teeth recurved 10. 

7. Corolla white; inflorescence branching with horizontally spreading branches. Leaves 
oblong-rhombic at base, entire, crenate-serrate above . . .12. M. peregrinum L. 

+ Corolla pinkish-violet or purple 8. 

8. Inflorescence branching; upper verticillasters confluent; stems coarse, thick, cover- 
ed with stellate hairs; leaves broadly ovate, nearly as wide as long 

14. M. purpureum Bge. 

+ Inflorescence not branching; stems numerous, slender; upper verticillasters not con- 
fluent; leaves narrower, ovate or orbicular 9. 

9. Leaves orbicular, obtusely toothed at margin; corolla purple 

13. M. goktschaicum N. Pop. 

+ Leaves ovate, entire, sparsely toothed only at apex; corolla lilac 

15. M. turkeviczii Knorr. 

10. Entire plant niveous-tomentose with fine flexuous hairs; leaves orbicular, with short 
unequal teeth 11. M. persicum C.A.M. 

+ Grayish-green plants, covered with bristly or stellate multicellular hairs . . . .11. 

11. Annuals, divaricately branching from base; leaves ovate, coarsely crenate; corolla 
white or pink, with long-exserted tube 7. M. catariifolium Desr. 

+ Perennials 12. 

12. Calyx-teeth nearly as long as calyx-tube; leaves reniform, coarsely crenate . . . . 
6. M. plumosum C.A.M. 

+ Calyx-teeth 1/3 to 2/3 as long as tube 13. 

13. Plant 9-25 cm high; stems solitary, not branching; cauline leaves obovate, finely 
toothed 9. M. nanum Knorr. 

+ Plant 30-60 cm high; stems branching; cauline leaves ovate 14. 

14. Leaves covered on both sides with stellate hairs, with simple hairs along veins be- 
neath; upper hp of corolla much longer than the lower . . 8. M. ieonuroides Desr. 

+ Leaves covered on both sides, more densely beneath, with fascicled as well as glandu- 
lar hairs; upper lip of corolla as long as the lower 

10. M. propinquum Fisch. et Mey. 



Section 1. Marrubium Benth. Lab. gen. et sp. (1834) 588. - Calyx-teeth 5-10; 
upper lip of corolla notched or 2-cleft; bracts hnear; leaves orbicular or ovate, toothed 
or crenate, rarely incised. 



Subsection 1. Decemdentata Briq. in Pflanzenfam. IV, 3a (1897) 230. - Calyx-teeth 
6-10, of which the auxiliary (commissural) as long as or shorter than the principal teeth. 



156 



1. M. vulgare L. Sp. pi. (1753) 583; M. B. Fl. taur.-cauc. II, 54; Benth. Lab. gen. et 
sp. 591 et in DC. Prodr. XII, 453; Ldb. Fl. Ross. Ill, 406; Boiss. Fl. or. IV, 703; Shmal'g. 
Fl. II, 331 ; N. Pop. in Protok. Obshch. estestv. Yur'evsk. univ. XXIII, 4, 141. - M. apu- 
lum Ten.Fl. Nap. atlas, IV (181 1-1838) 34, tab. 154. -M. hamatum Humb. et Kunth, 
Nov. gen. et sp. II (1817) 250. - M. anisodon C. Koch in Linnaea, XXI (1848) 696; 
Trautf. in Tr. Bot. sada, IX, 103. - M. vulgare var. lanatum Benth. in DC. Prodr. XII 
(1848)453. -M. vulgare var. arcuata Trautv. op. cit. V (1874) 471. -M. v. var. ma- 
crocalyx N. Pop., op. cit. 141 ; Grossg. Fl. Kavk. Ill, 288. - M. kusnezovii N. Pop., 
op. cit. (1916) 145.- Ic: Fedch and Fl. Fl. Evrop. Ross. 605, Figure 714. - Exs.: Fl. 
morav. No. 261 , Fl. Palest. No. 175. 

Perennial, 30-60 cm high, whitish-lanate ; stems simple or branching, covered with long 
236 appressed hairs; basal and cauline leaves orbicular-ovate, coarsely crenate-dentate; the 
terminal similar but smaller, rugose, pale green with scattered long hairs above, grayish be- 
neath with long bristly hairs, prominently veined, short-petioled; lower verticillasters very 
distant, the upper approximate; bracts subulate, recurved, densely covered with long hairs, 
shorter than or equaling calyx-tube; calyx with 10 divergent subulate hamate teeth, of 
these 5 longer, 1/3 to 1/2 as long as the long-hairy tube; coroUa white; the upper hp as 
long as or slightly shorter than the lower, 2-cleft at apex, the lower lip with broadly reni- 
form middle lobe and small lateral lobes; nutlets oblong-eUipsoid, spotted, finely tuber- 
culate. June -August. 

Roadsides, cultivated fields, wastelands. - European part: Bait., U. Dnp., M. Dnp., Bl., 
Crim., L. Don; Caucasus: Cisc, E. and S. Transc, Tal.; Centr. Asia: Ar.-Casp., Balkh., 
Kyz. K., Mtn. Turkm., Amu D., Syr D. Gen. distr.: Centr. Eur., Med., Bal.-As. Min., 
Iran, Ind. -Him., Dzu.-Kash. Described from N. Europe. Type in London. 

Note. We do not attach much taxonomic significance to the forms of this polymor- 
phous species, described by various authorities as varieties or even as independent species 
(mainly based on the character of vesture or the curvature of calyx-teeth) because they 
are not associated with definite geographical areas. 

Economic importance. Known as a medicinal plant. At one time it was used in W. Eu- 
rope as a remedy for chronic ailments of the respiratory tracts (the medicament was pro- 
duced in Russia). Horehound has also been advocated as a substitute for cinchona bark, 
but pharmacological tests have not so far yielded positive results. 

The aerial parts contain a crystalline bitter principle and amorphous bitter principles 
of unknown composition. 

A good nectariferous plant, producing a large amount of nectar. / 

2. M. altemidens Rech. fil. in Oesterr. Bot. Zeitschr. (1952) 37. - M. vulgaris var. 
oligodon Rech. fU. ap. Koie, Dan. Scient. Invest, in Iran 4 (1945) 44. 

Perennial, 30-100 cm high, erect, branching in upper part, coarse, flocculent-hairy in 
lower half, lanate above; lower leaves elUptic or orbicular, to 5 cm long and 4 cm wide, 
239 coarsely crenate, erose-crenate at apex; upper cauline and floral leaves resembling the 
lower but smaller, pale green, rugulose, velutinous above, gray beneath, densely covered 
with appressed fascicled and bristly hairs; basal and cauhne leaves with petioles 3-6 mm 
long, the upper short-petioled; inflorescence long, of 12-15 many-flowered verticillasters, 
these very distant in lower part, only the uppermost 2 or 3 approximate; bracts subulate- 
setaceous, as long as or slightly longer than calyx, the lower bracts partly declined, partly 

157 



(237) 




PLATE XV. 1 - Marrubium parviflorum Fisch. et Mey., general aspect, calyx, corolla, seed; 2 - M. 
persicum C.A.M., fragment of inflorescence, calyx, corolla; 3 - M. goktschaicum N. Pop., general 
aspect, calyx, corolla, seed. 



158 



appressed to calyx, covered with fine cellular and fascicled hairs; calyx with 10 unequal 
teeth, 5 longer teeth alternating with 5 shorter ones, patent, slightly arched-recurved or 
straight, with a ring of long hairs at throat at base of teeth, covered outside with simple 
and fascicled hairs; corolla pale yellow or pale pink, half as long again as calyx, the upper 
Up incised-notched, the broadly linear lobes rounded at apex, the lower lip with broadly 
reniform undulate-margined middle lobe and oblong round-tipped lateral lobes, the tube 
narrow at base, cupuUformly dilated above; nutlets dark brown, obovoid, obtusely 3- 
angled, smooth, blackish. 

Dry places, wormwood-grass and grass-mixed herbs steppe, often a weed. — Caucasus: 
E. Transc. (Gogenaker, reported by Rechinger); Centr. Asia: T. Sh. (W.), Pam.-Al., Mtn. 
Turkm. (Kopet-Dagh). Gen.distr.: Bal.-As. Min. (Anatolia), Iran., Ind. -Him. Described 
from Shakhrud-Bustam province in Iran. Type in Vienna. 

Note. The species is highly variable in the vesture of stem and leaves as well as in the 
shape of leaves. 

3. M. woronowii N. Pop. in Protok. Obshch. estestv. Yur'evsk. univ. XXIII, 4 (1916) 
146; Grossg. Fl. Kavk. Ill, 287. - Ic: N. Pop., op. cit. 117, Plate 4. 

Perennial, 20-40 cm high; stems ascending, reddish; lower leaves semiorbicular, cre- 
nate in upper part; bracts oblong, rounded at apex, tapering at base, twice as long as ver- 
ticillasters; outer bracts slightly shorter than calyx, rigid, subulate-spinescent; calyx with 
10 erect, subulate teeth, half as long as tube, the tube covered with stellate and simple 
hairs; corolla purple, exserted. April. 

Shrubs, in central mountain belt. — Caucasus: S. Transc. (western part). Endemic. 
Described from former Artvin district. Type probably in TbiUsi. 

4. M. praecox Janka in Oesterr. Bot. Zeitschr. XII (1875) 62; Boiss. Fl. or. IV, 702; 
Shmal'g. Fl. 33 1 ; N. Pop. in Protok. Obshch. estestv. Yur'evsk. univ. XXIII, 148; Grossg. 
Fl. Kavk. Ill, 288. - M. pannonicum Rchb. Fl. Germ. exc. (1833) 325; Benth. in DC. 
Prodr. XII, 452. - M. paniculatum Desr. in Lam. Encycl. Ill (1789) 716. - M. pesta- 
lozzae Boiss. Diagn. Ser. II, 4 (1859) 54. - Ic: Rchb. PI. crit. Ill, 86, tab. 300. - Exs.: 
GRF, No. 835. 

Perennial, 30-60 cm high, gray appressed-tomentose; stems covered with short fasci- 
cled hairs; branches directed upward; leaves oblong-ovate or oblong, entire at base, 
crenate-serrate in upper part, netted-rugose above, covered with monoradiant stellate hairs 
(i.e. with 1 elongate ray) and simple hairs, deeply netted-rugose beneath, prominently 
veined, more densely covered with closely appressed stellate (partly with elongate rays) 
and simple hairs, short-petioled; terminal (floral) leaves oblong-lanceolate, longer than 
verticillasters; inflorescence with many verticillasters, these 8-10-flowered, very distant 
in lower part, approximate above; bracts subulate, equaling or slightly shorter than calyx- 
tube, covered with stellate hairs with 1 elongate ray and bristly hairs; calyx 5-10-toothed, 
the teeth erect, unequal, lanceolate-subulate, rigid, acuminate, 1/3-1/2 as long as tube, 
the tube cylindrical, prominently nerved, densely covered with stellate hairs, some of 
these markedly monoradiant; corolla white, nearly twice as long as calyx, the tube with 
stellate and sparse simple hairs outside under throat, glabrous in lower half, the upper Up 
deeply cut into two lobes, the lower lip with reniform middle lobe and oblong-ovate 
lateral lobes; nutlets ovoid, finely tuberculate. June-August. 

159 



Steppes, fields. - European part: M. Dnp., V.-Don, Bes., Bl., Crim., L. Don, L. V.; 
Caucasus: Cisc, Dag., E. Transc. Gen. distr.: Centr. Eur., Bal.-As. Min. Described 
from Austria. Type in Vienna. 

5. M. parviflorum Fisch. et Mey. Ind. sem. hort. Petrop. II (1835) 33; Boiss. Fl. or. 
IV, 704; N. Pop. Protok. Obshch. estestv. Yur'evsk. univ. XXIII, 4, 146; Grossg. Fl. Kavk. 
Ill, 288. -M. peregrinum C.A.M. Verz.(1831) 28, non L. - M. radiatum Benth. in 
DC. Prodr. XII (1848) 452, non Delil.; Ldb. Fl. Ross. Ill, 405. 

Perennial, 40-60 cm high; stems 1-4, erect, branching above, whitish-grayish, densely 
covered with fine long branching stellate hairs, these with unequal rays; radical leaves 
many, white-tomentose, eUiptic or orbicular; cauline leaves oblong, tapering at base, fine- 
241 ly toothed, green above, with bristly and long hairs (mainly on the convex sections, i.e. 
between nerves), covered beneath with long, flexuous and fascicled hairs and sparse mono- 
radiant stellate hairs, petiolate; floral leaves similar to the cauline but smaller, subsessile, 
exceeding verticillasters; inflorescence long, lower verticillasters very distant, 8-10-flower- 
ed, the upper approximate; bracts curved, 1/3-1/2 as long as calyx, covered with flexuous 
and fascicled hairs; calyx prominently nerved, the tube 4.5-5.5 mm long; 5 or 10 irregu- 
lar, 2-2.5 mm long, teeth densely covered with unequiradiant or monoradiant stellate 
hairs (glandular hairs sometimes occur); corolla whitish-yellowish, the tube scarcely ex- 
serted from calyx, the upper lip short-notched, with 2 oval lobes, the lower lip with broad- 
ly elhptic undulate-margined or denticulate middle lobe, the elliptic lateral lobes as long as 
middle lobe. May-July. (Plate XV, Figure 1.) 

Lower and middle mountain belt, dry places. - Caucasus: E. and S. Transc, Tal. Gen. 
distr.: As. Min., Iran. Described from Talysh. Type in Leningrad. 

Subsection 2. Quinquedentata Briq. in Pflanzenfam. IV, 3a (1897) 230. - Calyx- 
teeth 5, very rarely 6-7 and then auxiUary (commissural) teeth small. 



Group 1. Stellata Briq. 1. c. - Calyx-teeth stellate-declined. 

6. M. plumosum C.A.M. Verz. (1831) 96; Benth. Lab. gen. et sp. (1834) 589; id. in 
DC. Prodr. XII, 448; Ldb. Fl. Ross. Ill, 403; N. Pop. in Protok. Obshch. estestv. Yur'evsk. 
univ. XXIII, 4, 157; Grossg. Fl. Kavk. Ill, 290. - M. leonuroides Desr. var. plumosum 
Boiss. Fl. or. IV (1872) 701; Shmal'g. Fl. II, 331. 

Perennial, 28-35 cm high; stems slightly curved, branching, covered with long hairs; 
lower leaves reniform, coarsely crenate, with petioles of medium length; upper leaves 
orbicular, short-petioled or sessile, grayish-green rugose above, sparsely covered with 
monoradiant stellate hairs, gray beneath with copious tightly appressed stellate hairs and 
prominent veins; verticillasters 4-6-flowered; bracts nearly as long as calyx, subulate- 
setaceous, densely covered with simple and monoradiant stellate hairs; calyx-teeth 5, 
patent, subulate-acicular, as long as tube, covered up to the naked point with multijointed 
242 and fascicled hairs; corolla pale pink, slightly exserted from calyx, the upper lip as long as 
the lower, cut to at least half its length into narrowly oblong lobes; the lower lip with 



160 



broadly flabelliform middle lobe and oblong lateral lobes; nutlets oblong, obtusely 
3-angled, brown, glabrous. June. 

Alpine mountain belt, taluses. - Caucasus: Dag. Endemic. Described from alpine 
belt of E. Caucasus. Type in Leningrad. 

7. M. catariifolium Desr. in Lam. Encycl. Ill (1789) 717; Benth. in DC. Prodr. XII, 
449; M. B. Fl. taur.-cauc. II, 53; Ldb. Fl. Ross. Ill, 403; Boiss. Fl. or. IV, 699; N. Pop. 
inProtok.Obshch.estestv. Yur'evsk. univ. XXIII, 4, 155; Grossg. Fl. Kavk. Ill, 289. - 
M. candidissimum C. A. M. sec. Boiss. 1. c. non L. — M. catariaefolium Desr. var. 
microphyllum Somm. et Lev. in Tr. Bot. sada, XVI (1900) 392. - Exs.: PI. or. exs. 
No. 121. 

Annual, 15-45 cm high; stems divaricately branching nearly from base, stellate-hairy, 
the hairs partly short, partly monoradiant (with 1 elongate ray); leaves thin, the basal and 
cauline ovate-oblong, coarsely crenate, pale green above, subglabrous or with sparse simple 
and stellate hairs with 1 elongate ray, covered beneath with copious short-rayed stellate 
and sparse fascicled hairs; lower leaves with petioles 10-20 mm long; upper leaves short- 
petioled; floral leaves resembling the cauline but smaller; flowers in 6-9 verticillasters, 
these very distant, only the uppermost 2 or 3 approximate; bracts as long as calyx, subu- 
late, spinescent, rigid, partly recurved, partly spreading, covered with monoradiant stellate 
and fascicled hairs; calyx half as long as tube, appressed-hairy, covered with monoradiant 
and fascicled hairs, with 5 short, lanceolate-subulate, patent teeth; corolla pink or white, 
the tube markedly exserted from calyx, the upper lip as long as the lower, cut into two 
oblong lobes, the lower lip with broadly infundibuliform middle lobe and oblong lateral 
lobes; nutlets ellipsoid, dark brown, finely tuberculate. June- August. 

Dry slopes, scrub ; a weed among crops. — Caucasus: Cisc, S. and E. Transc. Gen. 
distr.: As. Min. Described from Syria. Type in Paris. 

8. M. leonuroides Desr. in Lam. Encycl. Ill (1789) 715; Benth. Lab. gen. et sp. 588 et 
in DC. Prodr. XII, 450; Ldb. Fl. Ross. Ill, 404; Boiss. Fl. or. IV, 701 ; Shmal'g. Fl. II, 
331 ; Briq. in Pflanzenfam. IV, 3a, 331 ; N. Pop. in Protok. Obshch. estestv. Yur'evsk. 

243 univ. XXIII, 4, 151 ; Grossg. Fl. Kavk. Ill, 289. - M. astracanicum M. B. Fl. taur.-cauc. 
II (1808) 52; Stev. in Mem. Soc. Nat. Mosc. Ill, 266, non Jacq. - Ic: Rchb. PI. crit. Ill, 
tab. 299. 

Perennial, 60-65 cm high; stems coarse, slightly curved, branching, covered with short 
stellate and simple hairs; lower leaves ovate, coarsely incised-crenate; upper leaves oblong, 
covered above with stellate and 1 -rayed stellate hairs, beneath along veins with simple 
hairs, densely covered between the veins with short, 5-6-rayed stellate hairs; lower leaves 
with longer petioles, the upper short-petioled; floral leaves exceeding verticillasters, these 
many-flowered and forming a long inflorescence; lower leaves remote, the upper approxi- 
mate; bracts not spinescent, shorter than calyx, the outer divergent, the inner appressed 
to calyx, covered with long simple fine flexuous hairs; calyx 1/3-1/2 as long as tube, cov- 
ered with appressed stellate fascicled and sparse multicellular hairs; teeth 5, subulate, acu- 
minate, glabrous, short, patent and curved; corolla pink, the tube slightly exserted from 
calyx, glabrous at base, hairy above, the upper lip cut to middle into 2 lobes, much longer 
than the lower lip, this with broadly reniform middle lobe and oblong lateral lobes; nutlets 
elliptic, brown. May- July. 

161 



Dry slopes and weed-infested places from lower to upper mountain belt. — European 
part: Crim. (eastern part); Caucasus: Cisc, Dag. Gen.distr.: As.Min. Described from 
As. Min. Type in Paris. 

9. M. nanum Knorr. in Bot. mat. gerb. Bot. inst. AN SSSR, XV (1953) 346, fig. 
p. 347. 

Perennial, 9-25 cm high; stems single, not branching, densely leafy, covered with fme 
tangled hairs; lower and median cauline leaves obovate, 1.5-2 cm long, 1.1-1.6 cm wide, 
dentate; the petioles 8-10 mm long; floral leaves resembling the cauline but smaller, 
green, sparsely covered with bristly hairs above, with long coarse bristly hairs on the veins 
beneath, verticillasters 5 or 6, subapproximate, 8-10-flowered; bracts subulate, equaling 
calyx-tube, ca. 4 mm long, covered with long fine hairs; calyx covered with short bristles 
and long hairs only along the prominent nerves, long-hairy inside at base of throat, the 5 
triangular subulate teetli 2/5-1/2 as long as tube, their points glabrous; corolla pale pink, 
exserted from calyx, the upper hp as long as the lower, shallowly notched, the lower lip 
244 with broadly orbicular middle lobe and shorter oblong lateral lobes; nutlets ellipsoid, 
brown, obtusely 3-angled. May. 

Caucasus: S. Transc. Endemic. Described from Nakhichevan ASSR. Type in Leningrad. 

10. M. propinquum Fisch. et Mey. Ind. sem. hort. Petrop. 1 (1835) 33; Benth. Lab. 
gen. et sp. 743 et in DC. Prodr. XII, 450; Ldb. Fl. Ross. Ill, 404; Boiss. Fl. or. IV, 701 ; 
N. Pop. in Protok. Obshch. estestv. Yur'evsk. univ. XXIIl, 154; Grossg. Fl. Kavk. Ill, 289. 
-M. coerulescens Desf. Cat. Horti reg. Paris, ed. 3(1829) 395; Benth. in DC. Prodr. 
XII, 449. 

Perennial, 30-45 cm high; stems coarse, curved, divaricately branching above, covered 
with long tangled hairs; lower cauline leaves ovate, the middle and upper obovate, cune- 
ately tapering toward base, dark green, rugose above, with scattered short fascicled hairs, 
grayish-green beneath, the prominent veins densely covered with same as well as glandular 
hairs, petioles 8-10 mm long; floral leaves eUiptic-cuneate, subsessile; verticillasters re- 
mote, many-flowered, approximate only at summit of inflorescence; bracts slightly curved, 
partly recurved, slightly shorter than calyx; calyx prominently nerved, covered with multi- 
cellular and glandular hairs, inside with profuse multicellular hairs below throat, the 5 pa- 
tent subulate point-tipped teeth 2/3 as long as tube; corolla pink, the tube exserted from 
calyx, the upper lip 2-lobed, the lower with broadly elliptic middle lobe and oblong lateral 
ones. June-July. 

Dry slopes in mountain regions. - Caucasus: Tal. Gen.distr.: Iran. Described from 
Talysh. Type in Leningrad. 

Note. Both Popov and Grossgeim attributed glabrous bracts to this species, but in fact 
they are always rather densely covered with multicellular hairs. 

11. M.persicum C.A.M. Verz.(1831)95; Benth. in DC. Prodr. XII, 449; Ldb. Fl. 
Ross. Ill, 404; Boiss. Fl. or. IV, 700; Briq. in Pflanzenfam. IV, 3, 230; N. Pop. in Protok. 
Obshch. estestv. Yur'evsk. univ. XXIII, 4, 190; Gross. Fl. Kavk. Ill, 289. - Exs.: Fl. or 
exs. No. 292. 

Perennial, 35-45 cm high, niveous, densely tomentose-lanate with fme flexuous tangled 



162 



hairs; stems erect, branching; basal and cauUne leaves orbicular, unevenly crenulate, 
grayish -green, rugose on both sides, sparsely covered with flexuous, tangled hairs; floral 

245 leaves elliptic, short-petioled or subsessile; inflorescence with 6-9 verticillasters, these 
6-8-flowered, most distant, only the uppermost 2 or 3 approximate; bracts very small, 
subulate, twisted, 1/4-1/2 as long as calyx, covered with flexuous hairs; calyx-tube white- 
tomentose with shorter flexuous appressed hairs; calyx-teeth 5, subulate, spreading, rigid, 
spiny-tipped, 1/3-1/2 as long as tube; corolla lilac, not exserted from calyx, the upper 
lip slightly longer than the lower, shallowly notched into orbicular lobes, the lower 

lip with infundibuliform undulate-margined middle lobe and oblong lateral lobes. May- 
June. (Plate XV, Figure 2.) 

Dry stony foothill slopes. — Caucasus: W., E. and S. Transc, Tal. Gen. distr.: Iran. 
Described from Talysh (Zuvant). Type in Leningrad. 

Group 2. Microdonta Briq. in Pflanzenfam. 3a (1897) 230. - Calyx-teeth erect, 2/3- 
1/3 as long as tube. 

12. M. peregrinum L. Sp. pi. (1753) 582; Benth. Lab. gen. et sp. 590 et in DC. Prodr. 
XII, 452; M.B.Fl.taur.-cauc. 11,53; Ldb. Fl. Ross. Ill, 409; Boiss. Fl. or. IV, 694; 
Shmal'g. Fl. 33 1 ; N. Pop. in Mat. fl. Kavk. Ill, 590. - M. creticum Mill. Gard. Diet. 
VIII (1768) No. 3. - M. peregrinum var. creticum C. Koch in Linnaea, XXI (1848) 
696. -Ic: Rchb.Fl.Germ. tab. 23; Fl. Yugo-vost. VI, 161, Figure 683. 

Perennial, 30-60 cm high, gray-tomentose; stems few; branches horizontally spreading, 
covered with short appressed and fascicled hairs, below inflorescence with stellate and fas- 
cicled hairs; basal and cauUne leaves oblong-rhombic, entire at base, crenate-serrate above; 
terminal leaves smaller, lanceolate, brownish-green rugose above, covered with simple and 
monoradiant stellate or only with simple hairs, prominently veined beneath, with short 
fascicled hairs; inflorescence branching, with many subremote 8-10-flowered verticillas- 
ters; bracts as long as to equahng calyx-tube, subulate, thickish, gray, densely covered with 
fascicled and monoradiant stellate hairs; calyx campanulate, prominently nerved, densely 
covered with monoradiant stellate and fascicled hairs; teeth 5, strong, erect, triangular, 
becoming patent after flowering; corolla white, glabrous from base to middle, with stellate 
hairs above, the tube slightly exserted from calyx, the upper lip notched to middle into 2 

246 oblong lobes, the lower lip slightly shorter than the upper, with broadly reniform middle 
lobe and oblong lateral lobes; nutlets triangular-ellipsoid, dark brown, finely tuberculate. 
June- July. 

Dry steppe slopes. - European part: V.-Kama (along TaUev), Bl., Crim.; Caucasus: 
Cisc, according to Grossgeim in W. and E. Transc. Gen. distr.: Centr. Eur., Bal.-As. Min. 
Described from Centr. Eur. Type in London. 

13. M. goktschaicum N. Pop. in Protok Obshch. estestv. Yur'evsk. univ. XXIII, 4 
(1916) 160; Grossg.Fl. Kavk. 111,289. 

Perennial; rhizome curved, with numerous slender rootlets; stems numerous, 20- 
40 cm high, curved, purple, covered with long fine hairs, more densely so at base and 
below inflorescence; lower leaves ovate, 2.5-3 cm long, 1 .5-2 cm wide, with rather long 



163 



petioles, obtusely toothed; bracts ovate-elliptic, coarsely toothed, tapering at base, short- 
petioled or sessile, several times longer than verticillasters, pale green above with scattered 
simple hairs, rugose beneath, densely covered with stellate and simple hairs; verticillasters 
numerous, 10-12-flowered, distant in lower part of inflorescence, approximate above; 
bracts subulate, tough, spiny-tipped, straight, about equaling calyx, covered with very fine 
long white hairs and sparse stellate ones; calyx covered with simple fine long hairs and 
sparsely stellate-hairy; teeth 5, erect, subulate, 1/4-1/3 as long as tube; corolla lilac, long- 
exserted, the upper hp 2-cleft, the lower with broadly reniform undulate-margined middle 
lobe and oblong lateral lobes, the tube glabrous below middle, covered above, like hmb of 
coroUa, with stellate hairs; nutlets obovoid, dark brown, finely tuberculate. May- July. 
(Plate XV, Figures.) 

Steppe mountain slopes. — Caucasus: S. and W. Transc, Tal. Endemic. Described 
from Sevan Lake in Armenia. Type in Leningrad. 

14. M.puprureumBge.inMem.Acad. Sc. Petersb.ser. VII,XXI, 1 (1873) 162; 
Grossg. Opred. rast. Kavk. 329. — ? M. astracanicum Jacq. Ic. rar. I (1781-1786) 11, 
tab. 109; N. Pop. in Protok. Obshch. estestv. Yur'evsk. univ. XXIII, 4, 160; Grossg. Fl. 
Kavk. Ill, 289. - M. astracanicum var. raddeanum Alb. Prodr. Fl. Colch. (1895) 
201; N. Pop., op. cit. 162. 

Perennial, 20-35 cm high; stems ascending at base, coarse, sparsely covered in lower 
part, more densely above (especially beneath inflorescence), with stellate hairs and stellate 
|247 hairs with 1 long ray; lower leaves broadly ovate, suborbicular, coarsely crenate, with 
petioles 2-4 cm long; floral leaves oblong, cuneately tapering at base, toothed only at 
apex, twice as long as verticillasters, subsessile, grayish sericeous above with stellate (in- 
cluding monoradiant) hairs; inflorescence branching, the many-flowered verticillasters 
sub remote, the terminal closely approximate; bracts subulate, point-tipped, nearly as long 
as tube, sometimes reaching calyx -teeth, covered with simple hairs; calyx covered with 
stellate (including monoradiant) hairs, the 5 short erect teeth 1/3-1/2 as long as tube; 
corolla purple, the upper Hp slightly longer than the lower, deeply notched with 2 oblong 
lobes, the lower lip with infundibuliform middle lobe and oblong lateral lobes; nutlets 
elongate-ellipsoid, dark brown. June- July. 

Stony places in southern and middle mountain belts. — Caucasus: S. Transc, Tal. 
Gen. distr.: Iran. Described from Iran. Type in Paris. 

15. M, turkeviczii Knorr. sp. nov. in Addenda XIX, 352. 

Perennial, 34-36 cm high, woody at base; stems few, slender, slightly curved, purple- 
brown, covered with fascicled and simple hairs; radical and cauline leaves ovate, with 
short rounded teeth toward apex; upper cauline leaves oblong, tapering at base; floral 
and upper cauhne leaves 3-4 times as long as verticillasters, with petioles 20-30 mm long, 
pale green above, sparsely covered with bristly and monoradiant stellate hairs, gray be- 
neath, densely covered with appressed monoradiant stellate hairs and long hairs; verticil- 
lasters 7-10-flowered, distant, the uppermost 2 or 3 approximate; bracts as long as calyx, 
covered with long hairs; calyx with somewhat prominent nerves, the 5 erect subulate 
point-tipped teeth covered with long hairs except for glabrous point, the tube covered 
with stellate (including monoradiant) hairs; corolla lilac, the tube exserted from calyx. 



164 



glabrous at base, with stellate hairs near throat, the upper lip longer than the lower, deeply 
notched with 2 oblong lobes, the lower Up with broadly infundibuhform oval middle lobe 
and oblong-oval lateral lobes; nutlets dark brown, ellipsoid, 3-angled, finely tuberculate. 
May. 

Rocks. — Caucasus: S. Transc. Gen. distr.: E. Anatolia. Described from former Kars 
region, slopes near Araks River. Type in Leningrad. 

Note. Although this plant was collected outside the USSR, we consider it necessary 
to report it in the "Flora of the USSR," because it was collected so close to the border. 



Genus 1249.* Lagopsis**Bge. 

Bge. ex Benth. Lab. gen. et sp. (1834) 586. 

Calyx tubular or tubular-campanulate, 10-nerved, with 5 unequal teeth (2 longer than 
the others); corolla with long tube and 2-lipped limb, the upper lip erect, entire or shal- 
lowly notched at apex, the lower Up 3-lobed, patent, with broader middle lobe; stamens 
4, the lower longer than the upper but not exserted from corolla-tube; anthers 2-celled, 
the cells divergent; style 2-lobed at apex, the lobes short, subovoid, obtuse; nutlets 
oblong or ovaloid, 3-angled, obtuse at apex, smooth or alveolate. Perennial herbs, with 
palmatisect or pedate leaves. 

Four species distributed mainly in the Soviet Union but also occurring (2 species) in 
the Dzungaria-Kashgar area and in Mongolia. 

1. Flowers in long loose spicate inflorescences, forming 16-20 distant or partly approx- 
imate whorls; corolla white or pale pink 4. L. supina (Steph.) Ik.-Gal. 

+ Flowers in short dense spicate inflorescences; corolla cinnamon brown or yellow . 
2. 

2. Corolla cinnamon brown; leaves reniform-orbicular 3. 

+ Corolla yellow; leaves cordate or broadly ovate 2. L, flava Kar. et Kir. 

3. Bracts as long as calyx; leaves green on both sides, subglabrous 

3. L. eriostachya (Benth.) Ik.-Gal. 

+ Bracts 1/3-1/2 as long as calyx; leaves white-lanate on both sides 

1 . L. marrubiastrum (Steph.) Ik.-Gal. 

1. L. marrubiastrum (Steph.) Ik.-Gal. from Kryl. Fl. Zap. Sib. IX (1937) 2303. - Mo- 
lucella marrubiastrum Steph. in Mem. Soc. Nat. Mosc. (1809) 8, excl.ic; Bge. in Ldb. 
Fl. Alt. II, 817. - M. laniflora Willd. ex Benth. in Linnaea, XI (1837) 339. - Marrubi- 
249 um lanatum Benth. Lab. gen. et sp. (1834) 587; id. in DC. Prodr. XII, 442; Ldb. Fl. 
Ross. Ill, 402. - Lagopsis incana Bge. Suppl. alt. (1836) 59. - Ic: Ldb. Ic. pi. Fl. 
Ross. II, tab. 150. 

Perennial, 5-20 cm high; stems ascending, spreading, covered sparsely at base, densely 
below inflorescence, with white-lanate flexuous pubescence; leaves orbicular-reniform, 

* Treatment by O.E. Knorring. 
** From the Greek lagos, hare and opsis, appearance, face. 

165 



shallowly palmatipartite with 5 ovate or suboblong lobes, these with obtuse or rounded 
teeth at apex, both sides white-lanate with fine tangled hairs; lower leaves with long pu- 
bescent petioles; uppermost leaves short-petioled, smaller, rhombic, mostly with 3 orbi- 
cular-ovate lobes; flowers in terminal, oblong, ovoid, densely villous, compact spicate 
inflorescences, only 1 or 2 lowermost verticillasters subdistant; bracts setaceous, 1/3-1/2 
as long as calyx, densely covered with long fine hairs; calyx campanulate, densely covered 
all over with very long white hairs, the 5 triangular teeth acutely spinous-tipped; corolla 
cinnamon-brown, as long as or nearly as long as calyx, the upper lip entire, broadly ovate, 
distally hairy outside, the lower lip with 3 orbicular lobes; lower stamens twice as long as 
the upper; nutlets brown, ovaloid, tapering at base, subobtuse at apex, glabrous. 
June-July. 

Gravelly and stony places, in the desert-steppe belt and adjacent parts of the alpine 
mountain belt. — West Siberia: Alt. (Chu steppe, Kurai steppe, and others). Endemic. 
Described from Altai. Type in Leningrad. 

2. L. flava Kar. et Kir. in Bull. Soc. Nat. Mosc. XV (1842) 425. - Marrubium flavum 
Walp. Repertorium, III (1844-1845) 856; Benth. in DC. Prodr. XII, 448; Ldb. Fl. Ross. 
111,403. 

Perennial, 14-27 cm high; stems few, coarse, 4-angled, sparsely or densely tomentose- 
lanate; leaves cordate, shallowly palmatipartite with 3-5 oval or ovate lobes, obtusely 
toothed at apex, glabrous above or hirsute, more densely so beneath; petioles of lower 
leaves 2-3.5 cm, the upper 1 cm long; flowers in terminal ovoid-ovaloid spikes; verticil- 
lasters crowded, only the lowermost 2 or 3 distant; bracts one-third as long to as long as 
calyx-tube, covered with long hairs; calyx tubular-campanulate, covered with long hairs, 
the 5 subulate-acicular teeth unequal but approximately as long as calyx-tube; corolla 
yellow, the longer upper Up oblong-elliptic, distally hairy outside, the lower Up with 
250 broadly elliptic middle lobe and eUiptic lateral lobes; nutlets ovoid, pale brown, alveolate. 
July-August. (Plate XVI, Figure 2.) 

GraveUy places in alpine mountain belt. — Centr. Asia: Dzu.-Tarb., T. Sh. Qen. 
distr.: Dzu.-Tarb. Described from sources of Sarkan River (Dzungaria Ala-Tau). Type 
in Leningrad. 

3. L. eriostachya (Benth.) Ik.-Gal. in herb. (1936). — Marrubium eriostachyum 
Benth. Lab. gen. et sp. (1834) 586; id. in DC. Prodr. XII, 448. - Leonurus eriostachys 
Turcz. mss. ex Benth. 1. c. 586. - L. eriophorus Turcz. ex Steud. Nomencl. bot. II 
(1841) 25. — Lagopsis viridis Bge. Mon. gen. Molucc. ined. ex Benth. 1. c. 586. — 
Molucella mongholica Turcz. ex Ldb. Fl. Ross. Ill, 402. 

Perennial, 15-25 cm high; stems few, ascending, coarse, brown-purple, sparingly pubes- 
cent at base, densely so below inflorescence; leaves reniform-orbicular, deeply palmatifid, 
5-lobed, with ovate or oval obtusish teeth, green on both sides, subglabrous or short-hairy 
on the veins beneath, the margins revolute, cUiate; lower and middle leaves with petioles 
1-2 cm long, upper leaves broadly short-petioled; inflorescence spicate, oblong-ovoid, 
densely villous, only the lowermost 2 or 3 verticillasters distant; bracts subulate or acicular, 
rigid, nearly as long as calyx, covered with long hairs; calyx tubular-campanulate, complete- 
ly covered with long hairs; calyx-teeth 5, triangular, unequal, subulately long-acuminate. 



166 



(251) 




PLATE XVI. 1 - Lagopsis supina (Steph.) Ik.-Gal.; 2 - L. flava Kar. et Kir. 



167 



patent in flower; corolla cinnamon-brown, the upper lip about as long as the lower, 
broadly elliptic, long-hairy outside, the lower lip with broadly orbicular middle lobe 
and orbicular lateral lobes; nutlets ovaloid-oblong, rounded at apex. June-July. 

East Siberia: Ang.-Say. Gen. distr.: Mong.? Described from East Siberia. Type in 
Leningrad. 

4. L. supina (Steph.) Ik.-Gal. in herb. (1936). — Leonurus supinus Steph. ex Willd. 
Sp.pl. 111(1800) 116. - Marrubium incisum Benth. Lab. gen. et sp. (1834) 586; id. 
in DC. Prodr. XII, 447; Ldb. Fl. Ross. Ill, 402; Kom. Fl. Man'chzh. Ill, 349. - Krasch- 
nikowia (sic) geraniifolia Turcz. ex Ldb. 1. c. 402. 

Perennial, 15-35 cm high; stems few, ascending from base, often branching, densely 
covered with short hairs; leaves deeply incised, 3-lobed, the lobes with rounded or oblong 
253 teeth, green on both sides, sparsely hirsute above, hairy only on the veins beneath, with 
ciliolate margins; lower leaves v^th petioles 1 5-20 cm long, the upper short-petioled; 
inflorescence long, spicate, composed of 16-20 verticillasters, these 8-12-flowered, mostly 
subremote, 3 or 4 closely approximate; bracts about equaling calyx, curved, acicular, 
densely hirsute; calyx broadly campanulate, densely hirsute, the 5 unequal triangular 
point-tipped teeth 1/3-1/2 as long as calyx; corolla pale pink or white, slightly exserted 
from calyx, the upper Up longer than the lower, apically villous-hairy on the outside, the 
lower Up 3-lobed, with broadly elliptic middle lobe and elliptic lateral lobes; nutlets 
oblong-ovoid, brown, scaly. July-August. (Plate XVI, Figure 1.) 

Damp places on riverside terraces, roadsides. — East Siberia: Ang.-Say., Dau. Gen, 
distr.: Mong., Ch. and Jap. Described from East Siberia. Type probably in Berlin. 



Genus 1250.* Sideritis** L. 

L. Sp.pl. ed. 1 (1753)574. - Hesiodia Moench, Meth. PI. (1794) 391. - Burgsdorffia Moench, 1. c. 
392. - Empedoclea Rafm. Caratteri (1810) 78. 

Calyx tubular-campanulate, 5-10-nerved, sometimes slightly 2-lipped, with 5 erect, 
generally spiny teeth; corolla with tube enclosed in calyx, with 2-hpped limb; upper lip 
erect, nearly flat, entire or emarginate-bifid; lower lip spreading, 3-cleft, the middle lobe 
larger than the lateral, notched; stamens 4, didynamous, not exserted, the upper very 
short, with divaricately 2-celled anthers, the lower longer, mostly with deformed or re- 
duced anthers; style also enclosed in corolla-tube, bifid, the upper lobe short, truncate, 
the lower dilated, surrounding the upper at base; nutlets ovoid, smooth, obtuse or 
rounded at apex; flowers in 6- to many-flowered verticillasters, axillary or forming spicate 
inflorescences. Annual or perermial herbs or subshrubs, usually very hairy or tomentose, 
with entire or dentate leaves. 

* Treatment by S.V. Yuzepchuk. 

** From the Greek sideros, iron. The name was used for the plant in antiquity and also during 
the 16th century because of its presumed effectiveness in the treatment of wounds inflicted by 
various weapons. 



168 



Note. A Mediterranean genus containing many species. In the Soviet Union it is 
meagerly represented; most species occur in the Crimea. 

254 1. Perennials (subshrubs) 2. 

+ Annuals 8. 

2. Floral leaves mostly longer than wide, merely contiguous at margins or scarcely 
overlapping, rather small, up to 2 cm, usually not becoming yellow 3. 

+ Floral leaves nearly as wide or wider than long, markedly overlapping at margins, 
the largest usually more than 2 cm, becoming more or less distinctly yellow at 
flowering 5. 

3. Floral leaves small, 0.5-1 .3 cm long, 6-9 mm wide, narrowly triangular, firm, grayish- 
green, very hairy, differing very httle from upper cauUne leaves in consistency, color 
and indument 1. S. taurica Steph. 

+ Floral leaves mostly somewhat larger, to 1 .8 cm long, 1 .6 cm wide, more or less mar- 
kedly differing from upper cauUne leaves in thinner consistency, color (green or 
yellowish-green) and sparser indument 4. 

4. Floral leaves thin, dark green; verticillasters remote ... 2. S. chlorostegia Juz. 
+ Bracts coarse, turning yellowish; verticillasters mostly approximate (especially in 

upper part of inflorescence) 4. S. marschalliana Juz. 

5. Verticillasters remote. Plants of Crimean highland pastures (and parly of southern 
coast of the Crimea) 3. S. catillaris Juz. 

+ Verticillasters generally closely approximate. Plants of E. Crimea and W. Trans- 

caucasus 6. 

6. Plants of E. Crimea; floral leaves markedly yellowing, rather sparingly pubescent; 
flowers also rather sparingly pubescent; calyx eglandular 7. 

+ W. Transcaucasian plant ; floral leaves grayish-green or faintly yellowing; floral 

leaves and flowers copiously hairy; calyx often with glandular hairs 

7. S.euxinaJuz. 

7. Stems mostly simple, usually bearing 1 terminal inflorescence; verticillasters relative- 
ly few (2-9, rarely up to 12), the lowermost often remote . . . 6. S. conferta Juz. 

+ Stems often (though not always) branching, with well developed lateral inflores- 

255 cences in addition to terminal inflorescence; verticillasters (5) 8-15, all more or 
less closely approximate 5. S. imbrex Juz. 

8. Flowers yellowish-brown, subsessile; verticillasters more or less distant; at least 
lower internodes of inflorescence equaling calyx 9. 

+ Flowers purple, distinctly pediceled; verticillasters closely approximate; internodes 
of inflorescence shorter than calyx 10. S. balansae Boiss. 

9. All floral leaves herbaceous, green; calyx-tube cyHndrical, finally with constriction 
in upper part, the teeth often patent but straight 8. S. montana L. 

+ Upper floral leaves scarious, yellow, forming a kind of coma at the summit of inflo- 
rescence; calyx-tube obconical, without distinct constriction, the teeth suberect- 
patent and mostly arched-recurved 9. S. comosa (Rochel) Stank. 



Section 1. Empedoclea (Rafin. pro gen.) Benth. Lab. gen et sp. (1834) 571. - Peren- 
nial herbs; stems commonly woody at base, i.e. suffrutescent. Bracts unlike cauline leaves 



169 



at least in shape, usually also in consistency and vesture, mostly broadly cordate, subentire, 
acuminate, covering sessile flowers in verticillasters forming a spikelike inflorescence; calyx 
with 5 equal, acutely spinescent teeth. 

Note. The species of this section are confined in their distribution to the east Mediter- 
ranean area. Apparently all members of the section occurring in the USSR belong to one 
natural series that may be designated as Tauricae Juz.; it would evidently include some of 
the Balkan-Asia Minor forms. Until recently, all Soviet components of this series were re- 
cognized as species, S. taurica "M.B."; this, however, has proved to be an aggregate spe- 
cies, providing an excellent example of an ancient Crimean-Novorossiisk endemic form, 
with distinct Asia Minor connections, which segregated into several narrowly localized, 
concrete (elementary) species. Six of these are known in the Crimea, the seventh growing 
in the vicinity of Novorossiisk. 

It should be emphasized that determination of these species with the aid of a key does not 
always produce satisfactory results. Bornmiiller rightly pointed out that in the section Em- 
pedocleaof the genus Sideritis, as in many of the other genera of Labiatae of the Eastern 
flora (Nepeta, Marrubium, Salvia, Scutellaria, Stachys, not to mention Thymus), 
256 it is often very difficult to present the distinctive features of some of the forms in a brief 
description even though such forms would be acknowledged at a glance as species. It is 
even more difficult to draw up a key that would reflect the distinguishing characters in a 
clear-cut fashion. Frequently, only comparison with the original material ensures reliable 
determination, the primary consideration being, in Bornmiiller's words, "only the plant as 
a whole" (J. Bornmiiller, Neue und kritische Sideritis-Arten (Sectio Empedoclea) der 
vorderasiatischen Flora. Magyar Bot. Lap XXXI, 1932, 128). We may add that investiga- 
tion of extensive material usually helps in the matter. 

Economic importance. The Crimean species of this section are of value as a source of 
essential oil and as ornamentals. They are also used as a substitute for tea (especially the 
highland pasture forms — S. chlorostegia Juz., S. catillaris Juz.). Some of them (e.g. 
S. conferta Juz.) have been reported to contain valuable fixative resins. 

1. S. taurica Steph. ex Willd.Sp. pi. ed. 4, III (1800) 66; M. B. Fl. taur.-cauc. 11,43, 
p. p.; Ldb. Fl. Ross. 111,401, p. p.; Stev. Verzeichn. 281, p. p.; Boiss. Fl. or. IV, 709, 
p. p.; Shmalg. Fl. II, 330, p. p. - S. syriaca Pall. Tabl. phys. et topogr. taur. (1795) 53 
et in Nova Acta Ac. Sc. Petrop. X(1797) 312, saltem p. p. non L. — S. armeniaca |3. 
steveniana Bornm. in Mag. Bot. Lap. 31 (1932) 137. — S. scythica Juz. in Bot. mat. 
gerb. Bot. inst. AN SSSR, XllI (1950) 296, in footnote. - S. steveni Zefirov, Gubotsv. 
(Labiatae) Kryma (1951) 6 (rossice) s. str. - Ic: ?M. B. Centur. plant, rar. Ross, merid. 
(1810) tab. 39. 

Subshrub, with rather long branching woody caudex and brown multicipital root; 
flowering stems one to several, 20-40 cm long; annotinous shoots ascending, rosulate; 
whole plant very densely appressed-tomentose, gray or almost snow-white; radical leaves 
1.5-8.5 cm long, 0.6-1.6 cm wide, oblong-spatulate, gradually tapering toward base, at 
margin obscurely crenulate or (if sparsely pubescent) sharply denticulate, obtuse to sub- 
acute, sparingly hairy, rugulose; petioles 1-2 cm long; cauline leaves (I) 2-7 cm long, 
oblong-obovate to lanceolate, short-petioled to sessile, acute; inflorescence elongate, 
slender, 7-22 cm long, containing (6) 9-16 (18) verticillasters, rather compact or in lower 



170 



(257) 




PLATE XVII. 1 - Sideritis catillaris Juz., general aspect and inflorescence, bract, calyx, corolla; 
2 - S. taurica Steph. (S. scythica Juz.), inflorescence and bract; 3 - S. imbrex Juz., summit of 
stem with inflorescences. 



171 



part interrupted, with lower whorls 2-5 cm apart, the upper forming an uninterrupted 

259 spike; lateral branches, when present, 6-14 (18) cm long, bearing auxiliary inflorescences, 
these 3-8 cm long, distinctly pedunculate, resembling main inflorescence but much smaller; 
floral leaves small, 0.5-1 .3 cm long, 6-9 mm wide, cordate or the lower mostly narrowly 
triangular, gradually (the uppermost often abruptly) acuminate, entire, firm, herbaceous, 
grayish-green, rather densely tomentose, usually broadly white-tomentose-rimmed at mar- 
gin, with indistinct longitudinal nerves, the lowermost as long as or longer than calyx, the 
others mostly shorter (very rarely as long), successive pairs at most contiguous but never 
overlapping; calyx 6-8 mm long, loosely tomentose, the teeth lanceolate, erect, spinescent, 
keeled, about as long as calyx-tube, mostly fully protruding from under the floral leaves; 
corolla 8-10 mm long, at least half as long again as calyx, pale yellow, villous at throat, the 
tube completely enclosed in calyx (calyx-teeth slightly exceeding the corolla-tube), sparse- 
ly villous-tomentose outside in upper part; upper Hp emarginate or shallowly bifid, with 
obtusish semiovate or subtriangular lobes; lower lip reflexed, tripartite; lateral lobes twist- 
ed, narrowly triangular, subobtuse, spreading, the lower lobe broad, rounded-tetragonal, 
slightly notched at apex, concave. June-September. 

Limestone outcrops, taluses, stony steppe plots and slopes, pastures. — European part: 
Crim. (S. Crimea, western end, western part of foothills to Simferopol' inclusive). Ende- 
mic. Described from near Sevastopol' (Khersones). Type was in Berlin; isotypes (includ- 
ing chirotype of Stephan!) in Leningrad. 

Note. The very extensive synonymy of this species indicates, among other things, that 
authors who have recognized more than one species of this section in the Crimea have fail- 
ed to acknowledge the plant described above as the true S. taurica. This oversight is due 
to the fact that, in the Crimea, this plant is by no means the most widely distributed spe- 
cies of the section (it is much less common than S. catillaris Juz.). Another reason is 
that ever since the time of Reichenbach (1826) and well into the present, Bieberstein has 
been accepted as the author of S. taurica. According to him, it was an aggregate species, 
embracing at least three different forms (namely, S. taurica, S. catillaris Juz. and S. 
marschalliana Juz., among which Bieberstein especially set off the latter). In actual fact, 
the author of the name S. taurica is apparently Stephan and the author of the first de- 
scription is Willdenow. In describing the species, Willdenow cites "S. taurica Stephan in 

260 litt." The description itself could hardly fit better the material on S. taurica from 
Stephan's herbarium, preserved in the herbarium of the Botanical Institute of the Academy 
of Sciences of the USSR and labeled "legi in Chers. (oneso)taur.(ico)." It is indeed curi- 
ous that completely identical and unquestionably duplicate material is preserved in the 
same herbarium with the label "Herb. Fischer. (Sideritis) taurica Steph. Spont. Tauria," 
the name of the species being in Fischer's handwriting. It is clear that Stephan distributed 
duplicates of his S. taurica and there is no doubt that Willdenow based his description 
on one of them. 

We should point out that, even though we have included our S. scythica, described 
from the vicinity of Simferopol', in the synonymy of S. taurica Steph., we are not fully 
convinced of the absolute identity of these forms. Both the Sevastopol' plant (i.e. typical 
S. taurica Steph.) and the undoubtedly identical S. steveni Zefir. established from the 
vicinity of Syuren' station, have closely approximate verticillasters, while in S. scythica 
they are exceptionally distant (Plate XVII, Figure 2). The material on S. taurica Steph. 



172 



from the western part of its distribution area is far from sufficient and it is therefore im- 
possible to determine with certainty that the western and eastern forms differ consistently 
in these characters and warrant specific segregation. 

2. S. chlorostegia Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (195 1) 24. - S. tau- 
rica auct. nonnull. fl. taur. p. p. 

Perennial; root very long, woody, brown, caudex multicipital, producing 1 to few 
stems and several abbreviated rosulate sterile shoots (annotinous stems); radical leaves 
2-7 cm long, 0.5-1 .7 cm wide, oblong-spatulate, gradually tapering toward base, obtuse at 
apex, obscurely and finely crenulate, rather densely (or young leaves very densely) covered 
with subappressed or loose tomentum, gray or greenish or initially subniveous; petioles 
1.5-4 cm long; flowering stems (21) 25-50 cm long, suberect, very densely subappressed- 
tomentose, subniveous; cauline leaves 2.5-8 cm long, 0.5-2 cm wide, oblong-obovate to 
lanceolate, short-petioled to sessile, obtusish to subacute, with tomentum rather like that 
on grown radical leaves, often greenish, rugose where tomentum less dense; uppermost 
cauline leaves 2-4.5 cm long, obtuse, mucronate or acute; inflorescence rather slender, 
(6) 9-22 cm long, interrupted in lower part, unbranched or very rarely with subfurcately 
branching stem; verticillasters (5) 8-15(17), the lower always rather distant ((1) 2-4.5 
i261 (5.5) cm apart), the upper approximate to subremote; floral leaves medium-sized, 0.7- 
1 .8 cm long, 0.6-1 .6 cm wide, sessile, cordate, contiguous or mostly somewhat overlapping, 
the lowermost longer, the upper and uppermost shorter than verticillasters, abruptly and 
usually rather long-acuminate, submembranous, green, becoming brown in fruiting plants, 
loosely or very loosely tomentose, rather distinctly reticulate-nerved; calyx 7-9 mm long, 
loosely tomentose or subglabrous, the teeth lanceolate, straight, flexible, slightly patent at 
apex, lanate-villous, mostly exserted; corolla 10-11 mm long, half as long again as calyx, 
pale (sulfur) yellow, with tube enclosed in calyx, otherwise resembling corolla of S. tau- 
rica Steph. June-July. 

Stony mountain slopes, rocky places (mainly limestone). — European part: Crim. 
(Baidar highland pasture and southern coast fromBalaclava to Simeiz). Endemic. De- 
scribed from Baidar highland pasture near Bizyuk survey point. Type and isotypes in 
Leningrad. 

Note. This race occupies an intermediate position between S. taurica Steph. and 
S. catillaris Juz. Its differentiation, at least from the latter,. does not appear to be quite 
definite. Certain forms that occur in Ai-Petri, which we have included in S. catillaris 
Juz., lean slightly toward S. chlorostegia Juz. One of them is a plant which was record- 
ed in "Herbarium Florae Rossicae" as No. 73 (under the name S., taurica M. B.). 

3. S. catillaris Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIII (1950) 296, in adnot. - 
S. incana Habl. Fiz. opis. Tavr. obi. (1785) 148, ex descr. non L. - S. taurica M. B. Fl. 
taur.-cauc.II(1808)43,p.p.; Ldb. Fl. Ross. Ill, 401, p. p.; Stev. Verzeichn. 281, p. p.; 
Boiss. Fl. or. IV, 709, p. p.; Shmal'g. Fl. II, 330, p. p. - Exs.: GRF, No. 73 (saltem huic 
proxima). 

Subshrub, with woody base; root branching, producing ascending or prostrate brown- 
ish caudices, 1 or mostly several stems and also sterile rosettes(reduced annotinous stems); 
"■adical leaves 5-10 cm long, 0.7-2 cm wide, oblong-spatulate, gradually tapering to slightly 



173 



winged petiole 1-3 cm long, obtuse at apex, obscurely crenulate, very densely tomentose 
all over, snow-white or greenish-gray; stems 20-60 cm long, slightly ascending at base, 
otherwise erect, obscurely 4-angled, very densely appressed-tomentose, white or faintly 
greenish; cauline leaves subequal, 2.5-10 cm long, 0.7-2.1 cm wide, oblong-lanceolate to 

262 lanceolate, shortly winged-petiolate to sessile, the lowermost obtuse, others obtuse or 
acute, subniveous or grayish-green, often somewhat rugose where tomentum less dense; 
inflorescence (5) 8-23 cm long, rather thick, consisting of 5-1 1 (13) whorls, in lower part 
usually more or less interrupted, with internodes 1.2-4.2 (5.5) cm long, in upper part 
often spicate; lateral branches wanting or extremely few, with poorly developed inflo- 
rescences of second order; floral leaves medium-sized or the lower rather large, 1.2-2.2 
(2.5) cm long, 1-2 cm wide, broadly cordate, strongly overlapping, at apex abruptly short- 
acuminate, as long as or (the upper) shorter than verticillasters, submembranous, yellowish- 
green, loosely tomentose or subglabrous, with a rather distinct but thin network of veins; 
calyx 0.8-1 cm long, greenish, with arachnoid-tomentose tube, the teeth as long as or 
slightly shorter than tube, lanceolate or narrowly lanceolate, erect, softish, at margin long- 
tomentose, obscurely keeled; corolla 1-1.2 cm long, up to IH times the length of calyx, 
pale yellow, softly subappressed-pubescent outside except for glabrous tube; upper lip 
erect, flat, deeply cleft, with oblong obtuse lobes; lower lip tripartite, reclinate, its lower 
part entire; nutlets ca. 2.5 mm long, flattened, short-ellipsoid or ovoid, 3-angled, blackish, 
dull. July-August. (Plate XVII, Figure 1.) 

Rocks, stony places and slopes, mountain pastures. — European part: Crim. (highland 
pastures, sporadically adjacent part of southern coast). Endemic. Described from Roman- 
Kosh Mountain (Babugan-Yaila). Type in Leningrad. 

Note. This species is the most widely distributed member of the section in the Crimea. 
It comprises an overwhelming majority of specimens preserved in Soviet herbaria under 
the name "S. taurica." It goes without saying that prior to our investigations, the S. tau- 
rica of those authors who tried to distinguish more than one perennial species of Sideri- 
tis in the Crimea (Bornmiiller, Zefirov) corresponds first and foremost to S. catillaris. 

We have not succeeded in elucidating the nature of S. distans Willd. (Sp. pi. Ill (1800) 
66) which was described without indication of provenance. In Prodr. Fl. Balcan. (2 Bd., 
1929, 258), Hayek adopted this name for "S. taurica M. B. which he included in the 
synonymy of S. distans. It has been recently demonstrated (P.H. Davis, Additamenta 
ad Floram Anatoliae, III. Notes fr. the Royal Bot. Card. Edinburgh, Vol. XXI, No. 2, 
1952, 69) that this was done without any justification since both S. taurica and S. 
distans had been published by Willdenow at the same time, the first actually having 
priority in listing. If the material on which S. distans was based originated from the 

263 Crimea (which is very doubtful), then, according to the description, it would most hkely 
correspond to S. catillaris Juz. This, however, is strongly contradicted by the acute 
leaves ascribed to S. distans. 

4. S. marschalliana Juz. sp. nov. in Addenda XIX, 343. - S. taurica M.B. Fl. taur.- 
cauc. II (1808) 43, s. restr. (i. e. quoad pi. e Karassubasar) et auct. plus. fl. taur. p. p.; 
Ldb. Fl. Ross. Ill, 401, p.p.; Boiss. Fl. or. IV, 704, p.p.; Yuz. in Bot. mat. gerb. Bot. 
inst. AN SSSR, XIV, 28, non Steph. ex Willd. - ? S. syriaca Pall. Tabl. phys. et topogr. 
taur. (1795) 53 et in Nova Acta Ac. Sc. Petrop. X (1797) 3 1 2, p. p. - Ic: ? M. B. Centur. 



174 



plant, rar. Ross, merid. (1810) tab. 39 (nisi potius S. taurica vera ?); Rchb. Ic. bot. IV 
(1826) tab. 384, fig. 563 (nom. S. taurica M. B.). 

Subshrub, with robust woody root branching at the top and low caudex covered with 
brown bark and branching from base or summit; flowering stems 1 to (mostly) several, 
25-45 cm long; in addition to stems there are also several abbreviated rosulate, annoti- 
nous shoots; the whole plant, the stems and young leaves in particular, very densely cov- 
ered with appressed white tomentum, gray or (especially stems and young leaves) snow- 
white; radical and lower leaves 2-6 cm long, 0.5-1 .2 cm wide, oblong-spatulate, gradually 
tapering toward base, obscurely crenulate or sometimes (in sparingly pubescent leaves) 
finely serrate-dentate, obtuse to subacute at apex, rugose when indument weakly develop- 
ed; petioles 0.8-2 cm long; cauline leaves (1 .2) 1 .5-7.5 cm long, oblong-oblanceolate and 
(the upper) lanceolate, short-petioled or (the upper) sessile; lower leaves mostly subobtuse, 
the upper subacute to acute; inflorescence elongate-cyhndrical, 5-20 cm long, with (7) 
10-18 (22) verticillasters, rather slender or often fairly thick, somewhat interrupted in 
lower part, with lowest whorl mostly at a distance of 1-2.5 (3) cm from the others, tightly 
contracted above (sometimes throughout) in a dense uninterrupted spike; lateral branches 
wanting or fairly long, bearing mostly short contracted subsidiary inflorescences 4-6 cm 
long on peduncles ca. 7 cm long; floral leaves (except for lowermost which reach or often 
exceed base of the next pair) rather small or medium-sized, 0.6-1.8 cm long, 0.7-1.6 cm 
wide, cordate or broadly cordate, the lowermost more gradually, others abruptly attenuate 
to a rather long or (the upper) short firm spinous point, usually as long as or (in lower 
whorls) exceeding calyx, rather firm, chartaceous, grayish-green, later slightly yellowing, 
264 densely or sparingly tomentose, sometimes subglabrous, with narrow white-tomentose rim 
at margin, or sometimes scarcely rimmed, with more or less distinct longitudinal nerves 
and faintly discernible internerves, two of a pair contiguous at margins or rarely just over- 
lapping in lowermost part; calyx 6.5-9 mm long, loosely tomentose; teeth lanceolate, 
erect, soft, with a distinct dorsal nerve, about equaling, or much shorter than, calyx-tube, 
strongly protruding from under the floral leaves; corolla 0.9-1 .2 cm long, half as long 
again as calyx, pale (sulfur) yellow, villous at throat, the tube completely included in 
calyx, tomentose outside in upper part; upper lip erect, flat, deeply bifid, with oblong 
obtuse lobes ; lower lip tripartite reflexed, the lateral lobes narrow, distant, the lower lobe 
dilated, entire, concave. July. 

Chalk outcrops and taluses, chalky steppes. - European part: Crim. (Belogorsk district). 
Endemic. Described from Ak-Kai cliffs near Belogorsk (former Karasubazar). Type in 
Leningrad. 

Note. In the Bieberstein herbarium there are tags bearing the inscription "Sideritis 
taurica. Ex Taurica" attached to 6 specimens of Sideritis. Two of them refer to the 
above-described plant, two are very similar but most likely represent the true S. taurica 
Steph., and two belong to S. catillaris Juz. of highland pastures. Bieberstein indicated 
Karasubazar as the location of S. taurica ("circa Karassubasar frequens"), which is the 
reason why we earlier accepted the above-described plant as S. taurica. This was of 
course a mistake based on a misunderstanding (at that time we ascribed the authorship of 
S. taurica to Bieberstein, as then generally accepted; see note to the latter species). 

The extensive material that we collected at the classical location of S. marschalliana 
testifies to considerable variability of this species; it fluctuates in its characters between 



175 



S. taurica Steph. (which it approaches more closely) and S. imbricata Juz., and geogra- 
phically occupies an intermediate position between them. We assumed its hybrid deriva- 
tion (op. cit). As far as we know, neither the typical S. taurica nor S. imbricata now 
occurs in the vicinity of Belogorsk, and the Belogorsk form unquestionably deserves a 
specific name, all the more so because the possibility of its being acknowledged as a 
phylogenetically intermediate link between these two species cannot be ruled out. 

5. S. imbrex Juz. nom nov. — S. imbricata Juz. in Bot.mat.gerb.Bot.inst. ANSSSR. 
XIV (1951) 27, non H. Lindb. - S. taurica var. orientalis Zefirov, Gubotsv. Kryma 
(1951) 6 p.p. 

265 A rather low subshrub, with woody root and woody branching caudex with ascending 
or decumbent brown ramifications; sterile shoots short, rosulate; fertile stems 1 to several, 
simple or very often slightly branching in upper part, 12-30 (35) cm long; radical leaves 
2-6 cm long, 0.5-1.5 cm wide, oblong-spatulate, gradually tapering toward base, obtuse or 
very short-acuminate, nearly entire or obscurely crenulate; petioles 1-2 cm long; cauline 
leaves 1.5-6.5 cm long, oblong-oblanceolate or lanceolate, short-petioled or sessile, obtuse 
or the upper acute; all vegetative aerial parts very densely white-tomentose or old radical 
leaves (at base of fertile stems) and cauline leaves (especially the lower) usually less dense- 
ly hairy, bright grayish-green, the least pubescent (green) leaves strongly rugulose; inflores- 
cence 4-9 cm long, ellipsoid or short-cylindrical, rather dense, not interrupted, consisting 
of (5) 8-15 whorls, these very closely approximate or the lowermost whorl at a distance 
of 1-2 cm from others; lateral inflorescences wanting or very often branches borne in the 
axils of uppermost pair (very rarely 2 upper pairs) of cauline leaves developing lateral inflo- 
rescences, these closely approximate to and resembling the terminal inflorescence but 
somewhat shorter, 2-5 cm long; bracts imbricated, 0.8-2 cm long, 1-2.6 cm wide, sessile, 
cordate or broadly cordate, abruptly short-acuminate, the upper generally rounded at apex, 
with small mucro, entire or at times indistinctly incised-crenate; leaves of a pair broadly 
covering each other at margins; calyx 0.8-1 .2 cm long, shallowly tomentose, the teeth 
slightly shorter than tube, erect, narrowly lanceolate, soft, with prominent dorsal nerve, 
protruding to not more than half their length from under bracts or often completely con- 
cealed; corolla 1-1.5 cm long, the tube included in calyx, the limb exserted, pale yellow 
(sulfur-yellow), the upper dilated part of tube rather densely covered with appressed 
long-sericeous hairs. July. (Plate XVII, Figure 3.) 

European part: Crim. (eastern part). Endemic. Described from near Old Crimea 
(Agarmysh Mountain). Type in Leningrad. 

Note. Stephan already noted (Verzeichn. 281) the occurrence of unique forms of 
"S. taurica" with large bracts and closely approximate "whorls." He reported such a 

266 form for Kerchi; however, not one specimen of "S. taurica" from that area is known to 
us though, generally speaking, similar forms grow apparently in eastern Crimea. We have 
differentiated the above-described form, S. imbrex Juz. growing near Old Crimea, and a 
form growing at the southern coast of eastern Crimea (see above) as separate taxonomic 
units (even though they are very closely allied). 

6. S. conferta Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 26. - S. tauri- 
ca var. orientalis Zefirov, Gubotsv. (Labiatae) Kryma (1951) 6 (rossice) p.p. 



176 



Perennial, very similar to preceding species; stems 10-30 (36) cm high, usually simple; 
inflorescence slightly less dense, (2) 5-12 cm long, with (2) 5-9 (12) verticillasters, the 
lowermost 1 or 2 (3) often at a distance of up to 3 cm (though usually less) from the 
others; lateral inflorescences rarely present and then generally poorly developed; bracts 
of successive whorls not overlapping (the lower) or sUghtly overlapping, somewhat larger 
compared with S. imbrex, 1-2.6 cm long, 1.2-3 cm wide, gradually (the lower) or abrupt- 
ly (the upper) tapering at apex to larger, often slightly curved, nearly spinous mucro, some 
with few remote or soUtary fine obtuse or acute teeth at margin, two of a pair very broad- 
ly overlapping at margins; flowers slightly larger than in S. imbrex; calyx 8-12 mm long, 
the teeth often completely protruding from under the floral leaves; corolla 12-14 mm 
long; otherwise, similar to preceding species. June-July. 

Stony mountain slopes, limestone outcrops. — European part: Crim. (E.). Endemic. 
Described from vicinity of Karadag and Sudak. Type and paratype in Leningrad. 

Note. This and the preceding species are not always easily distinguished from each 
other. This is quite natural inasmuch as their extremely restricted distribution areas ad- 
join each other. However, the differences between them strike the eye when extensive 
material is examined. 

7. S. euxina Juz. in Bot. mat. gerb. Bot. inst. AN SSSR, XIV (1951) 28, in footnote. - 
S. taurica Lipsky, Issledov. Severn. Kavk. (1889-1890) 33-34; idem, Fl. Kavk. 423; 
Shmal'g. Fl. II, 330, p. p. et auct. plur. fl. Cauc. non Steph. nee M. B. — S. taurica var. 
glandulosa Albov. Prodr. Fl. Colch. (1895) 202. - S. steveni Zefirov, Gubotsv. (Labia- 
tae) Kryma (1951) 6 (rossice) p. p. (quoad pi. e distr. Novorossijsk). — Exs.: Fl. Cauc. 
exs. No. 145. 

Subshrub, with woody root and branching caudex; stems usually numerous, sturdy, 
ascending at base, 18-50 cm long; sterile shoots abbreviated, rosulate; whole plant gray 
267 with dense appressed or (especially in mature leaves) slightly flocculent white tomentum; 
radical and lower cauUne leaves 3-8 cm long, 0.5-1.5 cm wide, oblong-spatulate, gradually 
tapering toward base, mostly obscurely crenulate-serrulate, mostly subacute, rugose where 
sparingly pubescent; petioles 0.5-2 cm long; cauline leaves 2.5-7 cm long, oblong-oblan- 
ceolate to broadly lanceolate, short-petioled, the uppermost subsessile, acute, often non- 
duplicate and arched; inflorescence cylindrical, (4.5) 7-16 cm long, containing (5) 10-18 
whorls, rather thick, in lower part shghtly interrupted with lowermost whorl at a distance 
of 1 .5-3 cm from others, contracted above often throughout in uninterrupted spike; 
lateral branches often present, short or fairly long, bearing well developed auxiliary inflo- 
rescences, these (3) 4-8 cm long, compact or below slightly interrupted, on peduncles 0.5- 
7 cm long; floral leaves 0.8-1 .7 cm long, 1 .2-2 cm wide, thin, chartaceous to submembra- 
nous, pale grayish-green or faintly yellowish, mostly densely or very densely tomentose, 
with prominent longitudinal and short transverse nerves, often obscurely crenate, broadly 
cordate, abruptly attenuate to a short (or lowermost to a long), slightly recurved point, 
usually equaUng calyx, two of a pair usually widely overlapping at margins; calyx 11- 
12 mm long, heavily but loosely tomentose-villous all over, often with copious admixture 
of small short-stipitate glands; teeth erect, lanceolate, soft, with obscure dorsal nerve, 
about equahng or slightly shorter than tube, fully or mostly slightly protruding from 
under the floral leaves; corolla 14-15 mm long, small to half as long again as calyx, pale 



177 



yellow, densely villous at throat, the tube completely included in calyx, the upper part of 
the tube and nearly the whole limb densely sericeous-tomentose outside. June-August. 

Stony dry sunny slopes, mountain steppes, limestone rocks. Caucasus: Cisc. (Anapa 
and others), W. Transc. (Novorossiisk region). Endemic. Described from near Novorossi- 
isk. Type in Leningrad. 

Note. N.A. Busch was mistaken in stating that "S. taurica" was first found in the 
Novorossisk region by Lipskii. This statement appears on a tag attached to Duz's plant 
from near Novorossiisk distributed from the herbarium of the Yur'ev Botanical Garden. 
268 In the herbarium of the Boianical Institute of the Academy of Sciences of the USSR there 
are specimens of S. euxina collected in Novorossiisk in 1841 (probably by Radozhitskii). 
The plant was also collected by Bayern. There is no question but that neither Bieberstein 
nor Boissier knew of the existence of S. euxina; hence Kolakovskii served no purpose in 
placing S. taurica M.B. p.p. and S. taurica Boiss. p.p. in the synonymy ofS. euxina 
Juz. (Spisok rast. gerb. fl. SSSR, XII, 1953, 1 10). 

Albow's "variety" was probably based on specimens with generally depauperate tomen- 
tum, in which the glandular vesture of the calyx tends to be more conspicuous. 



Section 2. Hesiodia (Moench, pro gen.) Benth. Lab. gen. et sp.( 1834) 272. -Annuals. 
Floral leaves resembling cauline leaves but smaller, entire or weakly crenate, obtuse; calyx 
2-lipped, the 3 upper teeth connate at base. 

Note. In contrast to the preceding, the species of this section are distributed through- 
out the Mediterranean region. 



Series 1 . Man tanac Juz. Flowers bicolor, yellow with brown, on very short pedicels. 

8. S.montanaL. Sp.pl. (1753)575; Ldb. Fl. Ross. Ill, 401, p. p.; Shnial'g. Fl. II, 
330, saltem pro max. parte. - S. hirsuta Eichw. PI. nov. min. cogn. casp.-cauc. (1831- 
1833) 24, non L. - ? S. romana Georgi, Beschr. Russ. R. Ill, 5 (1800) 1081, non L. - 
Hesiodia bicolor Moench, Meth. pi. (1794) 392. - H. montana Dum. Fl. Belg. (1827) 
44. - Burgsdorfia montana Rchb. Fl. Germ. exc.(1830) 327. - Ic: Rchb. Ic. Fl. 
Germ. XVHI, 15 (1856) tab. 1226; Hegi, III. Fl. Bd. V, 4, fig. 3271 et 3272 (phot.). - 
Exs.: Rchb. Fl. Germ. No. 1236; Schultz, Herb. norm. No. 2248; Woloszcz. Fl. pol. 
No. 980; Baenitz, Herb. Eur. (e Nagy Enyed a. 1894 sine No.); Fl. exs. Reip. Bohem.- 
Slov.No.356. 

Annual; root curved, llexuous; stems 10-50 cm long, erect or ascending at base, simple 
or slightly branching at middle, patent-tomentose -villous, with internodes 1-3 (5) cm long; 
leaves 1-4 (5)cm long, 0.25-1 ( 1 .5)cm wide, spatulate or elliptic, obtuse to subacute or acu- 
minate, sliort-petioled, the upper subsessile, with 1-3(5) longitudinal lateral nerves on each 
side produced into spiny-tipped teeth, otherwise entire, dull yellow on both sides, covered 
with long, mostly sparse hairs; flowers short-pediceled, in very numerous, mostly 6-flower- 
ed whorls, forming long, interrupted moderately dense spikes; bracts not or slightly differ- 
271 ing from cauline leaves, mostly longer than flowers; summit of stems without bundle of 
bright green terminal leaves; calyx rigid, prominently nerved, slightly accrescent, 8-1 1 mm 



178 



(269j 




PLATE XVIII. 1 Sideritis montana L., general aspect, flower, corolla, fruiting calyx; 2 - S. co- 
mosa (Rochel) Stank., summit of stem, flower, corolla, fruiting calyx; 3 - S. balansae Boiss., general 
aspect, flower, fruiting calyx. 



179 



long in fruit, the tube subcylindrical or narrowly campanulate, with constriction in upper 
part, 10-nerved, loosely villous-hairy at throat; teeth ovate, villous, suberect to sHghtly 
patent, spiny-tipped, becoming fully spreading (but not curved), 3 connate into upper lip 
as long as tube; corolla shorter than cahoc, pale yellow, the limb reddish-brown close to 
margin, becoming very dark brown (nearly black) after flowering and on drying; tube ca. 
3 mm long, included in calyx; upper Up slightly convex, ca. 1 .5 mm long; lower lip only 
ca. 1 mm long, with very small lateral lobes; stamens ca. 1.5 mm long, slightly hairy; nut- 
lets 1 .5 mm long, 3-angled, grayish-brown, rounded at apex and slightly foveolate-punctate, 
otherwise smooth. June-September. (Plate XVIII, Figure 1.) 

Impoverished meadows, rocky places, steppes, wastelands, stony (often limestone) and 
sandy soil. — European part: U. Dnp., M. Dnp., V.-Don, Bes., Bl., L. Don, Crim. (mainly 
in northern part and foothills of mountainous Crimea); Caucasus: universal; Centr. Asia: 
Mtn. Turkm. (Kopet-Dagh). Gen. distr.: Med., Bal.-As. Min., Iran. Described from Italy. 
Type in London. 

Economic importance. A noxious weed. There are reports that it is poisonous to horses. 

9. S. comosa (Rochel) Stank. Opred. vysh. rast. Evrop. ch. SSSR (1949) 861. - S. 
montana ^. comosa Rochel ex Benth. in DC. Prodr. XII (1848) 446; Boiss. Fl. or. IV, 
707. — S. montana Ldb. Fl. Ross. I, 401 et auct. plur. fl. Ross. p. p. - Exs.: GRF, 
No. 525; Fiori et Beg. Fl. Ital. No. 1941. 

Annual; stems 15-40 cm long, simple or often branching from base, covered with long 
spreading or erect-spreading hairs, often becoming intensely red; leaves 0.6-3.5 cm long, 
0.1-0.6 cm wide, oblanceolate, gradually tapering to short petiole, the upper sessile, most- 
ly acute, obscurely toothed, grayish-green on both sides, mostly densely covered with long 
spreading or loosely appressed hairs; floral leaves markedly shorter and slightly wider than 
cauline leaves, often only slightly exceeding flowers, tlie uppermost without flowers in 
their axils, usually enlarged, lemon-yellow or golden-yellow, forming a bundle or coma at 
summit of stem; flowers in very distant verticillasters; calyx 5-7 mm long, the tube dilat- 
272 ed above, obconical or campanulate, nearly without constriction, the teeth lanceolate, 
suberect-patent and slightly recurved, shorter than tube; corolla small, but limb slightly 
larger than in S. montana, yellow with brown. Otherwise, similar to S. montana L. 
June-August. (Plate XVIII, Figure 2.) 

Dry, mostly swardless slopes, stony places, steppes. - European part: Bes., Bl. (rarely), 
Crim. (mountainous part, very common); Caucasus: Cisc, W. and E. Transc. Gen. distr.: 
Med., Bal.-As. Min. Described from Dalmatia, Rumelia and Asia Minor. Type unknown. 

Note. Up to recently S. comosa was considered as a form of little taxonomic impor- 
tance. Boissier did not even list it as a variety. Gams regarded it (wrongly without doubt) 
as a seasonal fall modification (cp. Hegi, III, Fl. V, 4, 2405). However, our discovery of a 
number of other distinctive characters, besides the basic character of the yellow color of 
the bracts crowning the inflorescence, and a different geographical distribution area, led us 
to agree with Stankov, the first to promote this form to the rank of species. Our observa- 
tions of S. com osa, mainly in the Crimea, gave us the impression of an aboriginal species 
widespread in the mountainous (southern) part of the peninsula. Conversely, S. montana 
appears to have penetrated into this area recently (from the north) and is far from being a 
synanthropic form. Intermediate forms between these two species often occur in the 
Crimea, but apparently only where they both grow together. 

180 



Series 2. Balansaeanae Juz. - Flowers purple (sometimes whitej, more or less dis- 
tinctly pediceled. 

10. S. balansae Boiss. Diagn. Ser. II, No. 4 (1859) 35; id. in Fl. or. IV, 707. - S. wo- 
ronowii Schischk. ex Grossh. Fl. Kavk. Ill (1932) 290; Grossg. Opred. rast. Kavk. 330. - 
S. purpurea auct. nonnull. fl. Cauc. non Talb. 

A small annual, pale green, 2-1 5 cm high; stems erect or ascending at base, simple or 
branching from base with short and stout branchlets, sparsely covered with long spreading 
hairs, often suffused with red in lower part; leaves to 1.5 cm long, 0.8 cm wide, eUiptic or 
oblong, obtuse, with few markedly asymmetrical acute teeth, sparsely or fairly densely 
hairy, rather long-petioled; inflorescence elongate, dense, the internodes shorter than 
calyx; floral leaves short-petioled, acute, otherwise resembling cauline leaves, as long as 
or barely exceeding flowers; verticillasters few or fairly numerous, 2-6-flowered; flowers 
273 on distinct pedicels to 2-3 mm long; calyx 5-9 mm long, prominently nerved, patent- 
hairy, sometimes slighdy reddening, the tube becoming dilated in lower part, constricted 
in upper part; calyx-teeth as long as or slightly shorter than tube, erect, ovate-lanceolate, 
abruptly attenuate to short mucro; corolla pinkish-purple (rarely white), almost included 
in calyx or slightly exserted; nutlets 1 -1 .5 mm long, grayish. May-July. (Plate XVIII, 
Figure 3.) 

Dry stony slopes. - Caucasus; S. Transc. (Armenian SSR, Nakhichevan ASSR). Q^n. 
distr.: Bal.-As. Min. (As. Min.). Described from Kilikia. Type in Geneva. 

Note. We have been unable to distinguish the Transcaucasian plant of this type (i. e. 
S. woronowii Schischk. ex Grossh.) from the original S. balansae Boiss. Grossgeim, 
too, in his original description of S. woronowii, mentioned no differences between them. 

S. balansae Boiss. is readily distinguishable from the Greek S. purpurea Talb., with 
which it was at times confused in the Soviet Union, by its low stem, short branches, 
scantier vesture throughout, approximate verticillasters, more pronounced dentation of 
floral leaves (often discernible even in upper bracts), differently shaped calyx, with nar- 
rower teeth and weakly developed mucro, smaller corollas. The uppermost tooth of the calyx 
of S. purpurea is larger and wider than the three lower teeth, these narrow and acumi- 
nate. As regards this character, it should be included in another section — Burgsdorfia 
(Moench) Briq. 



Tribe 2. NEPETEAE Benth. Lab. gen. et sp. (1834) 462; id. in DC. Prodr. Xll, 368; 
Briq. in Pflanzenfam. IV, 3a (1895) 207 et 233. - Calyx 5-toothed or 2-lipped, the lips 
dentate; corolla with exserted tube and concave upper lip; stamens 4, didynamous, the 
upper longer than the lower, ascending parallel to each other and lying below upper lip; 
stamens and style not included in corolla-tube. 



Genus 1251.* Agastache** Clayt. ex Gronov. 

Gronov. Fl. Virgin. (1762) 88; O. Ktze. Rev. gen. II (1891)511. - Vleckia Raf. in Med. Repor. New- 
York V (1808) 352. - Lophanthus sect. Chiastandra Benth. Lab. gen. et sp. (1834) 462. 

* Treatment by A.I. Poyarkova. 
** I'rom the Greek aga. many, and stachys, spike. 

181 



Calyx 15-nerved, tubular-obconical, straight, the throat oblique, without hairy ring 
inside; corolla with erect tube, gradually and slightly dilated toward limb, equaling calyx 

274 or very slightly exserted, without hairy ring inside; upper lip erect, bifid; lower lip tri- 
partite, the lateral lobes small, erect, and the middle lobe large, spreading, sessile (clawless), 
broad, crenate or undulate; all 4 stamens perfect, much longer than corolla, the upper 
curved forward, the lower upright; anther-cells at first nearly parallel, later slightly diverg- 
ing; style witli 2 short subequal lobes; disk regular, obscurely lobed; nutlets smooth, 
densely hairy at apex, attached by its basal part to the flat top of the disk at the gaps be- 
tween its lobes and close to its center. Tall perennial herbs, with toothed petiolatc leaves 
and small bisexual flowers; verticillasters forming terminal spikelike inflorescences. 

One species in E. Asia and about 8 in North America. 

1. A. rugosa (Fisch. et Mey.) O. Ktze. Rev. gen. 11(1891) 51 1. - Lophanthus rugo- 
sus Fisch. et Mey. in Ind sem. hort Petrop. I (1835) 31 ; Benth. in DC. Prodr. XII, 369; 
Maxim. Prim. fl. Amur. 218; Kom. Fl. Man'chzh. Ill, 349; Kom. and Alls. Opred. rast. 
Dal'nevost. kraya, II, 899. - Ic: Kom. and AHs., op. cit. Plate 272. 

Perennial; root robust, woody, obhque; stems to 1 mm [sic J long, to 7-8 mm thick, 
covered with very short appressed hairs, in upper part with leafy flowering branches; 
leaves thin (dry leaves chartaceous and more or less warped), to 8-9 (12) cm long and 
5-7 (8) cm wide, ovate or oblong-ovate, with cordate or rarely rounded base, acuminate 
or terminating in a long attenuate point, coarsely toothed, with very short appressed hairs 
on both sides and numerous punctate glands beneath; petioles 1/4-2/5 as long as blade; 
verticillasters crowded at end of stem and of axillary branches in dense cyhndrical inflo- 
rescences, these reaching 10 cm in length and 2 cm in width on the stem; terminal leaves 
at base of inflorescence similar to cauline leaves but smaller; floral leaves subtending the 
semiwhorls up to 5 mm long and ca. 1-1.2 mm wide, lanceolate-linear, long-acuminate; 
bracts similar but approximately half as long; pedicels 1-2 mm long, all densely covered 
with short spreading hairs; calyx (5) 7-8 mm long, more or less suffused with lilac or 
purple-lilac, sparsely covered with short simple spreading hairs (longer only at margin of 
teeth) and beset with sparse capitate white glandular hairs, the throat distincfly oblique; 
teeth lanceolate, the upper ca. 1/2 (2/5) as long as tube, the lower shorter; corolla (6.5) 
8-9.5 (10) mm long, bluish-lilac, puberulent above, the tube scarcely longer than calyx; 
upper Up 1 .25-1 .5 mm long, cleft approximately to middle into ovate lobes; middle lobe 

275 of lower Up strongly dilated, ca. 2 mm long, 3-3.5 mm wide, slightly notched at apex, with 
slightly undulate margin, the lateral lobes triangular, 0.3-0.5 mm long, 1 .3-1 .5 mm wide; 
stamens exserted from corolla for 3-5 mm; nutlets dark brown, obovoid, ca. 1 .5 mm long, 
0.6-0.7 mm wide, 3-angled, with obtuse apex, beset Uke ribs with upright rufous hairs. 

Fl. July-August, fr. August-September. (Plate XIV, Figure 2.) 

Rocks, rocky grass-covered slopes, often in shade of trees and shrubs, in small groups. - 
Far East: Uss. (southern part, in the north Bureinskii Range and near Khabarovsk). Gen. 
distr.: Jap. and Ch., Tib. (provinces in central and eastern China). Described from China. 
Type in Leningrad. 

Economic importance. This species is worthy of investigation as a source of essential 
oil, since all plant parts have a strong, pleasant peppermint scent. 



182 



Genus 1252.* Lophanthus** Adans. 

Adans. Fam. PI. II (1763) 194, 572, p. p.; emend. Benth. in Bot. Reg. XV (1829) post tab. 1282; 
emend. Briq. in Pflanzenfam. IV, 3a (1897) 234; Levin in Tr. Bot. inst. AN SSSR, V (1940) 268. - 
Vleckia Rafin. Fl. Tellur. Ill (1808) 89, p.p. - Sect. Psilonepeta Benth. gen. Nepeta L. in DC. 
Prodr. XII (1848) 392. 

Calyx tubular or tubular-campanulate, straight or slightly curved, 5-toothed, with 
straight or more or less oblique throat, rarely 2-lipped, 1 5-nerved, with hairy ring inside; 
corolla straight or curved; the tube exserted, dilated above, twisted; limb inverted, the 
upper (here the lower) lip 2-lobed, the lower (here the upper) lip 3-lobed, the large middle 
lobe broader than long; anther-cells nearly parallel or slightly diverging; pistil parted 
above into 2 thin lobes; nutlets oblong-ovaloid, slightly compressed, rounded on one side, 
on the other bilaterally tapering, smooth, brown. Perennial herbs; leaves oblong to broad- 
ly ovate, crenate-dentate or dentate; floral leaves usually small; stems above and inflores- 
cence usually leafless; cymes in axils of upper leaves; bracts small, linear-lanceolate, rarely 
broadly lanceolate. 

1 . Leaves with very prominent veins beneath, mostly plicate-rugose, to 1 5 mm long . 
2. 

+ Leaves different 3. 

2. Plant gray-tomentose, sometimes nearly white, rarely more or less green; leaves 
crisp-crenate ; calyx nearly 2-lipped; corolla 10-15 mm long; 2 stamens long- 
exserted 7. L. subnivalis Lipsky. 

276 + Plant short-hairy, grayish; leaves large, nearly incised-dentate; calyx straight at 

throat; corolla 1 5-18 mm long; stamens not exserted or slightly so 

8. L. elegans (Lipsky) Levin. 

3. All stamens not or scarcely exserted; calyx-teeth triangular-lanceolate; middle pe- 
duncles 7-12 mm long; plant 30-35 cm high; leaves broadly ovate, cordate at base, 
10-20 mm long; petioles 1-6 mm long 4. L. tschimganicus Lipsky. 

+ Two stamens long-exserted and 2 scarcely or not exserted 4. 

++ All stamens long-exserted (very rarely 2 included); petioles 5-20 mm long; middle 
peduncles 15-20 mm long; inflorescence usually leafless; floral leaves usually linear- 
lanceolate 6. 

4. Plant covered with rather long stiffish hairs or subglabrous; leaves cordate at base; 
calyx tubular-campanulate, usually much dilated above, 8-1 1 mm long, nearly 
oblique at throat, the teeth mostly broadly oval-lanceolate; corolla 13-20 mm long 
3. L. schrenkii Levin. 

+ Plant subglabrous or puberulent (rarely almost tomentose); leaves at base rounded, 
truncate, obtuse, rarely cordate; calyx 6-10 mm long, oblique at throat or nearly 
2-hpped, the teeth triangular-lanceolate; corolla 10-15 mm long 5. 

5. Calyx tubular, slightly dilated above, nearly 2-lipped, 6-8 mm long; corolla bluish- 
lilac; middle peduncles 3-7 mm long; leaves 10-25 mm long 

2. L. krylovii Lipsky. 

* Treatment by E.G. Levin. 
** From the Greek lophos, comb, helmet, and anthos, flower. 



183 



+ Calyx nearly tubular-campanulate, mostly markedly broadening above, more or less 

oblique at throat, 6-10 mm long; corolla blue; middle peduncles 8-12 mm long; 

leaves 15-40 mm long 1. L. chinensis (Rafm.) Benth. 

6. Calyx tubular-campanulate, the teeth oval-lanceolate or lanceolate 

5. L. schtschurowskianus (Rgl.) Lipsky. 

+ Calyx tubular, teeth narrowly lanceolate, rarely triangular-lanceolate, acute, often 

point-tipped 6. L. lipskyanus Ik.-Gal. et Nevski. 

1. L. chinensis (Rafin.) Benth. in Bot. Reg. XV (1829) post. tab. 1282; id in DC. 
Prodr. XII,369,p.p.; Ldb. Fl. Ross. Ill, 372, p. p.; Turcz. Fl. baic.-dahur. II, 401; Levin 
inTr. Bot.inst. ANSSSR, 1,V, 271. - L. obtusifolius Heyn. Nom. bot. hort. (1840) 
477. - Hyssopus lophanthus L. Sp. pi. (1753) 569. — H. resupinatus Moench, Meth. 
suppl. (1802) 134. -Vleckia chinensis Rafin. Fl. Tellur. Ill (1818) 89. -Nepeta lo- 

277 phantha Fisch. eX Benth. Lab. gen. et sp. (1834) 464. — Agastache lophanthus 0. 
Ktze. Rev. gen. 11 (1891) 51 1. - Ic: Jacq. Hort. vind. II (1772) tab. 182. 

Perennial, puberulent or subglabrous, glandular, green; stems usually many, 20-50 cm 
long, often divaricately branching; leaves ovate or oblong, obtuse or acute, at base round- 
ed, truncate or subcordate, crenate-dentate, 15-40 mm long, 10-25 mm wide; petioles 
2-8 mm long; floral leaves small, linear-lanceolate, approximately as long as cymes or the 
upper shorter, rarely longer; cymes in axils of upper leaves forming lax inflorescences; 
middle peduncles 8-1 2 mm long; calyx subtubular-campanulate, nearly straight or slightly 
curved, more or less obhque, 6-10 mm long, with hairy ring inside, teeth triangular-lanceo- 
late, 1/4-1/3 as long as tube, the upper 3 slightly broader; corolla blue, 10-15 mm long, 
about twice the length of calyx; corolla-tube exserted, dilated above, twisted, the limb 
inverted; upper (here the lower) lip 2-lobed; lower (here the upper) lip 3-lobed, the mid- 
dle lobe larger, wider than long, transversely notched, finely toothed, the lateral lobes 
smaller, broadly oval; upper (here lower) stamens and pistil long-exserted; lower (here 
upper) stamens not or scarcely exserted; anther-cells slightly diverging; pistil divided 
above into two thin lobes; nutlets oblong-ovaloid, slightly constricted, smooth, brown. 
June-August. 

Stony slopes, cliffs, debris, ravines, steppes and meadows. — East Siberia: Ang.-Say., 
Dau. Gen. distr.: Mong., Ch. (Hopei). Described from China. Type in London. 

2. L. krylovii Lipsky in Tr. Bot. sada, XXIV (1905) 122; Kryl. Fl. Zap. Sib. IX, 2305; 
Levin in Tr. Bot. inst. AN SSSR, V, 274. 

Perennial, puberulent, glandular, green to grayish (f. glabrata Kryl.) or whitish-gray, 
with nearly tomentose leaves (f. canescens Kryl.); stems numerous, 20-30 cm long, 
erect or slightly ascending, divaricately branching almost from base, the branches nearly 
as long as stems; leaves ovate or ovate-triangular, obtuse to subacute, at base subcordate, 
obtuse or truncate, crenate-dentate, 10-25 mm long, 8-20 mm wide; petioles 3-8 mm 
long; lower leaves much smaller; floral leaves generally small, linear-lanceolate, usually 
shorter than cymes, rarely longer; cymes in axils of upper leaves, forming loose inflores- 
cences, these subsessile or short-peduncled; middle peduncles 3-7 mm long; calyx tubu- 

278 lar, sHghtly dilated above, erect, nearly 2-lipped, 6-8 mm long, with hairy ring inside; 
teeth triangular-lanceolate, 1/4-1/3 as long as tube, the upper slightly broader; corolla 



184 



bluish-lilac, curved, 10-15 mm long, about twice the length of calyx; corolla-tube sHglitly 
exserted, gradually dilated above, twisted, the limb inverted; upper (here lower) lip 2- 
lobed, with broadly oval lobes, lower (here upper) lip 3-lobed, the middle lobe larger, 
wider than long, notched, finely toothed, the lateral lobes smaller, broadly oval; upper 
(here lower) stamens and pistil long-exserted, the lower (here upper) not or slightly ex- 
serted; anther-cells slightly diverging; pistil divided above into two thin lobes; nutlets 
oblong-ovaloid, slightly compressed, smooth, brown. June-August. 

Alpine belt, dry stony slopes and plateaus, placers and glacial moraines. — West Siberia: 
Alt. (especially Altai and Naryn Range); Centr. Asia: Dzu.-Tarb. (Tarbagatai, Saur). Gen. 
distr.: Mong. (Mongolian Altai). Described from Altai (Argut River). Type in Leningrad. 

3. L. schrenkii Levin in Bot. mat. gerb. Bot. inst. ANSSSR, VII, 10 (1937-1938) 218; 
ibid, Tr. Bot. inst. AN SSSR, V, 276. - L. chinensis Benth. in DC. Prodr. XII (1848) 
369, p. p.; Ldb. Fl. Ross. Ill, 372, non Benth. - Hyssopus lophanthus Ldb. Fl. alt. 
Ill (1830) 398, non L. - Ic: Bot. mat. gerb. Bot. inst. ANSSSR, VII, 10, 219. - Exs.: 
HFAM,No.204. 

Perennial, hairy or subglabrous, green, very rarely grayish, glandular; stems erect or 
slightly ascending at base, 20-40 (80) cm long, branching; leaves ovate, cordate at base, 
obtuse or more or less acute, crenate-dentate, 15-35 (50) mm long, 10-20 (40) mm wide, 
the petioles 3-15 (30) mm long; upper leaves more or less rounded at base; floral leaves 
shorter than cymes, often much shorter, small, linear-lanceolate, broadly or narrowly 
ovate; cymes in axils of upper leaves, forming lax inflorescences, these subsessile or short- 
peduncled; middle peduncles 2-1 mm long; calyx tubular-campanulate, mostly much 
dilated above, straight or slightly curved, somewhat oblique, 8-1 1 mm long, with hairy 
ring inside; teeth broadly oval -lanceolate, rarely triangular-lanceolate, 1/4-1/3 as long as 
tube, the upper three slightly broader; corolla 13-20 mm long, nearly twice the length of 
calyx, the tube exserted, dilated above, twisted, the limb inverted; upper (here lower) lip 
2-lobed, lower (here upper) Up 3-lobed, the middle lobe larger, broader than long, trans- 
versely notched, finely toothed, the lateral lobes smaller, broadly oval; upper (here lower) 
stamens not or slightly exserted; anther-cells slightly diverging; pistil long-exserted, divid- 
ed into two thin lobes; nutlets oblong-ovaloid, slightly compressed, smooth, brown. 
May- July. 

Rocks, stones, shale taluses, stony slopes, grass plots. — Centr. Asia; Balkh. (Kent and 
Bektau-Ata mountains), T. Sh. (Kara-Tau, Talass Ala-Tau, Chatkal Range, Kirghiz Ala-Tau, 
Chu-Ili mountains), Dzu.-Tarb. (Dzungaria-Ala-Tau, Saur). Gen. distr.: Dzu.-Kash. (E. 
T. Sh.). Described from Bektau-Ata mountains. Type in Leningrad. 

4. L. tschimganicus Lipsky in Tr. Bot. sada, XXIII (1904) 212; Levin in Tr. Bot. inst. 
ANSSSR, 1,V, 277. 

Perennial, short-hairy, glandular; green; stems many, erect or slightly ascending, 30- 
35 cm long, branching; leaves broadly ovate, usually obtuse, cordate at base, crenate-dentate, 
10-20 mm long, 6-15 mm wide; petioles 1-6 mm long; lower leaves often subreniform, 
small, obtuse or tapering at base; uppermost leaves subsessile, often linear-lanceolate; 
cymes in axils of upper leaves, forming lax peduncled inflorescences; middle peduncles 
7-1 2 mm long; bracts small, linear-lanceolate; calyx tubular or tubular-campanulate, erect. 



185 



slightly oblique, 7-9 mm long, with hairy ring inside, the teeth triangular-lanceolate, 1/4 
to 1/3 as long as tube, the upper 3 slightly broader, corolla dingy lilac, 13-18 mm long, 
about half the length of calyx; tube much exserted, slightly dilated above, twisted, the 
limb inverted; upper (here lower) hp 2-lobed; lower (here upper) lip 3-lobed, the middle 
lobe larger, broader than long, transversely notched, finely toothed, the lateral lobes 
smaller, broadly oval; all stamens included or nearly so; anther-cells slightly diverging; 
pistil long-exserted, divided above into two thin lobes; nutlets oblong-ovaloid, shghtly 
compressed, smooth, brown. June-August. 

Grassy slopes. - Centr. Asia: T. Sh. (Tashkent Ala-Tau: Chimgan, Chatkal River). 
Endemic. Described from Chimgan. Type in Leningrad. 

5. L. schtschurowskianus (Rgl.) Lipsky in Tr. Bot. sada, XVIII (1901) 93; Levin in 
Tr. Bot. inst. AN SSSR, ser. 1, V, 279. — L. schtschurowskianus ^. kulabensis Lip- 
sky, 1. c. p.p. — Nepeta schtschurowskiana Rgl. in Izv. Obshch. lyub. est. antr. i etn. 
XXXIV, 2 (1882) 63, 66. - Nepeta ourumitanensis Franch. in Ann. Sc. Nat. ser. VI 
XVIII (1884) 230. 

Perennial, subglabrous or puberulent, glandular, green; stems erect, rarely ascending, 
30-70 cm long, branching; leaves ovate-cordate or ovate-oblong, usually rounded at base, 
280 crenate-dentate, 20-40 mm long, 10-30 mm wide; petioles 5-20 mm long; floral leaves 
small, linear-lanceolate; inflorescence leafless, rarely all leaves large and then inflorescence 
leafy; cymes in axils of upper leaves, forming lax long-peduncled inflorescences; middle 
peduncles 15-20 mm long; calyx tubular-campanulate, straight, very slightly obhque, 8- 
10mm long, with hairy ring inside; teeth oval-lanceolate or lanceolate, 1/4 to 1/3 as long 
as tube, the upper 3 slightly broader; corolla dingy lilac (?), 15-20 mm long, 2 to 3 times 
the length of calyx, the tube exserted, dilated above, twisted, the limb inverted; upper 
(here lower) lip 2-lobed; lower (here upper) Up 3-lobed, the middle lobe larger, broader 
than long, transversely notched, finely toothed, the lateral lobes much smaller, broadly 
oval; all 4 stamens long-exserted, rarely only 2 or none; anther-cells slightly diverging; 
pistil long-exserted, divided into two thin lobes; nutlets oblong-ovaloid, slightly com- 
pressed, smooth, brown. May-July. 

Mainly in Central Asian juniper belt, grassy slopes, scrub, dry stony slopes, dry river- 
beds, alpine stands, 2 100-3000 m. — Centr. Asia: Pam.-Al. (Turkestan Range, basin of 
Zeravshan, Malyi Alai, Alai Range, Trans-Alai Range), T. Sh. (Fergana Range). Described 
from Alai Range. Type in Leningrad. 

6. L. lipskyanus Ik.-Gal. et Nevski in Tr. Bot. inst. AN SSSR, 1, IV (1937) 325; Levin, 
ibid. V (1940) 281. — L. schtschurowskianus j3. kulabensis Lipsky in Tr. Bot. sada, 
XVIII (1901) 83, p.p. 

Perennial, plant subglabrous or puberulent, glandular, green; stems erect, rarely ascend- 
ing, 30-70 cm long, branching; leaves ovate-cordate or ovate-oblong, usually rounded at 
base, crenate-dentate, 20-40 mm long, 10-30 mm wide; petioles 5-20 mm long; floral 
leaves very small, linear-lanceolate; inflorescence leafless, rarely all leaves large and then 
inflorescence leafy; cymes in axils of upper leaves, forming loose, long-peduncled inflo- 
rescences; middle peduncles 15-20 mm long; calyx tubular, straight, very slightly oblique, 
8-9 mm long, with hairy ring inside; teeth narrowly lanceolate, rarely triangular-lanceolate. 



186 



acute, often point -tipped, 1/4 to 1/3 as long as tube; corolla blue with lilac tinge, 15- 
20 mm long, 2-3 times the length of calyx; tube exserted, dilated above, twisted, the Umb 
inverted; upper (here lower) Up 2-lobed, lower (here upper) lip 3-lobed, the middle lobe 
larger, broader than long, transversely notched, finely toothed, the lateral lobes smaller, 

281 broadly oval; all 4 stamens long-exserted, rarely only 2 or none; anther-cells slightly 
diverging; pistil divided into two thin lobes; nutlets oblong-oval, slightly constricted, 
smooth, brown. May-August. 

Mainly in Central Asian juniper belt. Slopes in large-herb-and-grass steppes, stony 
juniper-covered slopes, alt. 2000-3000 m. — Centr. Asia: Pam.-Al. (Samarkand moun- 
tains, basin of Zeravshan, Zeravshan Range, Kugitang, Kulyab, Shugnan). Endemic. 
Described from Ala-Kisrak east of Kulyab. Type in Leningrad. 

7. L. subnivalis Lipsky in Tr. Bot. sada, XXIII (1904) 209; Levin in Tr. Bot. inst. 
AN SSSR, ser. 1, V, 284. - L. subnivalis jS. tomentosa Lipsky, op. cit. 212. — L. 
tomentosus Rgl. in Izv. Obshch. lyub. est., antr. i etn. XXXIV, 2 (1882) 63, non Forst. 
(1776). — Agastache tomentosa 0. Ktze. Rev. gen. (1891) 511. — Nepeta newes- 
skyi Lipsky, op. cit. 226. 

Perennial, gray-tomentose, sometimes nearly white, rarely greenish (a. virescens (Lip- 
sky) 0. at B. Fedtsch.), glandular; stems numerous, erect, rarely ascending, 10-50 cm 
long; branches from near base and often reaching length of stem; leaves ovate, subcor- 
date or rounded at base, obtuse, generally plicate-rugose, with prominent veins and crisp- 
crenate margins, 5-15 mm long, 3-12 mm wide, short-petioled; upper leaves sessile, often 
all leaves subsessile; petioles 0.5-7 mm long; floral leaves very small, linear-lanceolate; 
inflorescence leafless; cymes in axils of upper leaves, forming loose short-peduncled or 
subsessile inflorescences; middle peduncles 2-8 mm long; calyx tubular, slightly dilated 
above, more or less curved, 2-lipped or nearly so, 6-9 mm long, with hairy ring inside; 
teeth triangular-lanceolate, 1/4 to 1/3 as long as tube, the upper 3 slightly broader; corolla 
lilac, 10-15 mm long, about twice the length of calyx, the tube exserted, slightly dilated 
above, twisted, the Umb inverted; upper (here lower) Up 2-lobed; lower (here upper) lip 
3-lobed, the middle lobe larger, broader than long, transversely notched, finely toothed, 
the lateral lobes smaller, broadly oval, upper (here lower) stamens and pistil long-exserted, 
the lower (here upper) not or slightly exserted; anther-cells slightly diverging; pistil divid- 
ed above into two thin lobes; nutlets oblong-ovaloid, sUghtly compressed, smooth, brown. 
June-September. 

Alpine belt, stony slopes, taluses, pebbles, juniper shale, high-mountain steppes, near 

282 glaciers, 2100-2400 m. — Centr. Asia: Pam.-Al. (Turkestan Range, basin of Zeravshan, 
Zeravshan Range, Gissar Range, Peter the First Range, Alai Range, Alai valley, Trans-Alai 
Range), T. Sh. (Mogol-Tau). Described from Gissar Range. Type in Leningrad. 

8. L. elegans (Lipsky) Levin in Tr. Bot. inst. AN SSSR, ser. 1,V(1940) 288. -Nepe- 
ta elegans Lipsky in Tr. Bot. sada, XXIII (1904) 224 and XXVI (1909) 502. 

Perennial, puberulent, grayish, glandular: stems numerous, 30-70 cm high, erect, 
branching; branches nearly from base, often as long as stems; leaves ovate-triangular, 
oblong-ovate or oval, rounded or truncate or tapering at base, coarsely dentate to nearly 
incised, usually plicate-rugose, very prominently veined, 6-13 mm long, 3-7 nmi wide. 



187 



petioles 2-6 mm long; upper leaves very small, much shorter than cymes, sessile; inflores- 
cence leafless or somewhat leafy; cymes in axils of upper leaves, forming lax peduncled 
inflorescences, middle peduncles 5-12 mm long; bracts small, linear-lanceolate; calyx 
straight, tubular, scarcely dilated above, straight at throat, 6-9 mm long, with hairy ring 
inside at throat, the teeth linear-lanceolate or lanceolate, two-thirds as long to nearly as 
long as tube; corolla blue, 1 5-18 mm long, l-lVi times the length of calyx, the tube long- 
exserted, gradually dilated above, twisted, the limb inverted; upper (here lower) lip 2- 
lobed, lower (here upper) lip 3-lobed, the middle lobe larger, broader than long, trans- 
versely notched, finely toothed, the lateral lobes smaller, broadly oval; stamens not ex- 
serted or the upper (here the lower) slightly so; anther-cells slightly diverging; pistil 
exserted, divided above into two thin lobes; nutlets oblong-oval old, slightly compressed, 
smooth, brown. May-September. 

Stony slopes, rock crevices, taluses. — Centr. Asia: Pam.-Al. (Darvaz, Roshan, Shugnan). 
Gen. distr.: Iran (Afghanistan, Badakhshan). Described from Afghanistan Badakhshan 
(Omar on Pyandzhe). Type in Leningrad. 



Genus 1253.* Schizonepeta** Briq. 

Briq.in Pflanzenfam. IV, 3a (1897) 235. - Saussuria Moench, Meth. pi. (1794) 388, non Saussurea 
DC. 1870 nom. const. - Nepeta sect. Schizonepeta Benth. Lab. gen. et sp. (1834) 468. 

Calyx 15-nerved, obconical, slightly curved, with oblique throat, without hairy ring 
inside; corolla-tube glabrous inside, curved, above rather strongly expanding at the neck, 
283 exserted from calyx; upper lip erect, slightly concave, bifid; lower lip spreading, tripartite, 
the middle lobe unguiculately tapering at base, its upper part strongly broadening, notch- 
ed at apex, the margins crenate, involute, the lateral lobes much smaller; all 4 stamens 
perfect, the upper ascending under the upper lip, the lower projecting forward; anther- 
cells parallel at first, later diverging at nearly a right angle; style split above into two 
subequal lobes; disk with 4 well developed lobes, the lower lobe sometimes much larger 
than the others; nutlets smooth, glabrous, with basal attachment between disk lobes; 
areola rather indistinct. Perennial or annual herbs, with pinnatisect (sometimes partly 
simple) leaves and small flowers forming terminal spikeUke inflorescences. 

Three species are included in the genus: two from Siberia and one from N. China. 

1. Leaves twice pinnatisect or divided into thin lobes; calyx-teeth aristate, corolla 

whitish 2. S. annua (Pall.) B. Schischk. 

+ Leaves once pinnatifid, dissected, rarely divided into broader lobes, sometimes all 

entire; calyx-teeth not aristate; corolla blue-violet ... 1. S. multifida (L.) Briq. 

1 . S. multifida (L.) Briq. in Pflanzenfam. IV, 3a (1897) 235; Kryl. Fl. Zap. Sib. IX, 
2315. - Nepeta multifida L. Sp. pi. (1753) 572 (non L. fil. 1781). - N. lavandulacea 
L. fil. Suppl. (1781) 272; Bge. in Ldb. Fl. Alt. 11,404; Benth. Lab. gen. et sp. 468; in 

* Treatment by A.I. Poyarkova. 
** From the Greek schizein, to split, and the Latin nepeta - name for catmint. 



188 



DC. Prodr. XII, 370; Ldb. Fl. Ross. Ill, 372; Turcz. Fl. baic.-dahur. II, 402; Kom. Fl. 
Man'chzh. Ill, 353; Kom. and Alls. Opred. rast. Dal'nevost. kraya, II, 900. - Saussuria 
pinnatifida Moench, Meth. (1794) 388. — Nepeta lobata Rudolph ex Steud. Nomencl. 
(1821) 552. - Lophanthus multifidus Benth. in Bot. Reg. XV, ad calc. No. 1282. - 
Ic: Rchb. Ic. bot. tab. DXXX, fig. 726; Tr. SOPS, ser. sibirskaya, 3 (1932) Fig. 19. - 
Exs.: GRF, No. 1234a, 1234b. 

Perennial; root woody, longitudinally splitting, passing into woody rhizome, this 
branching above ; stems few, 1 5-60 cm high, 1 .5-3 mm thick, erect or ascending, simple 
or with 1 or 2 pairs of mostly short fertile branches in upper part, covered with rather 
long spreading hairs, especially in lower part and at nodes, and short tightly appressed 
hairs; leaves 1 .5-6 cm long, 1-4 cm wide, with sparse appressed fine hairs above and long 

284 coarser hairs beneath usually confined to veins; lower leaves on petioles about as long as 
blade, orbicular-ovate to oblong-ovate, coarsely and obtusely toothed, rarely pirmatilobate; 
middle leaves sometimes similar to the lower or deeply pinnatifid or pinnatisect pinnatipar- 
tite, with 3-5 (7) entire (the lower sometimes denticulose) obtuse or rarely acute segments 
or lobes; upper leaves short-petioled, usually more deeply cut than the middle, with acute 
or acuminate lobes; sometimes all leaves entire or all pinnatisect; verticillasters 8-20- 
flowered, crowded in cylindrical, spicate, usually dense, rarely lax inflorescences 3-13 cm 
long, 0.7-1.5 cm thick, the lowermost 1 or 2 (3) verticillasters sometimes distant; lower 
floral leaves green, pinnatipartite, the next entire, the others (and often all) broadly ovate, 
strongly attenuate or acuminate, terminating in a long point, mostly bright blue, entire, 
with short appressed hairs on the outside, the margin long-ciliate ; bracts 1.5-2.5 mm long, 
ovate or lanceolate -ovate; pedicels 0.5-1 mm long; calyx 4.5-5 mm long, glaucescent, 
puberulent and beset with minute short-stipitate whitish glands and much larger orbicular 
sessile yellow glands, at margin of throat with 5 small lumpy thickenings between the 
teeth; teeth triangular, acute or acuminate; corolla bluish-violet, 8-10 mm long, twice 

the length of calyx, puberulent outside, with longer hairs on lobes of lips; upper Up ca. 
2.5 mm long, cut to 1/3 or deeper into ovate acuminate lobes; lower lip half as long again 
as the upper, with lateral lobes of similar shape but half as large again, the middle lobe 
obovate in upper part, obtusely crenulate, abruptly attenuate to rather long claw; stamens 
exceeding upper Up; nutlets brown, oblong, rounded at apex, pointed at base, 1.5-1.7 mm 
long, 0.6-0.8 mm wide. Fl. June-August, fr. July-September. (Plate XIV, Figure 3.) 

Steppe zone and adjacent parts of forest zone, meadows, exposed grassy and stony 
slopes of river banks, hills and mountains; in mountains reaching the limit of forest vege- 
tation. — West Siberia: Ob (southern part, rarely), Irt., Alt.; East Siberia: Ang.-Say., 
Dau., Lena-Kol. (southern part); Far East: Ze.-Bu., Uda, Uss. Gen. distr.: Mong., Ch. 
(Manchuria). Described from Dauria. Type in London. 

Note. Very often specimens of S. mul+ifida occur with female flowers (female dioe- 
cism); such flowers are distinguished by the corolla-tube being less dilated above and the 
Umb narrower, the middle lobe differently shaped (narrower), staminodes with very small 
rudiments of anthers, and short filaments included in the tube. 

285 Economic importance. The essential oil of N. multifida is translucent, pale yellow and 
has a pleasant, delicate peppermint odor; it is suitable for perfuming soap (Prokhorova 
andLebedev, 1932). 



189 



2. S. annua (Pall.) B. Schischk. in Sched. Herb. Fl. Ross. X, f. LXIV (1936) 72; Kryl. 
Fl. Zap. Sib. IX, 2314. - Nepeta annua Pall, in Acta Acad. Petr. 1779, II (1783) 263. - 
N. multifida L. fil. Suppl. (1781) 273, non L. (1753). - N. botryoides Sol. in Ait. 
Hort. Kew. ed. I, 2 (1789) 287; Benth. Lab. gen. et sp. 468; in DC. Prodr. XII, 370; Ldb. 
Fl. Ross. 111,372. -N.bipinnatifida Cav. Ic. rar. I (1791) 36. - Schizonepeta botry- 
oides Briq. in Pflanzenfam. IV, 3a (1897) 235; O. and B. Fedch. Perech. rast. Turk. V, 
139. -Ic: Pall. I.e. tab. 12; Cav. 1. c. tab. 49; Tr. SOPS, ser. sibirskaya, 3 (1932) Fig. 
18. -Exs.: GRF,No.3150. 

Annual; stems solitary or few, 9-40 cm long, to 4.5 mm thick, erect or ascending, 
usually branching from base, covered with simple retro rse hairs (longer and spreading in 
lower part) and minute whitish sessile capitate glands; axillary branches leafy, terminating 
in inflorescence, usually branching in turn, the branches sometimes fertile; leaves to 4 
(4.5) cm long, 3 cm wide, broad-ovate or oblong-ovate, deeply bipinnatisect or divided into 
Hnear, oblong or some into oblong-ovate, entire lobes, obtuse or rounded at apex, covered 
on both sides (more densely beneath) with fine appressed hairs sparsely interspersed with 
small capitate, white glands and larger orbicular flat yellow resinous glands; petioles in 
lower leaves as long as or slightly longer than blade, in upper leaves short; verticillasters 
6-10-flowered, crowded at summit of stem and axils of branches into narrow, spicate, 
dense inflorescence, sometimes loose in lower part, 2-10 (12)cm long, 7-15 mm wide 
(without corollas); lower bracts pinnatipartite, the others entire, linear-lanceolate, acumi- 
nate, approximately as long as calyces; bracts linear-subulate, short; pedicels 1-3 mm 
long, all covered, hke calyx, with simple spreading articulate hairs interspersed with stipi- 
tate or sessile capitate whitish glands and yellow sessile resinous glands; calyx 5-6 mm 
long, with distinctly obhque throat; teeth ovate, abruptly attenuate to aristate point, the 
upper 3 ca. half as long as tube, the lower smaller; corolla 7.5-8 mm long, about half as 
long again as calyx, whitish, pubescent and glandular outside, with dense fascicle of long 
hairs on upper Up and middle lobe; upper lip shallowly bifid, with semiorbicular lobes; 
286 lower lip half as long again as the upper; the middle lobe apparently concave, cuneate, 
unguiculate, the very broad limb usually emarginate, remotely and unevenly crenate, the 
lateral lobes half as long as middle lobe, broadly ovate; stamens with blue anthers, the 
upper scarcely exserted; nutlets dark brown, oblong, with rounded apex and pointed base, 
1.5-1.6mmlong, 0.6-0.7mm wide. Fl. July-August; fr. from first half of August. 

Steppes, on stony slopes and rocks; mountains 1000-2000 m in steppe and desert coe- 
noses, stony and gravelly slopes, pebble-beds of mountain streams. — West Siberia: Alt.; 
East Siberia: Ang.-Say. (western part). Gen. distr.: Mong. (mountains and foothills of 
western and central Mongolia). Described from Central Siberia, between Yenisei and Yus 
rivers. Type in Paris. 

Economic importance. The aerial part of the plant (dried) yields 0.56% oil which is 
pale yellow, very mobile, soluble in alcohol and in odor reminiscent of some species of the 
genus Thymus. It contains up to 55% thymol (Pigulevskii and Chistova, 1935). The oil 
appears to be of little interest to the perfume industry, but it can be used as a source of 
thymol. 



190 



Genus 1254.* Nepeta** L. 

L. Sp.pl. (1753) 570, p. p. - Cataria Adans. Fam. 11(1763) 192 . 

Plants usually with bisexual flowers (some of these displaying gynodioecism or gyno- 
monoecism), more rarely gynodioecious or dioecious. Calyx (13-17)-15-nerved, cylindri- 
cal, tubular or obconical, rarely campanulate (often enlarged, ovoid in fruit), curved or 
straight, naked inside, with more or less oblique, rarely 2-lipped or almost straight throat; 
corolla small or medium; tube without hairy ring inside but sometimes with a fringe of 
bristly hairs at throat, narrow in lower part, usually abruptly expanding, rarely gradually 
dilated toward limb; upper Up erect or slightly inclined forward, tripartite, the middle lobe 
much larger than the lateral, concave or nearly flat or with drooping lobules, entire or den- 
tate; throat of limb broad, bordered by base of lateral lobe; stamens parallel, ascending 
under upper Up of coroUa, the upper stamens about equaling the latter, the lower stamens 
287 slightly (rarely much) shorter than the upper, aU perfect; anther-ceUs ellipsoid, usuaUy 
diverging at an angle of 180°, rarely at an angle approaching 90°, their cavities not fusing; 
in female flowers stamens transformed into staminodes, concealed in broadened part of 
coroUa-tube; disk 4-lobed, usuaUy regular, sometimes the lower lobe distinctly larger than 
the others; style 2-lobed, the lobes subequal; nutlets obovoid or elUpsoid, rarely ovoid, 
smooth, granular or tuberculate, each nutlet attached by 2, often fusing, small, basal, gen- 
erally oblique surfaces to 2 adjacent disk lobes. Herbs varying in habit, mostly perennials, 
rarely annuals, very rarely subshrubs, with dentate leaves; upper floral leaves mostly 
strongly reduced, bracteiform; bracts usually not longer than calyx, narrow, very rarely 
as long as flowers or exceeding them. Flowers in half-whorls, usuaUy forming verticiUasters, 
these distant or crowded in spicate or capitate inflorescences, or flowers disposed in re- 
mote pairs of cymes forming racemose or paniculate inflorescences. 

The genus contains some 250 species (82 in the USSR), that are distributed only in the 
Old World, from the Pacific Ocean (Japan, Korea, Maritime Territory) to the Atlantic 
(Spain, Morocco, Canary Islands). A very large number of the species grow in the moun- 
tains of the ancient Mediterranean region, from foothills to the alpine belt. 

Note. The widest variation of type and the greatest abundance of species within the 
genus Nepeta is to be found in two regions: Southwest Asia (especiaUy Iran) and western 
Himalayas including adjacent Hindu Kush. In the USSR most of the species are concen- 
trated in the mountains of Transcaucasia and Central Asia. The origin of the Central Asian 
species is connected with the Himalaya-Hindu Kush development center of Nepeta, since 
most of them belong to sections that are exclusively or predominantly represented in the 
Himalayas and in Afghanistan (Glechomanthe, Capituliferae, Spicatae). There are also 
East Asian links via the section Macronepeta. The Caucasian species, with few exceptions, 
belong to sections of the Southwest Asian center (Schizocalyx, Micronepeta, Oxynepeta, 
Micranthae, and subsection Tuberculatae of Cataria). 

In the century that has passed since the publication of Bentham's monograph (1834 
and in DC. Prodr. 1848), the genus Nepeta as a whole has not been revised. A new 
monographic treatment is obviously necessary in view of the accumulation of vast material 

* Treatment by A.I. Poyarkova. 
** Pliny's name for one of the Labiatae; from Nepeta, Nepet (modern Nepi), an Etrurian city. 



191 



and the fact that the number of species known to science has more than doubled. Cat- 

288 mints of Southwest Asia have been investigated more thorouglily than others thanks to 
the work of Boissier, Bunge, Stapf, Bornmiiller, Bordzilovskii, Rechinger and others. In 
the USSR studies worthy of attention, other than the treatment by Boissier, are those by 
such Russian botanists as Meier (1831), Regel' (1880), Lipskii (1900, 1904, 1909), Gross- 
geim (1928, 1932, 1944, 1949) and Kudryashev (1947). Briquet's classification (Pflanzen- 
fam. IV, 3a, 1897), the second for the genus as a whole, is successively linked up with the 
classification proposed by Bentham and was later considerably revised in parts by Boissier in 
application to the flora of the Near East. Briquet's contribution consists mainly in break- 
ing down Boissier's composite subsections into smaller and more natural entities. In the 
present volume we have followed Briquet's system in that part which, in our view, corre- 
sponds to the phylogenetic relations in the genus. It appears natural to us to interpret the 
following earlier established supergeneric categories as sections: Orthonepeta Benth., Oxy- 
nepeta Benth., Capituhferae Benth., Micranthae Boiss., Longiflorae Boiss. (renamed Schizo- 
calyx Pojark. on formal grounds) and Spicatae Benth. The rest of the system has been 
more or less revised. In reconstructing the system we proceeded from phylogenetic series 
(in the Komarov sense), combining closely related series in subsections and related subsec- 
tions in sections. Much attention was focused on the structure of the flower and in partic- 
ular on details of corolla structure, which had been largely ignored by earlier authors. 

In the course of our work it proved necessary to remove a number of Central Asian species 
from Nepeta and consequently to set up two new genera, Kudrjaschevia Pojark. and 
Drepanocarium Pojark. 

Economic importance. Species of Nepeta yield essential oil, but only a few of them 
have been investigated in this respect. Some of the species studied were found to be of 
little interest to the perfume industry, while the essential oil of others, e. g. N. podosta- 
chys, is highly valued. Deserving of special interest are the Central Asian species among 
which (in sections Capituliferae, Glechomanthe, Spicatae) are some with a very strong, 
persistent and mostly agreeable scent. Some species have long been used as ornamentals 
(N. sibirica L., N. grandiflora M.B.), and their number could undoubtedly be increas- 
ed. Most species are nectariferous. 

1 . Perennials, with woody, more or less thick root and often with well developed 
rhizome 2. 

+ Annuals, with short and slender root 68. 

2. Flowers in large globular terminal head; floral leaves and bracts numerous, violet, 

289 large, exceeding flowers, 1.5-2.3 cm long; leaves cuneate, with unequally incised or 
deeply dentate undulate margin 3. 

+ Floral leaves and bracts small, 2-8 (10) mm long, not exceeding calyx, narrow, 

inconspicuous; leaves different 4. 

3. Floral leaves subtending verticillasters obovate-cuneate or lanceolate-eUiptic, bracts 
linear-lanceolate; flowers large; calyx 9-12 mm long; corolla 18-22 mm long . . 
2. N. lipskyi Kudr. 

+ Lower floral leaves subtending verticillasters in the head cuneate; other floral leaves 
and bracts linear or narrowly linear, finely long-acuminate; flowers smaller; calyx 
7-8 mm long; corolla 15-1 8 mm long 1. N. longibracteata Benth. 



192 



4. Leaves spinose and unevenly sharp-toothed; plant very densely leafy; flowers in 
distant verticillasters; cymes 2-5-flowered; floral leaves resembling cauline leaves; 

corolla with long tube gradually expanding into narrow throat 

3. N. glutinosa Benth. 

+ Teeth of leaves not spinescent; aggregate characters different 5. 

5. Plants usually gynodioecious or dioecious; calyx straight, with nearly straight (rare- 
ly slightly obUque) throat and straight erect subulate teeth 6. 

+ Flowers bisexual* ; calyx with more or less obhque or 2-Upped throat, the teeth not 
subulate, at least the lower more or less curved 14. 

6. Flowers in cymes grouped at ends of stem and axillary branches into small lax pyra- 
midal racemes; flower in cyme bifurcations sessile; corolla completely (rarely only 
to limb) included in calyx; lower Up of corolla obUquely ascending, nearly parallel 
to upper lip (section Oxynepeta) . 7. 

+ Flowers in cymes grouped at ends of stems and axillary branches into mostly long 
narrow racemes; flower in cyme bifurcations pediceled; corolla included in calyx to 
dilated part of tube, rarely higher; lower lip of corolla more or less downturned, 
nearly perpendicular to upper hp (section Orthonepeta) 11. 

7. Corolla azure or blue; stipitate glands usually absent or, rarely, few, confined to 
petiole (series Parviflorae) 8. 

+ Corolla yellow; all plant parts (except corolla) densely beset with short stipitate 

capitate lustrous glands 75. N. erivanensis Grossh. 

8. Calyx 6-7.5 (8) mm long, tube subglobular in fruit; cymes very compact; stem 
usually villous all over or at least in lower part, with long spreading hairs .... 
71. N. parvifloraM.B. 

+ Calyx (8) 9-12(13) mm long, violet-blue, the tube narrowly cylindrical or ovoid- 
cylindrical in fruit; stem with different vesture or glabrate 9. 

9. Stem, leaves and calyx scabrous, with very short hairs antrorsely curved from thick- 
ened base; floral leaves and often several pairs of upper cauhne leaves entire; nutlets 
prominently tuberculate 73. N. schischkinii Pojark. 

+ Stem with short hairs intermixed, at least in lower part, with long ones (simple white 
and pellucid viscous); calyx covered at least in lower part with long fine white 
spreading hairs; all cauline leaves and at least lower floral leaves dentate; nutlets 
indistinctly foveolate-tuberculate 10. 

10. Plant densely leafy, leaves being much longer than internodes; leaves dark green, 
dull; stem covered densely in lower part, sparingly above (often also on lower side 
of leaves), with long fine pellucid viscous hairs; calyx-tube cylindrical-ob conical in 
flower, subovoid in fruit 74. N. kopetdaghensis Pojark. 

+ Plant sparsely leafy; leaves much shorter than internodes, pale green, more or less 
glossy; stems covered with short upcurved hairs, mixed on ribs with longer simple 
white hairs; calyx-tube narrowly cylindrical in flower, cylindrical or rarely ovoid- 
cylindrical in fruit 72. N. ucrainica L. 

1 1 . Flowers yellowish or cream-colored with yellow lower Up; nutlets hairless at apex 
(cycle Sulphureae) 12. 

* Individuals with both female and bisexual flowers or with female flowers alone occur occasionally. 



193 



+ Flowers pale violet (lilac), rarely pinkish or white, with lilac lip; calyx and bracts 

more or less violet-tinged or green; nutlets usually hairy at apex 

67. N. pannonica L. 

12. Plant all gray, with dense, very fine soft implexed hairs; cauline leaves sessile. . . 
69. N. sintenisii Bornm. 

+ Plant green, with minute hairs indiscernible without magnifying glass; cauline leaves 
(except the upper) distinctly petiolate 13. 

13. Flowers small; calyx 5.5-7 mm long; corolla 8-10 mm long, included in calyx near- 
ly to the limb 68. N. sulphurea C. Koch. 

291 + Flowers larger; calyx 8-10 mm long; corolla 12.5-16 mm long, included in calyx 

only to base of dilated part of tube 70. N. komarovii E. Busch. 

14. Stems, inflorescence, lower and often also upper side of leaves covered with thick 
layer of loose, snow-white tomentum; verticillasters numerous, mostly sessile, only 
the lower short-peduncled; corolla tube slightly exserted; nutlets oblong .... 
27. N.olgaeRgl. 

+ Indument rather dense , not tomentose or if so then grayish ; aggregate of other char- 
acters different 15. 

15. Inflorescence of very distant pairs of small (2) 4-6-flowered capitate semiverticels; 
floral leaves and bracts ovate-lanceolate or lanceolate; most semiverticels sessile, 
only the lower on peduncles 0.7-2 cm long 26. N. ladanolens Lipsky. 

+ Flowers in semiverticels or in verticillasters composed of noncapitate cymes or, if 
semiverticels capitate, then many-flowered with peduncles 1 .5-5 (9) cm long, the 
floral leaves and bracts narrowly linear to fihform 16. 

16. Flowers in many-flowered hemispherical capitate remote semiverticels with develop- 
ed peduncles; floral leaves and bracts narrowly linear 17. 

+ Flowers in cymes or in noncapitate semiverticels, these remote or grouped in pairs 
in verticillasters or crowded in spicate terminal inflorescence 19. 

17. Stems leafy nearly to summit, with numerous leafy axillary branches, these branch- 
ing in turn; general inflorescence paniculate . . . .25. N.badachschanica Kudr. 

+ Stems leafy and branching only in lower part; elongate leafless peduncles produced 
above in axils of small bracteiform leaves; general inflorescence a sparse raceme 
18. 

18. Peduncles straight; leaves firm, thick, covered on both sides like petioles and young 
stems, with very short dense tomentum 24. N. pseudofloccosaPojark. 

+ Peduncles arched; leaves thin, pubescent or rather loosely tomentose; petioles and 
young stems with arachnoid indument of long hairs. . . 23. N. vakhanica Pojark. 

19. Corolla-tube half as long again as calyx, gradually expanding toward limb, without 
clearly deUneated neck 20. 

292 + Corolla-tube abruptly dilated into wide clearly deUneated neck 21. 

20. Calyx cyUndrical, tubular (scarcely dilated above), the teeth straight, gradually acu- 
minate, 1/3-2/5 as long as tube 4. N. knorringiana Pojark. 

+ Calyx obconical, dilated above, the teeth 2/3 as long as tube, abruptly terminating 
in a long slender recurved point 5. N. subhastata Rgl. 

21 . Calyx with a fairly broad elHptical slit situated in front, below base of lower teeth, 
and occupying half to nearly 2/3 of the tube; throat strongly obUque, nearly 2- 
lipped (section Schizocalyx) 22. 

194 



+ Calyx without frontal slit 30. 

22. Corolla (20) 22-27 mm long, the tube 2/4 times as long as calyx and S-SVz as long as 
limb (series Longitubae) 23. 

+ Corolla 1 1 .5-20 mm long, the tube 1 Vi-l times as long as calyx and 1 Vi-lVi times as 
long as limb 24. 

23. Flowers in few-flowered cymes; upper cymes sessile, crowded in a loose, (6) 10-12- 
flowered head, the other 2-5 pairs distant, short-peduncled; calyx 7-9 mm long, the 
teeth 4% to 6 times shorter than tube 58. N. longitubaPojark. 

+ Flowers in semiverticels clustered at ends of stem and axillary branches in a dense, 
many-flowered head, sometimes 1 pair of semiverticels remote; calyx 9-12 mm 
long, the teeth 4 to 5H times shorter than tube .... 59. N. sosnovskyi Asker. 

24. Flowers in dense semiverticels, the upper pairs of these sessile, crowded in a terminal 
head, the lower 1 to 4 remote, pedunculate; nutlets smooth (series Brevifoliae) . . 
25. 

+ Flowers in distant peduncled cymes, these forming a racemose or paniculate inflores- 
cence; nutlets very finely tuberculate (cycle Fissae) 26. 

25. Inflorescence of a lax 8-14-flowered head and 1-2 (3) remote pairs of semiverticels; 

lower peduncles 1.2-2.5 cm long; leaves to 17 mm long and as wide 

65. N.brevifoliaC.A.M. 

+ Inflorescence consisting of a dense (20) 30-50-flowered head and 2-4 pairs of re- 
mote semiverticels; lower peduncles 18-30 mm long; leaves to 35 mm long and 
wide 66. N. lamiifolia Willd. 

26. Stems, petioles and lower side of leaves (especially veins) densely covered with long 
white patent articulate hairs; all plant parts (except corolla) densely covered with 

very small subsessile short-stipitate capitate glands 

61. N. trautvetteri Boiss. et Buhse. 

+ Long patent white hairs and capitate glands absent 27. 

27. Stems and leaves velutinous to the touch, with very short simple hairs, these bloom- 
like on stems, corolla 11. 5- 15 mm long, the tube exserted from calyx for 1/3 its 
length 62. N. velutina Pojark. 

+ Indument not velutinous, longer and sparser; corolla 17-19 mm long, tube exserted 
to about the middle 28. 

28. Stems developing long axillary flowering branches nearly all way up from base; 
cymes remote, nearly all (except the upper) diffusely dichotomous; general inflores- 
cence a large, very lax panicle 60. N. fissa C. A. M. 

+ Plant less branching, stems commonly with few axillary branches; upper semiverti- 
cels approximate, the lower distant, only the latter distinctly dichotomous, general 
inflorescence a more or less contracted raceme, rarely a panicle 29. 

29. Stems and leaves scabrous to the touch, with short recurved hairs; leaves with ovate 
or triangular teeth; calyx (7.5) 9-10.5 mm long, the teeth 4^2 to SVz (6) times short- 
er than tube 64. N. iberica Pojark. 

+ Stems and leaves smooth to the touch; leaves crenate; calyx 6.5-8 mm long, the 

teeth 3 1/^ to 4^/i times shorter than tube 63. N. daghestanica Pojark. 

30(21). Middle lobe of lower lip of corolla concave, cup-shaped, with coarsely toothed 
involute margins, without basal bulge; nutlets more or less distinctly tuberculate 
(smooth only in N. cataria L.) (section Cataria) 31. 



195 



+ Middle lobe of lower lip of corolla with basal swelling wedging out toward its notch- 
ed apex, the flat lobules drooping or almost horizontally spreading; nutlets always 
smooth 47. 

31. Nutlets smooth; calyx 5-7 mm long, ovoid in fruit, corolla 7.5-10 mm long, dingy- 
white; flowers in dense compound semiverticels forming dense racemes at ends of 
stem and axillary branches 33. N. cataria L. 

+ Nutlets tuberculate (magnifying glass!); flowers large; aggregate of characters 

different 32. 

32. Flowers in verticillasters crowded at summit of stem in dense spicate inflorescence 

294 (sometimes 1 or 2 verticillasters distant); calyx almost straight, the throat scarcely 
obhque; stems erect, simple; leaves upright (series Betonicaefoliae) 33. 

+ Flowers in verticillasters or cymes, these all remote or only the upper loosely group- 
ed; aggregate of other characters different 35. 

33. Stems and leaves densely gray-pubescent; calyx almost tomentose with implexed 
hairs, 8.5-10 mm long 39. N, betonicifolia C.A.M. 

+ Bright green plants with sparser and hardly visible (magnifying glass!), minute fine 
appressed hairs; calyx covered with straight antrorse hairs (on tube sometimes 
spreading) 34. 

34. Corolla lilac, 18-23 mm long, the tube exserted from calyx for 2-3 mm; semiverti- 
cels with up to 20 flowers; leaves commonly very sparsely hairy, the largest 4-9 cm 
long, 1.5-4 cm wide 40. N. somkhetica Kapell. 

+ Corolla violet-blue, 15-1 7 mm long, tube exserted from calyx for 1.5-2 mm; semi- 
verticels with up to 10 flowers; leaves densely hairy, the largest 2.5-4.8 mm long, 
1.8-2.7 mm wide 41. N. grossheimii Pojark. 

35. Stems 1-3, simple; leaves upright, the upper parallel to stem; calyx slightly curved, 
the throat scarcely obhque, the teeth subequal; inflorescence a raceme of remote 
pairs of semiverticels (series Strictifoliae) 36. 

+ Stems many, strongly branching; leaves more or less horizontally spreading; calyx 
distinctly to strongly curved, the throat conspicuously obhque or nearly 2-lipped, 
the 2 lower teeth much smaller than the upper 38. 

36. Leaves lanceolate or lanceolate-ovate, frequently acute or acuminate, bright green, 

with appressed hairs, sharply dentate; calyx 7-9 mm long 

42. N. strictifolia Pojark. 

+ Leaves ovate, mostly obtuse, crenate; calyx 6-7.5 mm long 37. 

37. Stems and leaves covered with appressed minute hairs (magnifying glass!); leaves 
ovate or oblong-ovate; calyx covered with straight antrorse or, in lower part, spread- 
ing hairs 43. N. alaghezi Pojark. 

+ Stems or leaves with spreading crisp hairs; leaves broadly ovate-cordate; calyx 

densely villous or tomentose-villous, with crisp implexed hairs 

44. N.buhsei Pojark. 

38. Uppermost calyx-tooth longer than the rest, vertically projecting; calyx-tube strong- 
ly curved, the throat nearly 2-hpped; nutlets minutely tuberculate (strong magnifying 
glass!), the pellucid tubercles scarcely raised above surface; inflorescence a more or 

295 less pyramidal raceme or panicle, in upper part sometimes subspicate (cycle Hetero- 
dontae) 39. 



196 



+ Uppermost tooth of calyx not markedly different from others, the tube not strongly 
curved, the throat moderately oblique; nutlets finely and unevenly foveolate-tuber- 
culate, the tubercles not paler than the rest of the surface; inflorescence a narrow 
raceme 42. 

39. Corolla included in calyx up to the limb 40. 

+ Corolla included in calyx at most to the dilated part of tube 41. 

40. Calyx long-villous or tomentose-villous; inflorescence a dense panicle, subspicate in 
upper part; leaves green, with rather long coarse hairs, ovate to lanceolate . . . . 
34. N. cyanea Stev. 

+ Calyx densely covered all over with short appressed or partly subappressed hairs; 
inflorescence a rather lax raceme or panicle; leaves grayish or yeUowdsh with pro- 
fuse short hairs, oblong or narrowly lanceolate 

37. N. biebersteiniana (Trautv.) Pojark. 

41. Leaves yellowish-green, narrowly lanceolate or oblong; calyx puberulent; axillary 

branches long, virgate, the lower longest, about equaling stem 

35. N.kubanica Pojark. 

+ Leaves grayish, usually oblong-ovate; calyx villous; axillary branches subequal, 

much shorter than stem 36. N. czegemensis Pojark. 

42. Inflorescence a 2-sided loose raceme, consisting of remote many-flowered semiver- 
ticels; large plants, to 150cm high, strongly branching . . .38. N. grandiflora M.B. 

+ Inflorescence a more or less 1 -sided raceme, consisting of remote or (near summit 
of stem) more or less crowded verticillasters; plants to 30-50 (80) cm high (cycle 
Mussinianae) 43. 

43. Calyx with long white spreading hairs; stems covered with long simple spreading 
hairs; lower side of leaves similar but hairs subappressed; leaves to 3-4 cm long, 
rounded at apex; corolla pale, blue 49. N. noraschenica Grossh. 

+ Calyx finely pubescent or arachnoid-tomentose; stems and leaves covered with 

short fine appressed, rarely subappressed hairs; corolla darker, violet-blue . . . 44. 

44. Stems erect, 30-80 cm long, to 2.5-3 mm thick, grayish with very fine appressed 
hairs; inflorescence to 30 cm, lax, of 8-11 pairs of remote, 2-6-flowered semiver- 

ticels; leaves to 3.3 cm long, 2-3 cm wide, triangular-ovate 

47. N. hajastana Grossh. 

+ Stems ascending, slender, 0.7-1.5 (2) mm thick; inflorescence shorter; leaves . . 
usually smaller 45. 

45. Hairs on lower side of leaves grayish-velutinous, very fine (strong magnifying glass!), 
flexuous, implexed; flowers small; calyx (5) 6-7.5 mm long; corolla (9) 10-13 mm 
long 45. N. mussiniiSpreng. 

+ Indument of leaves consisting of coarser straight nonimplexed hairs, flowers larger 
46. 

46. Calyx 8-1 1.5 mm long; corolla 16-20 (22) mm long; all or at least the 3-4 upper 
verticillasters closely crowded at summit 48. N. transcaucasica Grossh. 

+ Flowers smaller; calyx 6.5-9 mm long; corolla 1 1-14 mm long; all verticillasters 

distant, even the upper not confluent ... 46. N. reichenbachiana Fisch. et Mey. 

47(30). Flowers in dense spicate terminal inflorescence, this sometimes interrupted in 
lower part, i.e. with 1-3 (5) distant verticillasters; calyx obconical (section Spi- 
catae) 52. 



197 



• 
+ Flowers in distant simple or compound cymes; calyx tubular (section Macronepeta) 
48. 

48. Flowers in compound loose many-flowered cymes; peduncles, except the upper, 
much longer than cymes; corolla 14-18 (20) mm long . . .30. N. formosa Kudr. 

+ Flowers in mostly simple dense few-flowered cymes or semiverticels; peduncles 
shorter than axillary inflorescences and even shorter than flowers; corolla (17) 
20-35 mm long 49. 

49. Leaves narrow, oblong or lanceolate, only the lowermost sometimes oblong-ovate 
(series Sibiricae) 50. 

+ Leaves broad, broadly ovate or cordate (series Gontscharovianae) 51. 

50. Flowers very large; calyx 9.5-14 (15) mm long; corolla 2.5-3.5 cm long; petioles 
of lower leaves short, 1/7 as long as blade 29. N. sibirica L. 

+ Flowers smaller; calyx 8.5-9.5 mm long; corolla 1.7-2 cm long; petioles of lower 
leaves longer, 1/5-1/3 as long as blade 28. N. manchuriensis S. Moore. 

51. Stems simple or in upper part with 1-3 pairs of short slender branches; leaves up- 
right above, nearly parallel to stem, obtuse or rounded at apex; petioles 1/9-1/6 as 
long as blade; corolla 22-27 mm long, the narrow part of its tube exserted to the 
middle 31. N. gontscharovii Kudr. 

297 + Stems strongly branching, the branches long, stout, branching in turn; leaves diver- 
gent, acute, petioles 1/5-2/7 as long as blade; corolla 19-24 mm long, the narrow 
part of its tube exserted from to 1/3 its length . . . 32. N. consanguinea Pojark. 

52. Calyx with straight throat, the linear-subulate teeth about as long as tube; bracts 
nearly as long as calyx; flowers in narrowly cylindrical spicate inflorescence . . . 
6. N. podostachys Benth. 

+ Throat of calyx oblique, the teeth lanceolate or narrowly triangular, 1/3-1/2 as long 
as tube; bracts shorter than calyx 53. 

53. All verticillasters usually in globular or ovoid-globular terminal inflorescence, some- 
times 1 (2) distant; cauline leaves more or less orbicular, small, the largest to 1.5 
(2) cm long and as wide (series Supinae) 54. 

+ Upper verticillasters in cyUndrical or ovoid inflorescence, the others (1) 2-5 distant; 
cauline leaves ovate to lanceolate-linear 58. 

54. Hairs (best seen on inflorescence) strongly branched, the branches consisting of 
several cells (segments), often with secondary ramifications; narrow part of corolla- 
tube not exserted; plant gray-tomentose 22. N. pamirensis F ranch. 

+ Hairs unbranched or sporadically with 1-2 rudimentary (unicellular) branches; nar- 
row part of corolla tube exserted to 1/3-1/2 its length 55. 

55. Plant grayish or wliitish, with dense tomentum on all parts; axillary branches usual- 
ly short, sterile 20. N. kokanica Rgl. 

+ Plant greenish, both leaves and inflorescence less densely hairy, not tomentose; axil- 
lary branches usually long, fertile 56. 

56. Narrow part of corolla-tube exserted from cal)^^ to 1/4-1/3 its length (2-3 mm); 
stems, petioles and leaves covered only with simple white thickish (not arachnoid) 
hairs; capitate sessile glands present only on calyx, bracts and sometimes upper part 

of stem (under inflorescence); leaves with large coarse acute teeth 

21. N. pseudokokanica Pojark. 



198 



+ Narrow part of corolla-tube twice as long as calyx; stems, petioles and leaves cover- 
ed with very thin subarachnoid pellucid simple hairs and copious small whitish 
sessile capitate glands; leaves with obtuse teeth 57. 

57. Flowers large; calyx 9-11 mm long; corolla 20-27 mm long; inflorescence many- 
flowered; leaves mostly cuneate at base; plants usually strongly branching . . . 
19. N.buschiiSosn. et Mand. 

+ Flowers smaller; calyx 7-8.5 mm long; corolla 14-20mm long; inflorescence few- 
flowered; leaves mostly cordate or rounded at base; plants usually few-branched 
18. N. supina Stev. 

58. Cauline leaves small, 1-2.5 (2.7) cm long; middle lobe of lower lip of corolla with 
entire or notched margin except for a fairly deep slit at apex (series Densiflorae) . 
62. 

+ Cauline leaves large, 3 .5-7 (9) cm long; middle lobe of lower lip of corolla large- 
toothed (series Bucharicae) 59. 

59. Cauline leaves narrow, ovate-lanceolate or lanceolate; corolla 10-13 mm long; nut- 
lets ca. 1.5mm long 16. N. alatavica Lipsky. 

+ Cauline leaves broad, ovate, oblong-ovate or orbicular; corolla and nutlets larger . 
60. 

60. Stems, petioles, lower side of leaves and inflorescence covered with dense, whitish, 
very short, tomentose-velutinous or velutinous indument; leaves faintly appressed- 
crenulate to subentire 15. N. maussarifii Lipsky. 

+ Indument not so short, not velutinous, more or less spreading, crisp, interspersed 

with long thin hairs, or with predominance of the latter 61. 

61. Leaves orbicular-ovate; stems and lower side of leaves covered with long thin sub- 
arachnoid hairs ; inflorescence with 1 distant verticillaster 

17. N. schugnanica Lipsky. 

+ Leaves oblong-ovate or ovate; stems and lower side of leaves covered with short 
crisp hairs interspersed with occasional long ones; inflorescence with 1-4 distant 
whorls 14. N. bucharica Lipsky. 

62. Axillary branches elongate, leafy, all or most terminating in inflorescence . . . 63. 
+ Axillary branches short, slender, always sterile, often rudimentary 66. 

63. Flowers small; calyx 4.5-5.3 mm long; corolla 8.5-9.5 mm long; nutlets oblong, 
1 .5-1 .6 mm long; inflorescence a narrow spike without distant whorls; axillary 
branches numerous, about equaling the stem 12. N. tytthantha Pojark. 

+ Flowers larger; calyx 7-8 mm long; corolla 16-18 mm long; nutlets large, 1. 6- 

2.5 mm long, broad 64. 

64. Cauline leaves lanceolate or oblong-ovate, with 3-4 remote teeth, green, with very 
short pubescence (magnifying glass!); calyx 9-1 2 mm long, covered with spreading 
not implexed hairs; inflorescence usually with 1 distant whorl; plants to 25 cm 
high 8. N. transiliensis Pojark. 

+ Cauline leaves ovate, with numerous acute teeth, grayish on both sides with dense 
pubescence; plants higher 65. 

65. All parts of plant, especially stems and calyx, with simple hairs as well as copious 
short capitate glandular hairs; inflorescence consisting of terminal spike and 2-4 
remote verticillasters; leaves cordate or truncate at base, mostly ovate-triangular . 
13. N. odorif era Lipsky. 



199 



+ Hairs predominantly simple, interspersed with few capitate glandular hairs; inflores- 
cence usually consisting only of an ovoid-spherical or spherical-ovoid terminal spike, 
1-2 distant verticillasters infrequently occurring on main stem; leaves with cuneate 
base, mostly rhombic-ovate 9. N. kokamirica Rgl. 

66. Cauline leaves lanceolate (occasionally ovate-lanceolate) with 2-4 remote teeth, 

some entire; calyx 8-10 mm long; corolla 15-16 mm long 

7. N. densiflora Kar. et Kir. 

+ Cauline leaves ovate or elliptic-ovate; flowers smaller 67. 

67. Inflorescence usually with 3-4 (1-5) remote verticillasters; stems and leaves covered 
with very short simple 1-3-cellular hairs; calyx with short not implexed hairs; cau- 
Hne leaves with few remote mostly recurved teeth . . . . 1 1 . N. pulchella Pojark. 

+ Inflorescence usually with 1 (2) remote verticillaster; stems and leaves with longer 
many-jointed simple hairs; calyx with long thin implexed hairs; teeth at leaf margin 
often not recurved 10. N. mariae Rgl. 

68(1). Flowers in remote, sessile, few-flowered verticillasters; bracts (except the upper- 
most) to 2^/4-3 times as long as semiverticels, lanceolate-linear; upper hp of corolla 
curved at apex, with a small notch; middle lobe of under lip with flat drooping 
lobules 82. N. satureioides Boiss. 

+ Inflorescence and bracts different, upper lip of corolla not curved at apex, cleft for 
300 1/3-3/4 its length; middle lobe ot lower lip concave, the lobules with upturned 

margins 69. 

69. Calyx straight, with nearly straight throat, the teeth upright, straight, equal or near- 
ly so; middle lobe of lower lip of corolla directed upward, with basal swelling; nut- 
lets smooth, lustrous; floral leaves not shorter than calyx of flowers of the cymes or 
semiverticels which they subtend, usually much longer, mostly broad, petiolate 
(section Micronepeta) 70. 

+ Calyx curved or straight, with oblique or 24ipped throat; lower 2 teeth much short- 
er and narrower than the upper, all teeth except the uppermost curved forward; 
middle lobe of lower Up of corolla downturned, smooth, without swelUng; nutlets 
tuberculate or granular; floral leaves and bracts small, narrow, shorter than calyx, 
not petiolate (section Micranthae) 75. 

70. Cauline and floral leaves oblate, reniform or semiorbicular (occasionally ovate- 
rhombic); bracts subtending semiverticels foliaceous, spatulate or lanceolate-spatu- 
late, tapering to long petiole 78. N. spathuiifera Benth. 

+ Cauline and floral leaves ovate or lanceolate; leaves subtending semiverticels or 

cymes of a different shape, tapering to short petiole 71. 

71. Inflorescence consisting of terminal head and several pairs of distant cymes; leaves 
subtending cymes and bracts (often also floral leaves) rigid, with thick nerves, 
spinescent; calyx-teeth separated by narrow sinus; lobes of upper lip of corolla 
short, semiorbicular (series Pungentes) 72. 

+ Inflorescence consisting of terminal head and sometimes 1 (rarely 2) verticillasters; 
leaves subtending semiverticels not rigid, not spinescent, ovate or lanceolate-linear; 
calyx-teeth separated by rounded sinus; lobes of upper lip oblong-ovate (series 
Bracteatae) 74. 



200 



72. Terminal head formed by 1-2 pairs of cymes, lax, 5-10 (13)-flowered; all cymas 
below it with peduncles (1) 2-5.5 cm long 81. N. microcephala Pojark. 

+ Terminal head formed by 3-4 pairs of cymes, compact, (12) 16-30-flowered; 2 or 
3 (rarely 1) pairs of remote cymes sessile or subsessile, the others with peduncles 
1-2 cm or 2-6 cm long 73. 

73. Calyx, bracts, lower side of leaves and upper part of stems covered with simple hairs 
and smaller short-stipitate or sessile glands; floral leaves subtending the terminal 
head not exceeding its width (usually about equaling bracts), rigid, curved, always 
spinescent 79. N. pungens(Bge.) Benth. 

+ Calyx, bracts, lower side of leaves and stem covered only with simple soft hairs; 

floral leaves subtending the terminal head exceeding its width, usually leaflike, flat, 
not curved and often not spinescent 80. N. fedtschenkoi Pojark. 

74. Cauline and floral leaves ovate, toothed; leaves subtending semiverticels large (to 
1 cm long), ovate, entire, forming, together with floral leaves, a kind of involucre 
around the head 76. N.bracteata Benth. 

+ Cauline and floral leaves lanceolate-linear or linear, few-toothed or entire; leaves 
subtending semiverticels to 7 cm long, narrow, ovate-oblong or narrowly elliptic, 
entire, not involucral 77. N. daenensis Boiss. 

75. Flowers medium-sized; calyx 6.5-9 mm long; corolla 11-15 mm long; nutlets 
coarsely tuberculate (series Amoenae) 76. 

+ Flowers small; calyx 3.5-8.5 mm long; corolla 3.8-9.5 mm long 77. 

76. Upper cymes loosely approximate (not forming a compact spicate inflorescence), 
the others sparse, all usually loosely dichotomous, with peduncles 3-9 cm long; 

leaves subtending cymes and bracts linear-spatulate or linear-oblong 

50. N. amoena Staf. 

+ Upper cymes crowded in a dense spicate inflorescence, the others compact, only 
the lowermost sometimes dichotomous, with peduncles 1-4 cm long; leaves sub- 
tending cymes and bracts narrowly Unear 51. N. troitzkii Sosn. 

77. Flowers in remote pedunculate cymes; calyx curved, with 2-lipped throat (series 
Micranthae) 78. 

+ Inflorescence consisting of a terminal many-flowered spicate terminal head and some- 
times 1 (2) pairs of distant sessile semiverticels; calyx straight, with oblique throat 
(series Ispahanicae) 81. 

78. Corolla included in calyx up to Hmb; uppermost tooth of calyx mostly half as long 
and sometimes as long as upper lip of corolla; flowers very small; calyx 3.6-5 
(6.5) mm long; corolla 3-8 mm long 79. 

+ Corolla more exserted, the Umb and the dilated part of tube not included in calyx; 
flowers larger; calyx (4) 5.5-8.5 mm long; corolla 7.5-9.5 mm long 80. 

79. Cymes rather loose, mostly loosely dichotomous, the axes of the first order 8-13 mm 
long; general inflorescence loosely racemiform or paniculate; leaves thin, charta- 
ceous, sparsely covered with simple hairs and sometimes with few glandular hairs 
52. N. micrantha Bge. 

+ Cymes dense , often forming a globular head, sometimes only the lower dichotomous, 
the axes of the first order not exceeding 2-3 mm; aggregate inflorescence a compact 
raceme; leaves firm, densely covered with capitate glandular hairs intermixed with 
simple ones 54. N. pallida C. Koch. 

201 



80. Stems, leaves and axial parts of inflorescence with powdery indument of very small 
capitate glandular hairs; cymes lax, 3-8-flowered, mostly loosely dichotomous 

(axes elongate); leaves thin, chartaceous, the veins not prominent 

55. N. saccharata Bge. 

+ Stems, leaves and axial parts of inflorescence grayish, densely covered with simple 
jointed hairs; cymes compact, 6-12 (l6)-flowered, only the lower sometimes 
dichotomous 53. N. meyeri Benth. 

81. Stems, petioles and part of inflorescence densely beset with simple hairs and short- 
stipitate glands; fruiting calyx submembranous; middle lobe of lower lip of corolla 
entire; bracts straight, upright 56. N. ispahanica Boiss. 

+ Short-stipitate glands absent, indument of simple hairs only; calyx indurated in 
fruit; middle lobe of lower Up of corolla large-toothed; bracts arcuately curved. 
57. N. santoana M. Pop. 



Section 1 . Glechomanthe Pojark. sect. nov. in Addenda XIX, 353. - Calyx with 
oblique or 2-Upped throat; corolla with long tube, gradually and rather weakly dilated 
above; middle lobe of lower lip more or less horizontal, flat without sweUing at base, 
gradually tapering to broad base or with rather long broad claw; lateral lobes large, elon- 
gate, ovate, nearly as long as upper lip and as wide as its lobes; lower stamens generally 
much shorter than the upper, with anthers at throat level or even somewhat concealed in 
tube, rarely slightly shorter than the upper; flowers in remote few-flowered cymes or in 
a terminal capitate inflorescence. Perennial plants. 
303 Type of section — N. longibracteata Benth. 

Note. In the structure of reproductive organs, the species of the section Glechomanthe 
do not differ substantially from those segregated in the genus Glechoma that differ con- 
spicuously in habit. Particularly noteworthy is the similarity in the structure of the corolla, 
that finds expression in a complex of characteristic details which can be observed separate- 
ly, or in combination with other characters, in the flowers of certain Nepeta species. 
Some of the features common to the Glechomanthe section of Nepeta and Glechoma 
are: 1) shape of the corolla-tube; 2) elongate lateral lobes of lower lip; 3) shape and 
small size of its middle lobe. In two subsections, Callistegia and Brachystegia, the struc- 
ture of the androecium is the same as in Glechoma. Among species in the subsection Catan- 
therae the androecium is structurally of the type characteristic for nearly all species of 
Nepeta. By way of this subsection, the section Glechomanthe links up with the most 
closely allied sections Macronepeta and Spicatae. The section Glechomanthe almost obli- 
terates the boundary between the genus Nepeta and the genus Glechoma and compels 
a reexamination of the relationship between them, taking into account all their morpho- 
logical and anatomical* features. 



O.F. Bilimovich (Tr. Voronezhsk. Gos. univ., VII, Bot. otd. 1935) found striking differences in the 
structure of the pericarp of the nutlets of Glechoma hederacea L. and three species of Nepeta 
which she investigated. However, since the latter belongs to closely allied sections (Cataria and 
Orthonepeta), no generahzed conclusions can as yet be drawn from this work. 



202 



Subsection 1. Callistegiae Pojark. subsect. nov. in Addenda XIX, 353. — Flowers in 
large dense terminal heads formed by crowded verticillasters; floral leaves resembling the 
cauline; leaves subtending verticillasters and bracts numerous, longer than flowers, suffus- 
ed with violet; calyx obconical, with very oblique nearly 2-lipped throat; lower stamens 
much shorter than the upper, with anthers at the level of throat; anther-cells divergent at 
nearly a right angle; nutlets smooth. Low alpine plants; leaves cuneate, with undulate, 
deeply crenate margin; stems many, the aerial parts densely beset with squamiform leaves. 

Type of subsection — N. longibracteata Benth. 

This subsection contains two closely related species which constitute the natural phylo- 
genetic series Longibracteatae Pojark. 

1. N. longibracteata Benth. Lab. gen. et sp. (1834) 737; in DC. Prodr. XII, 392; Hook. 
Fl. Brit. Ind. IV, 660; Lipskii in Tr. Bot. sada, XXVI, 572, p.p.(excl. var. latebracteata 
04 Lipsky); 0. and B. Fedch. Perech. Rast. Turk. V, 142. — Glechoma longibracteata 
Ktze. Rev. gen. (1891) 518. - Ic: Jacquem. Voy. Ind. Bot. tab. 137. 

Perennial; root woody, knotty, branching in upper part, passing into short rhizome, 
splitting longitudinally below into coarse fibers; stems 8-12 cm long, 1-1 .5 mm across, 
weak, precumbent with remote lower (underground) internodes, branching, often whitish 
in upper part with long fine jointed hairs, in lower part with sparser short hairs and small 
whitish glands, often reddening; axillary branches developed, often terminating in inflo- 
rescence; leaves at 2-3 lower nodes brownish, squamiform, linear-lanceolate, sessile; other 
cauline leaves 0.8-1 .5 mm long, 0.5-1 .2 mm wide, long-petioled, obovate-cuneate, ovate- 
rhombic or ovate, with cuneate or truncate-cuneate base, coarsely crenate-incised, the 
upper sometimes 3-lobed, rather densely covered on both sides with soft long fine hairs; 
leaves subtending the head similar to cauline, those subtending the lower semiverticels 
cuneate, colored or sometimes green, the others bracteiform; inflorescence usually globu- 
lar, rarely somewhat elongate, 1.5-3 (3.5) cm long; bracts linear or narrowly linear, much 
longer than flowers (to 17-19 mm long, 0.5-0.6 mm wide), violet, densely beset at margin 
with long simple hairs and outside with small glandular ones; pedicels 1-1.5 mm long; 
calyx straight, narrowly obconical, 7-8 mm long, 1.6-2.5 mm wide, with strongly oblique 
throat; teeth narrow, lanceolate-triangular, long-acuminate, the lower slightly narrower 
than the upper and slightly longer than calyx-tube, the upper as long as tube or nearly so, 
all teeth, like nerves of tube, densely covered v^th simple long fine hairs and outside vdth 
very small glands; corolla azure, finely pubescent outside, 15-1 8 mm long; tube 9-1 1 mm 
long, slender, curved, gradually dilated above, exserted for nearly 1/3 its length; upper 
hp concave, cleft to the middle or sUghtly less, the lobes broad, obtuse or subtruncate, 
sometimes crenulate at apex; lower lip much larger, the middle lobe oblate, short-clawed, 
broadly emarginate at apex, with remotely and coarsely crenate margin, whitish in middle 
part, with large blue spots, the lateral lobes obovate; upper stamens and style slightly 
shorter than upper lip. Fl. July-August. 

Alpine mountain belt, on shding stony taluses. - Centr. Asia: Pam.-Al. (E. Pamir). 
Gen. distr.: Ind.-Him. (Kashmir) and Dzu.-Kash. (area adjacent to Pamir). Described 
from N. Himalaya, Kan-Ran-Gkhan-Ti, from the Jacquemont specimen. Type in Paris. 

305 2. N. lipskyi Kudr. in Bot. mat. gerb. Inst. bot. i zool. AN UzSSR, IX (1947) 20. - 



203 



Dracocephalum kokanicum Rgl. in Izv. Obshch. lyub. est. antr. i etn. XXXIV, 2 
(1882)66; 0. Fedch., ibid, CIII, 24 (1902) 121. - N. longebracteata var. latebrac- 
teata Lipsky in Tr. Bot. sada, XXVI (1909) 572. — Glechoma lipskyi Pojark. nom. 
altera. 

Perennial; stems 5-10 cm long; root long, robust, vertical, splitting into fibers, branch- 
ing above, the underground perennial parts of stems slender, often elongate, branching; 
stems numerous, ascending, white-pubescent, often red in lower part, 1-1.5 mm across, 
with short sterile axillary branches; leaves at 2-3 lower nodes brownish, squamiform; 
other cauline leaves green, softly grayish-pubescent on both sides, the lower side with 
denser pubescence and often copiously punctate-glandular, obovate-cuneate or rhombic- 
ovate, rarely ovate, 1-1.5 (2) mm long, 0.7-1.2 mm wide, usually tapering to short petiole, 
the margin largely crenate or nearly incised in upper part; semiverticels sessile, forming a 
compact many-flowered capitate globose inflorescence 2-5 cm long and as broad; leaves 
subtending inflorescence green or violet but not differing from cauline leaves in size or 
shape; leaves subtending semiverticels lanceolate-elliptic or obovate-cuneate, long-tapering 
toward base, acute at apex, with large acute teeth, covered outside, like bracts, with very 
small capitate glands interspersed with few punctate glands and fringed with long white 
jointed hairs; bracts as long as or slightly longer than flowers, hnear-lanceolate, long- 
acuminate, violet; pedicels 1-1 .5 mm long; calyx 9-12 mm long, straight, with strongly 
obUque throat, violet or more or less greenish, finely glandular outside; teeth narrowly 
lanceolate, long-acuminate, at margin long-ciUate, subequal or the two lower slightly 
longer, the three upper (1/2) 2/3 as long as tube, the two lower nearly as long; corolla 
18-22 mm long, glandular outside, the tube long, exserted from calyx for approximately 
3 mm, gradually passing into narrow neck 4-5 mm long, 3-3.5 mm wide; upper lip deeply 
cleft into suborbicular lobes; lower lip with large middle lobe (ca. 4 mm long, 6 mm 
wide) and rounded-tetragonal recurved lateral lobes ca. 2 mm long, 2.5 mm wide; nutlets 
brown, broadly ellipsoid, 2-2.5 mm long, 1 .25 mm wide. Fl. July-August; fr. from end of 
July. (Plate XIX, Figure 1.) 

Taluses in alpine and subalpine belts. — Centr. Asia: Pam.-Al. (Alai, Turkestan and 
Gissar ranges). Endemic. Described from Dzhiptyk pass in Alai Range. Type in 
Leningrad. 



306 Subsection 2. Brachystegiae Pojark. subsect. nov. in Addenda XIX, 353. - Flowers 
in remote 1-5-flowered semiverticels; floral leaves resembling cauline leaves but the up- 
permost much smaller; bracts lanceolate, half as long as calyx; calyx obconical, with 
oblique throat; lower stamens much shorter than the upper; anthers diverging at nearly a 
right angle, at throat level or slightly lower; nutlets rugulose, pointed at apex. Perennial 
plants, with strongly thickened woody rhizome and sessile leaves. One species. 
Type of subsection — N. glutinosa Benth. 

3. N.glutinosaBenth. Lab. gen. etsp. (1834)735; id in DC. Prodr. XII, 377; Hook. 
Fl. Brit. Ind. IV, 660. - N. badamdarica Lipsky in Tr. Bot. sada, XXVI (1909) 574; 
0. and B. Fedch. Perech. rast. Turk. V, 141. — Glechoma glutinosa Pojark. in sched. 

Perennial, forming large tufts, 40-70 (and probably more) cm high, with strong lada- 
num odor; root sturdy, to 2-3 cm across, often twisted or bent, woody, splitting at end 

204 



(307) 




PLATE XIX. 1 - Nepeta lipskyi Kudr., general aspect, flower, nutlet in ventral and dorsal view; 
2 - N. glutinosa Benth., general aspect, flower, nutlet in dorsal view; 3 - N. knorringiana Pojark., 
general aspect, flower, nutlet in ventral view; 4 - N. subhastata Rgl., side view of calyx. 



205 



into fibers, passing into rhizome at the top, densely covered all over with imbricated 
brov^'n squamiform lanceolate or ovate nerved and toothed leaves 0.8-1 .8 cm long; stems 
strong, to 4-5 mm thick, erect or slightly ascending, rarely branching, covered below with 
imbricated leaves squamiform, densely leafy above, with long glandular hairs and numer- 
ous sessile glands; axillary branches slender, partly short, sterile, and partly elongate, flori- 
ferous; all leaves sessile, vdth broad semiamplexicaul base, the lower brownish, others pale 
green, viscous with a usually dense cover of long glandular hairs intermixed with sessile 
glands, the middle largest, 1 .3-3 cm long, 0.8-2.2 cm wide, cordate-ovate, the upper ovate, 
all acute; the veins coarse, impressed above, very prominent beneath, terminating in un- 
equal teeth (large usually alternating with small), all acute, mostly curved; flowers in (2) 
4-5-flowered semiverticels in axils of 4-8 upper pairs of leaves, the lower on peduncles 
3-5 (10) mm long, remote, the upper subsessile, approximate, sometimes almost confluent; 
^ all leaves subtending inflorescence resembling the cauline but the upper smaller, sometimes 
309 shorter than calyx, ovate, lanceolate or obovate-lanceolate, with few small teeth; pedicels 
1-2.5 mm long; bracts and leaves subtending semiverticels narrowly lanceolate to linear, 
nerved, approximately half as long as calyx; calyx 8-12 mm long, 2-3 mm wide, erect, 
obconical, with (13) 15 very prominent thick nerves, like other parts of inflorescence 
(except corolla) densely covered with sessile glands and jointed (2-8-celled) glandular 
hairs; throat oblique; teeth ovate-triangular to lanceolate-triangular, the upper 1/4- 
1/3 as long as tube, the lower slightly longer and narrower, 1 /3-2/5 as long as tube; corol- 
la bluish or azure, glandular, sparsely so outside, most copiously on limb, 18-22 mm long, 
twice the length of calyx, the long slender tube much exserted and gradually expanding 
into Umb, not forming any distinct neck; upper lip erect, 2.5-3 mm long, cleft to slightly 
below middle into obtuse lobes, ca. 2 mm wide; lower Hp half as long again as the upper, 
its middle lobe reniform, ca. 2.5 mm long, 4-5 mm wide, coarsely crenate-incised, the 
lateral lobes similar to lobes of upper Up in size and shape; stamens with anthers diverging 
at nearly a right angle, the upper 2/3 as long as upper lip; lower stamens much shorter, 
barely reaching the base of upper lip, with slightly smaller anthers; style nearly equaling 
the upper lip; individuals occur with only female flowers in which the rudiments of the 
stamens are included in the tube; nutlets ellipsoid, tapering at both ends, pointed at apex, 
greenish-brown to cinnamon-brown, transversely rugulose, 2-3 mm long, 1-1.3 mm wide. 
Fl. July-August; fr. from August. (Plate XIX, Figure 2.) 

Alpine belt, 3500-4200 m in steppe plant communities. — Centr. Asia: Pam.-Al. (W. 
Pamir, south only in Bakhan). Gen. distr.: Iran (northeast Afghanistan), Ind.-Him. 
(Kashmir), Dzu.-Kash. (western part). Described from NW Himalayas. Type in Paris. 

Note. In one of the herbarium specimens, some female flowers were observed in which 
stamens were included in the corolla-tube and had very small sterile anthers. 



Subsection 3. Catantherae Pojark. subsect. nov. in Addenda XIX, 353. - Flowers in 
remote cymes or semiverticels; floral leaves resembling the cauline leaves but the upper 
much smaller; bracts lanceolate-subulate, much shorter than calyx; calyx tubular or tubu- 
lar-obconical, with obUque or 2-lipped throat; lower stamens slightly shorter than the 
upper, with anthers diverging at an angle of 180°, lying under flat upper lip; nutlets finely 
papillose. The subsection comprises two Central Asian species. 

Type of subsection — N. knorringiana Pojark. 

206 



4. N. knorringiana Pojark. sp. nov. in Addenda XIX, 353. 

Perennial; plant with oblique rhizome, ca. 1 cm across, covered with remnants of black- 
ish-brown squamiform leaves; stems 13-25 cm long, 0.7-1 .3 mm thick, suberect, simple, 
densely leafy, with internodes to 3.5-4 cm long, covered with fine spreading white simple 
jointed hairs; leaves firm, glaucescent, densely covered on both sides with flat yellow resin- 
ous glands and fine simple hairs, very sparsely so above, rather densely beneath, especially 
along nerves where intermixed with few capitate sessile whitish glands, triangular-ovate, 
with cordate base or the upper with truncate base, obtuse or rarely acuminate at apex, 
along margin with 4-5 large triangular mostly acute teeth, the middle ones largest, to 1.3- 
2cm long, 1.1-1.6 cm wide; lower floral leaves resembling cauHne leaves, others smaller, ovate 
or rhombic-ovate, acute, toothed; semiverticels 2-5 pairs, 2-5-flowered, 1-2 pairs remote, 
other 2-3 pairs (or rarely all) loosely approximate at summit but not forming a head; pe- 
duncles 3-5 mm, the lower to 8-10 mm long; pedicels 0.5-1.5 mm long; leaves subtending 
semiverticels and bracts linear-subulate, phcate, puberulent; bracts 1/3-2/7 as long as 
calyx; calyx 8-9 mm long, violet, 15-nerved, covered with fine simple spreading 3-5-jointed 
hairs sometimes interspersed with few sessile capitate glands, tubular, with oblique throat; 
teeth lanceolate, gradually acuminate, straight, subequal, 2.25-2.7 (3) mm long, the upper 
2/5, the lower 1/2 times as long as tube; corolla 20-24 mm long, sparingly pubescent out- 
side (with traces of bright pink on lips); tube about twice the length of calyx, slightly 
curved above; upper lip 3-4 mm long, cleft to 1/3 into obtuse lobes 2-2.2 mm wide; 
middle lobe of lower lip cuneate-flabelliform, 3.5-4.5 mm long, 4-5.5 mm wide, gradually 
tapering from middle to wide base, emarginate and obscurely crenulate at apex, the lateral 
lobes ovate, 2.5-3 mm long, 1 .7-2.5 mm wide; upper stamens and style equaling the upper 
corolla-lip, lower stamens slightly shorter; anthers broadly elliptic; nutlets 1.6-1. 8 mm 
long, 0.7 mm wide, narrowly ellipsoid, 3-angled, with a rib on ventral side, pointed at apex, 
brown, densely papillose at apex, sparsely elsewhere. Fl. and fr. last half of June. (Plate 
XIX, Figure 3.) 

Stony slopes in high-mountain belt. — Centr. Asia: Pam.-Al. (eastern part of Alai 
Range). Endemic. Described from Tara River, tributary of Kara-Kul'dzha River. Type 
in Leningrad. 

5. N. subhastata Rgl. in Izv. Obshch. lyub. est. antr. i etn. XXXIV, 2 (1882) 64; O. 
and B. Fedch. Perech. rast. Turk. V, 143. 

Perennial, sometimes almost suffruticose; root thick, to 1 cm across, woody, more or 
less twisted, splitting at the end into fibers, passing at the top into a thick tough short 
rhizome bearing remnants of old stems; stems 8-25 cm long, ascending, slightly curved, 
slender, 0.8-1.5 mm across, with internodes 2.5-5 cm long, not branching, leafy all over, 
scabrous, densely covered (especially in upper part) with very short 1-2 (3)-cellular simple 
spreading hairs intermixed with small sessile glands; leaves pale, glaucescent-green, densely 
beset beneath (at first also above) with very short hairs and sparse yellow punctate aroma- 
tic glands, ovate, acute or obtuse, with cordate or sometimes deeply notched base (in 
upper leaves cuneate), the margin with large obtuse or rarely acute teeth; leaves on middle 
part of stem largest, 1.3-2 cm long, 1-1.8 cm wide; petioles of cauline leaves 1/6-1/3 as 
long as blade, with indument as on stems; lower floral leaves resembling the cauline but 
sessile, others small, 0.7-1 .2 mm long, 0.3-0.4 mm wide, elliptic, finely toothed or entire; 



207 



flowers in dense few-flowered semiverticels, these (except 1, rarely 2) crowded near sum- 
mit to form a loose head 3-4 cm long (without corollas); upper peduncles 2-5 mm, the 
lower to 1 cm long; bracts 1/3-2/5 as long as calyx; bracts and leaves subtending semiver- 
ticels lanceolate to linear, with minute hairs, the margin ciUolate; pedicels 0.5-1.7 mm 
long; calyx 7.5-9 mm long, narrowly obconical, in fruit subcampanulate, 13-nerved, with 
nearly 2-lipped throat, densely covered with short thick spreading hairs more or less inter- 
spersed with sessile capitate white glands; teeth lanceolate, abruptly terminating in a long 
recurved point, the 3 upper 2/5-1/2 as long as, the 2 upper [should be lower] about equal- 
ing, the tube; corolla 18-23 mm long, sparsely hairy outside, the tube slightly curved, ex- 
serted 5-7 mm from calyx, gradually expanding toward limb; upper hp erect, 3-4 mm 
long, cleft to 1/4-1/3 into 2 broad obtuse lobes; lower lip almost twice as long as the 
upper, its middle lobe cuneate-flabelliform, 4.5-5 mm long, 5-5.5 mm wide, gradually 
tapering to short claw, coarsely crenate, rather deeply emarginate, the lateral lobes ovate, 
2.5-3.3 mm long, 2-3 mm wide; upper stamens and style about equaling the upper lip; 
312 lower stamens reaching the middle of upper lip; nutlets eUipsoid, 3-angled, 2.5 mm long, 
0.8 mm wide, pale brown, unevenly and finely plicate, minutely tuberculate. Fl. June; 
fr. toward last half of July. (Plate XIX, Figure 4.) 

Stony slopes in high-mountain belt. — Centr. Asia: Pam.-Al. (eastern part of Turkestan 
Range and western part of Alai Range). Described from northern slope of Turkestan 
Range, Chichikta on Chiburgan River. Type in Leningrad. 

Note. This unique species was first described from the incomplete collections of 
0. Fedchenko (without flowers) from the upper reaches of the Isfara River; it was only 
much later that flowering specimens were collected on Alai Range (Kutban-Kul lake) and 
made it possible to establish its taxonomic position. 



Section 2. Spicatae (Benth.) Pojark. comb. nov. — Sect. Pycnonepeta § Spicatae 
Benth. in DC. Prodr. XII (1848) 371. - Sect. Eunepeta § Spicatae Boiss. Fl. or. IV (1879) 
637; Briq. in Pflanzenfam. IV, 3a, 236. - Flowers in verticillasters forming a dense spicate 
terminal inflorescence, this sometimes interrupted in lower part (i. e. 1-3 (5) lower verticil- 
lasters remote); bracts linear-subulate; calyx obconical, mostly slightly gibbous dorsally, 
with straight or oblique, sometimes 2-lipped throat; corolla medium-sized, the curved 
tube abruptly expanding into large neck; middle lobe of lower lip with sweUing at center, 
tapering from broad base, with horizontal or slightly drooping lobules; nutlets smooth. 
Perennial plants. 

This section is most diversified and most widely represented in the western Himalayas; 
a large number of species occur in Central Asia and Afghanistan, but very few in Iran. 
Two grow in the Caucasus. 



Series 1. Podostachydes Pojark. — The upper or all verticillasters forming an oblong 
cylindrical spicate inflorescence; calyx with straight throat and subequal linear-subulate 
teeth; narrow part of corolla-tube completely included in calyx or barely exserted; middle 
lobe of lower lip of corolla entire; bracts equaling or exceeding calyx; nutlets lustrous. 

In addition to N. podostachys distributed in Central Asia, tliis series contains N. kan- 
dagarica Kudr. (SE Afghanistan) and N. assurgens Hausskn. et Bornm. (SE Iran). 

208 



6. N. podostachys Benth. in DC. Prodr. XII (1848) 372; Boiss. Fl. or. IV, 639; Franch. 
PI. Turkest. 137; Lipsk. in Tr. Bot. sada, XVIII, 95; 0. and B. Fedch. Perech. rast. Turk. 

313 V, 139. -N. maracandia Bge. in Mem. div. sav. Ac. Sc. Petersb. VII (1851) 434; Boiss. 
Fl. or. IV, 640. - N. paulseni Briq. in Bot. Tidsskr. 28 (1908) 235. - Ic: Briq. 1. c. 
fig. 2, in Tr. Sredneaz. n.-i. opytn. st. efiro-masl. rast. 1 (1932) Fig. 39. 

Perennial; root woody, coarsely fibrous, more or less twisted, longitudinally splitting, 
passing above into more or less branching rhizome; stems many, 30-50 (70) cm high, in- 
durescent at base and beset with squamiform leaves, arcuately ascending to suberect, 
slender, subterete, sparsely puberulent, in lower part subglabrous, with very distant upper 
internodes, branching; lower branches short, rudimentary, sterile, the upper leafy, fertile 
and developing short sterile branches; leaves thin, more or less densely covered with basal- 
ly thickened spreading simple hairs, sometimes subglabrous; cauline leaves lanceolate or - 
oblong, small, to 2.5 cm long, 0.8 cm wide, acuminate, gradually tapering to slender peti- 
ole, rarely (mainly the lower) ovate-lanceolate to triangular-ovate, with rounded-cuneate 
or subtruncate base, each margin with 2-4 remote teeth, these acute or obtuse, patent, 
often large, almost lobelike; inflorescence dense, compact, oblong-cylindrical, to 9 cm 
long, 1 .6 cm wide, the lower 1-2 (3) verticillasters sometimes distant; floral leaves lanceo- 
late or oblong-elliptic, mucronate, entire, the upper bracteiform, mostly slightly shorter, 
rarely slightly longer than calyx, linear-subulate, very finely point-tipped but not spines- 
cent; pedicels 0.75-2 mm long; calyx 5-8 mm long, narrowly obconical, 15-nerved, dense- 
ly covered like bracts, with short-stipitate capitate glands and, on the nerves, with simple 
many-jointed hairs, the throat straight, the teeth long-acuminate from lanceolate base, the 
upper slightly longer than the lower, equaling or slightly exceeding the tube; all teeth, as 
well as bracts, cilia te or villous at margin, with long implexed pellucid many-jointed hairs; 
corolla 8.5-1 1 mm long, sparsely puberulent outside; tube barely exceeding calyx-teeth, 
slender, curved, abruptly expanding into a broad neck, 1-2 mm long, 1 .8-2.5 mm wide; 
upper lip 1.7-2 mm long, erect, cleft to middle into oblong-ovate lobes, the middle lobe 
2-2.5 mm long, 2.5-3.5 mm wide, cordate, strongly bulging at center especially near base, 
with entire margins, the lateral lobes semiorbicular-triangular, 0.7-1 mm long, 1 .5-2 mm 
wide; upper stamens as long as upper lip of corolla; style slightly exserted; nutlets black, 
lustrous, ellipsoid, 1.4-1. 6 mm long, 0.8-1 mm wide. Fl. end of June to first half of Sep- 
tember; fr. from July. 

314 Upper zone of wood and scrub belt; juniper groves, rosariums, in alpine belt in mea- 
dows and steppe coenoses, open stony and gravelly slopes. — Centr. Asia: Pam.-Al. Gen. 
distr.: Iran (eastern part — E. Afghanistan) and Ind.-Him. (southwestern part — Kashmir). 
Described from Afghanistan, mountains near Kabul. Type in Kew. 

Note. This polymorphous species has an extensive distribution area, but it has not 
been possible to discern any geographical pattern in its variability. Consequently, neither 
N. maracandia Bge. (type from vicinity of Samarkand) nor N. paulseni Briq. (type 
from Yashil'-Kul' lake in Pamir) has been recognized as an independent species. Most dis- 
tinctly diverging from the type, and possibly deserving of attention, is a form distributed 
in the rosarium belt in the western part of the Darvaz area which is particularly conspicu- 
ous because of its height (more than 1 m) and the large size of all parts, including inflores- 
cence and flowers. 

Economic importance. The essential oil of this plant has a pleasant odor and may be 
used in the perfume industry, but the yield does not exceed 0.04%. 

209 



Series 2. Densiflorae Pojark. - Upper or all verticillasters in cylindrical or ovoid spi- 
cate inflorescence; calyx-throat oblique, the teeth lanceolate, rarely lanceolate-triangular, 
(2/3) 1/2-2/5 as long as tube; corolla with tube included in calyx to base of neck (rarely 
slightly less), the middle lobe of under lip entire or scarcely emarginate; bracts much 
shorter than calyx; nutlets dull; cauline leaves small, to 10-25 (30) mm long. 

This series is nearly endemic to the upper mountain belt of Central Asia; only one 
species grows beyond its range, in southwestern Altai. 

7. N. densiflora Kar. et Kir. in Bull. Soc. nat. Mosc. XIV (1841) 725; Ldb. Fl. Ross. 
Ill, 373; Benth. in DC. Prodr. XII, 372; Kryl. Fl. Zap. Sib. IX, 2313. - N. saposhniko- 
wii Nik. et Plotn. in Bot. mat. gerb. Bot. sada, VI, 1 (1926) 20. 

Perennial, rhizome slender, creeping, branching at end, covered at nodes, like the lower 
part of stems, with dark brown squamiform leaves; stems to 25-35 cm long, ascending at 
base, erect above, 1 .5-2.25 mm thick, with internodes 5-6 cm long, covered more sparsely 
in lower part, with spreading crisp simple hairs interspersed with very small glandular hairs; 
axillary branches slender, always sterile, the lower with 4-7 pairs of leaves and developed 
internodes, the upper mostly rudimentary, short; leaves bright green, sparsely covered on 
both sides with short simple hairs interspersed, more copiously beneath, with minute glan- 
315 dular hairs and punctate yellow oil glands; cauline leaves 1 5-30 mm long, 2-10 mm wide, 
lanceolate or rarely some oblong-ovate, gradually acuminate from cuneate or rounded- 
cuneate base, usually entire in lower part, with 1-4 small distant antrorse teeth mostly 
near apex; lower leaves with petioles 2-4 mm long, others sessile; leaves of sterile lateral 
branches smaller and narrower, oblong-elliptic or lanceolate-linear, often obscurely tooth- 
ed or entire; pedicels 0.5-1 .25 mm long; inflorescence in flower ovoid or oblong-ovoid, 
usually with 1 remote verticillaster, rarely without it, subtended by small lanceolate green 
or more or less colored terminal leaves; bracts bluish-violet, lanceolate-linear, long-acumi- 
nate, 7-1 1 mm long, not exceeding calyx, densely covered with very small glandular hairs 
intermixed with long simple hairs, these forming a dense fringe along the margin; calyx 
bluish-violet, 8-10 mm long, covered outside, especially on nerves, with long hairs and 
densely beset with very small glandular ones, the throat oblique; teeth narrowly triangular, 
long-acuminate, the upper and lower similar in shape, the 2 lower sometimes 0.5-1 mm 
shorter, the upper ca. 1/2-2/3 as long as tube; corolla pubescent outside, blue, 15-16mm 
long, its tube as long as calyx; upper Up 3 mm long, cleft to middle or often for 2/3 into 
2 obtuse lobes, 2-2.5 mm wide; middle lobe of lower lip 3-4 mm long, (6.5) 7-8 mm wide, 
with deep and broad notch at middle, the lateral lobes semiorbicular-triangular, 0.75-1 
(1.5) mm long, 2.3-3 mm wide; upper stamens slightly exceeding upper Up; nutlets dark 
brown, broadly ovoid, 2 mm long, 1-1.25 mm wide. August. 

Alpine belt, in stony places. - West Siberia: Alt. (southwestern part only - Narym 
Range, Kaba region). Gen. distr.: Mong. (Mongolian Altai). Described from Narym 
Range, Dzhaidak River. Type in Leningrad. 

8. N. transiliensis Pojark. in Bot. mat. gerb. Bot. inst. AN SSSR, XV (1953) 286. 
Perennial; root knotty, vertical or ascending, longitudinaUy splitting into coarse fibrous 

sections, branching at the top, passing into a short multicipital rhizome; stems 20-30 cm 
long, 1 .5-3 mm across, ascending, often slightly arched, with internodes to 5 mm long. 



210 



branching from base, crisp-hairy, the hairs short, simple, jointed and interspersed with 

316 smaller sessile glandular hairs; axillary branches (except for 1-2 uppermost pairs) elongate, 
with 6-7 developed internodes, usually terminating in inflorescence, this in turn producing 
short axillary branches; leaves and stems densely covered with short simple glandular hairs 
intermixed with punctate yellow glands; basal leaves squamiform, brown, amplexicaul; 
cauline leaves 1 .5-2.3 cm long, 0.5-0.8 cm wide, oblong-ovate to lanceolate, gradually 
acuminate above, with narrowly cuneate base, the margins toothed nearly from base, the 
teeth few, equidistant, acute, patent; lower leaves short -petioled, the upper sessile; leaves 
of axillary branches smaller and narrower; floral leaves Unear-lanceolate, green, sessile; 
semiverticels sessile, forming a terminal oblong ovate or subcylindrical inflorescence, this 
4-7 cm long on main stem and usually accompanied by 1 more or less remote verticillaster; 
bracts lanceolate, 5-7 mm long, always shorter than calyx and, like the latter, blue-violet, 
covered with short scablike glandular hairs and spreading jointed simple hairs, the margin 
ciliate; pedicels 0.5-1 mm long; calyx 9-1 2 mm long, 3.5-4 mm wide; teeth oblong- 
triangular, acuminate, the upper 2.5-3.5 mm long, (2/7) 1/3-1/2 as long as tube, the lower 
sometimes slightly shorter and narrower than the upper; corolla (violet-blue?) 17-18 mm 
long, pubescent outside, the curved tube as long as calyx or slightly (0.5-0.75 mm) exsert- 
ed; upper Up 3-3.5 mm long, cleft to 2/3 into obtuse broad lobes, 2.3-2.5 mm broad; 
middle lobe of lower lip on stipe 0.6-0.8 mm long, 4 mm long, 7-8 mm wide, the lateral 

' lobes 0.7-1 mm long, 2.7-3 mm wide; upper stamens and style slightly exceeding upper 
lip; nutlets elongate-ovoid, 2.5-3 mm long, 1-1.2 mm wide, brown. Fl. from July; fr. 
September. 

Stony places, rock crevices in alpine belt. - Centr. Asia: T. Sh. (eastern part of Zailiisk 
Ala-Tau). Endemic. Described from Karach Range, upper reaches of Turgen' River. Type 
in Leningrad. 

Note. Very similar to N. densiflora in habit but easily distinguished by various char- 
acters outlined in the key. 

9. N. kokamirica Rgl. in Tr. Bot. sada, VI (1879) 358; 0. and B. Fedch. Perech. rast. 
Turk. V, 141. - Ic: Gartenflora, XXIX, tab. 1030. 

Perennial; root knotty, splitting, passing into branching rhizome, this, Uke lower part 
of stem, covered with brown squamiform leaves; stems numerous, ascending or decum- 
bent in lower part, 1 5-50 cm long, 1 .5-3 mm thick, with internodes to 7 cm long, grayish, 
densely covered with crisp spreading jointed simple hairs interspersed with few very short . 

3 1 7 glandular hairs; axillary branches numerous, often equaUng stem, with long internodes, 
branching, many terminating in inflorescence; leaves densely grayish-tomentose on both 
sides, small; cauHne leaves to 17 mm long, 8 mm wide, ovate or rhombic-ovate, acute, 
acuminate or obtuse, cuneate at base, toothed along whole margin above base, the lower 
with petioles 1/3-1/2 as long as blade, the upper with petioles ca. 2 mm long; leaves on 
axillary branches narrower, oblong-ovate, rarely sublanceolate or suboblong; verticilla- 
sters usually all included in terminal inflorescence, in flower broadly ovoid or suborbicular, 
rarely lower verticillasters on main stem more or less remote; leaves subtending semiverti- 
cels lanceolate or linear-lanceolate; bracts linear, 4.5-6 mm long, much shorter than calyx, 
both bracts and calyx blue-violet and densely covered with long spreading sericeous-arach- 
noid jointed hairs (forming a dense fringe at margins) intermixed with short glandular hairs 



211 



and punctate yellow glands; calyx 7-8 mm long, 2-3 mm wide; teeth oblong-triangular, 
2-2.5 mm long, 1-1 .5 mm wide, acuminate, all equal or the lower slightly narrower, (2/5) 
1/2 as long as tube; corolla light blue, pubescent outside, 16-18 mm long, the tube exsert- 
ed from calyx for 1 .5-2 mm; upper lip (2.5) 3 mm long, cleft to 2/3 into obtuse lobes; 
middle lobe of lower lip 4.5-5 mm long, 7-8 mm wide, tapering to a broad claw 0.4- 
0.6 mm long, the lateral lobes (0.7) 1-1.25 mm long, 2.5-3.5 mm wide, triangular flat ; 
upper stamens not exceeding upper lip; nutlets oblong-ellipsoid, 1 .6-2 mm long, 0.7- 
1.25 mm wide, blackish-brown. Fl. June-July; fr. from middle of July. 

From alpine belt to upper timberline, taluses and stony places. - Centr. Asia: Dzu.- 
Tarb. (Dzungaria Ala-Tau). Gen. distr.: Dzu.-Kash. (Boro-Khoro Range, Kuldzha region). 
Type in Leningrad. 

10. N. mariae Rgl. in Tr. Bot. sada, VI (1879) 359; Lipsk. in Tr. Bot. sada, XVIII, 97; 
O. and B. Fedch. Perech. rast. Turk. V, 140. 

Perennial; root multicipital, knotty, longitudinally spUtting into coarsely fibrous sec- 
tions, in upper part to 8 cm across; stems 1 2-40 cm high, 1 .2-2.5 mm across, covered with 
a mL-1-.. of crisp simple jointed hairs (these usually fairly long on upper part of stem and 
under nodes) and very small glandular hairs; lateral branches all sterile, very short and 
slender, only few with developed internodes; leaves mostly densely covered on both sides 
318 with very short (longer only on veins) simple crisp hairs intermixed with small glandular 
hairs; cauline leaves 10-23 mm long, 8-12 mm wide, the lower with petioles 2.5-4 mm 
long, the upper with petioles 1-2 mm long, broadly ovate, with broadly cuneate or rounded- 
cuneate, sometimes truncate base, acute or obtuse, the upper sometimes acuminate, the 
marginal teeth large, acute or obtuse, closely approximate; leaves of lateral branches 
oblong-elliptic or ovate-lanceolate; inflorescence with 1, rarely 2 remote verticillasters, 
its terminal part 1.5-4 cm long, at flowering globose (smaller inflorescences) to oblong- 
elHpsoid; bracts and calyx bluish-violet, very densely beset with small glandular hairs and 
sparse sessile yellow punctate glands, with long white simple hairs forming a dense fringe 
along the margin and rather densely covering calyx mainly along the nerves; bracts much 
shorter than calyx, linear or lanceolate-linear; calyx 7.5-9 mm long, the teeth triangular- 
lanceolate or oblong-triangular, acuminate, the upper 1/3-1/2 as long as tube; corolla 12- 
15 mm long, pubescent outside, blue, its tube not exceeding calyx; upper lip 2.5-4 mm 
long, cleft to middle into obtuse broad lobes; middle lobe of lower lip 3-3.5 mm long, 
5-6 mm Wide, the lateral lobes 0.8-1 mm long, 2-3 mm wide, sometimes undulate; sta- 
mens not exceeding upper lip; nutlets dark brown, obovoid, 2.2-2.4 mm long, 1 .25- 
1 .4 mm wide. Fl. July -August; fr. from last half of August. 

Alpine and subalpine belts, also upper part of the forest belt, wet grass plots, stony 
placers, river banks. — Centr. Asia: T. Sh. (Fergana, Chatkal', Pskemsk ranges, eastern 
part of Talass Ala-Tau), Pam.-Al. (Turkestan and Zeravshan ranges). Endemic. Described 
from W.Tien Shan. Type in Leningrad. 

1 1. N. pulchella Pojark. in Bot. mat. gerb. Bot. inst. AN SSSR, XV (1953) 282, Fig. 1. 
Perennial; root ascending, woody, longitudinally spUtting at lower end into fibrous 

sections, branching in upper part, multicipital, extending into short rhizome, this densely 
covered with brown scales; stems few (up to 10), 25-35 cm long, erect, robust, 1.5-2.5 mm 



212 



across at base, with mixture of very short profuse simple hairs and very small scabUke glan- 
dular hairs; axillary branches developing nearly at all nodes except for 2-4 of the lower- 
most, but usually rudimentary, very slender, with 2-4 short internodes, always sterile; 

319 lowermost leaves squamiform, brown, long-persistent; other cauline leaves pale green, 
covered (like stems) with very short hairs visible only under magnifying glass and sparse 
punctate yellow glands, to 27 mm long and 12 (18) mm wide, oblong-ovate or oblong- 
elliptic, occasionally ovate, acute or obtuse at apex, cuneate at base, the margin unevenly 
and acutely toothed from base or only in upper part, the petioles 3-4 mm long; upper 
leaves subsessile, often entire; leaves on sterile axillary branches oblong or narrowly lan- 
ceolate, remotely serrate or the upper sometimes entire; floral leaves sessile, the lower 
resembling cauline leaves but narrower, lanceolate, the upper bracteiform; verticillasters 
dense, sessile, the lower 2-4 (5) remote, others forming a dense spicate inflorescence, this 
(1.5) 2-4 cm long, oblong-ovoid or subcylindrical (ovoid in weak specimens); pedicels 2- 
3 (5) mm long; bracts lanceolate or linear-lanceolate, sometimes as long as calyx-tube but 
mostly half as long, partly or rarely completely violet-blue, all densely beset with very 
small glandular scablike hairs; long white simple hairs covering the nerves and forming a 
dense fringe around the margin; calyx violet-blue, 6.5-8 (10) mm long; teeth triangular- 
lanceolate, long-acuminate, the upper approximately 2/5 as long as the tube, the lower 
slightly shorter and narrower; corolla plain pale blue, puberulent outside, 10-13 mm long, 
the tube exserted only about 1 mm, curved, abruptly expanding into neck; upper lip ca. 
2.5 mm long, cleft to 2/3-3/5, the lobes broad, ovate, rounded above, 1.5-1.75 mm long, 
1.75-2 mm wide; middle lobe of lower lip 3-3.5 mm long, 5-6 mm wide, horizontally 
spreading, with a shallow notch at apex and slightly sinuate margin; lateral lobes rounded, 
0.75 mm long, 1 .75-2 mm wide; upper stamens and style as long as upper hp; anthers 
blue; nutlets brown, obovoid, obtusely 3-angled, 2.3-2.5 mm long, 1.3-1. 7 mm wide. Fl. 
second half of June-August; fr. from last half of July. 

Subalpine zone and upper timberline, northern slopes in meadow coenoses. — Centr. 
Asia: T. Sh., western part of Talass Ala-Tau. Endemic. Described from Kshi-Kainda 
River (basin of Aksu River). Type in Leningrad. 

Note. N. pulcheila recalls N. mariae Rgl. and N. kokamirica Rgl. N. mariae Rgl. 
differs from N. pulcheila in having broad, ovate leaves, denser pubescence, dense, large, 
acute teeth along the margin, the inflorescences usually only with 1, rarely 2 remote semi- 
verticels and shorter corolla-tube. N. kokamirica Rgl. is distinguished by well developed, 

320 long axillary branches usually bearing inflorescences, only 1-2 remote semiverticels on the 
main inflorescence, larger (16-18 mm long) corollas and longer and denser hairs covering 
the stems and leaves. 

12. N.tytthanthaPojark. in Bot. mat. gerb. Bot. inst. ANSSSR, XV (1953) 288, 
Fig.-2. 

Perennial, strongly branching; stems 40-60 cm long, ascending, 2.5-3 mm across in 
lower part, branching from base; branches long, many as long as stem, sometimes longer, 
all usually terminating in inflorescence; stems and axillary branches densely covered (at 
least in lower part) with very short white simple spreading hairs and very small glandular 
ones; leaves pale green, glaucescent, densely covered, especially when young, with hardly 
visible (magnifying glass!) hairs similar to those on stems but interspersed with punctate 



213 



yellow glands, on stems oblong-ovate; uppermost leaves lanceolate, 16-20 mm long, (6) 
8-12 mm wide (smaller on axillary branches), rounded or subcordate or some cuneate at 
base, gradually acuminate above, at margin with large acute, more or less appressed teeth, 
all leaves distinctly petiolate, the petioles 1/8-1/3 as long as blade, shorter in upper leaves 
than in lower; lower bracts resembling cauline leaves, the others bracteiform; inflorescence 
dense, spicate, narrow, cylindrical or oblong-ovoid, 2.5-4 cm long, commonly without 
remote verticillasters, rarely one subremote; semiverticels sessile; pedicels 0.5-1.5 mm 
long; bracts 1.5-3. 5 mm long, linear-lanceolate or linear, reaching the middle of calyx-tube 
or the largest the base of teeth, at margin with a fringe of long white 3-5 (6)-jointed simple 
hairs; calyx small, (4.5) 5-5.3 mm long, 1.3-1 .5 mm wide, tubular-obconical, slightly 
curved, nearly 2-hpped, with strongly obhque throat, densely covered outside with very 
small glandular hairs interspersed with punctate glands, and with long jointed simple hairs 
along nerves and at margin of teeth, the teeth oblong-triangular, acuminate, the 3 upper 
1/3-2/5 as long as tube; corolla blue-azure, sparsely pubescent outside, 8.5-9.5 mm long, 
the tube 4.5-5 mm long, not exserted, abruptly expanding at front into neck; upper hp 
2-2.5 mm long, deeply cleft into 2 broad obtuse lobes; middle lobe of lower lip sessile, 
ca. 2.5 mm long, 5 mm wide, broadly and rather deeply notched; lateral lobes rounded, 
0.5-0.75 mm long, 1.5-1 .75 mm wide; style and upper stamens slightly longer than upper 
lip; nutlets oblong, tapering at both ends, brown, 1 .5-1 .6 mm long, 0.6 mm wide. Fl. 
June-July; fr. from August. 
321 Along banks of mountain streams in subalpine belt. — Centr. Asia: Pam.-Al. (southern 
slope of Gissar Range). Endemic. Described from Maikhura River, tributary of Varzob 
River. Type in Leningrad. 

Note. Differs sharply from all species of the Densiflorae series in having small flowers 
and in specific shape of leaves. Particular resemblance to N. kokamirica Rgl., from which 
it differs in having, besides small flowers, also inflorescences, usually without remote 
whorls, different pubescence of stems and leaves, the latter being smaller. 

13. N. odorifera Lipsky in Tr. Bot. sada, XXIll (1904) 228; O. and B. Fedch. Perech. 
rast.Turk. V, 143. 

Perennial, to 85 cm high and to 2.5-6 mm across, strongly branching nearly from 
base; branches well developed, almost aU terminating in inflorescences; stem and branches 
grayish due to rather long crisp, more or less interwoven, hairs and also densely covered 
with very short glandular hairs; leaves on both sides densely (to tomentum) covered with 
simple hairs interspersed with glandular ones; lower side of leaf covered with closely 
crowded, punctate yellow glands, on stems 1 .5-2. 5mm long, 1 .5-2. 2mm wide, cordate or 
ovate-triangular, with cordate or truncated base, acute or short-acuminate, sharp-toothed, 
lower leaves on petioles 4-6 (8) mm long, upper on short petioles 2-3 mm long; leaves of 
branches smaller, of the same shape as on main stem. Semiverticels sessile or on short, 1.5- 
5 mm long, pedicels; lower two-four verticillasters remote, others forming oblong elliptical 
spicate inflorescence, on stem 4-6 cm long (on lateral branches smaller or ovoid); bracts 
and calyces purple-violet, densely covered with short glandular hairs, along nerves and at 
margin long, white, simple hairs; bracts from narrow-lanceolate to narrow-linear, 2.5- 
4.5 mm long, sometimes equaling length of calyx-tube, sometimes reaching the middle of 
it; calyx 7-8 mm long, 2-2.5 (3)mm wide, narrow, cylindrical-ob conical, three upper teeth 



214 



oblong-triangular or lanceolate-triangular, acuminate, 1/3-1/2 time as long as tube, two 
lower teeth usually narrower, sometimes longer to lanceolate-linear; corolla 16-17 mm 
long, with tube exserted approximately to 1/3 of its length from calyx, upper Up 3- 
4mm long, notched to about the middle to form two obtuse broad lobes; lower lobe of 
middle Up sessile, 3-3.5 mm long, 5-7.5 mm wide, with or without small apical notch, 
lateral lobes triangular, 1 .5 mm long, 2.5 mm wide; nutlets brown, oblong, 2.5 mm long, 
0.75-1 mm wide. Fl. to September; fr. September. 

Subalpine and alpine belts. — Centr. Asia: Pam.-Al. (Darvaz and surrounding area). 
Endemic. Described from Kugi-Frush Mountains. Lectotype in Leningrad. 



322 Series 3. Bucharicae Pojark. — Upper verticillasters forming a short-cylindrical or 
ovoid terminal inflorescence, the lower 1-3 remote; calyx with obUque throat and lanceo- 
late or triangular teeth; middle lobe of lower lip of corolla with remotely and coarsely 
crenate margin; bracts usually shorter than calyx; cauline leaves ovate to lanceolate, the 
largest (3.5) 4-9 mm long. Plants of subalpine and alpine belts. 

Besides the Central Asian species, N. clarkei Hook. fil. (W. Himalayas) and N. pubes- 
cens Benth. (E. Afghanistan) should also be included in this series. 

14. N. bucharica Lipsky in Tr. Bot. sada, XXIII (1904) 217; O. and B. Fedch. Perech. 
rast.Turk. V, 140. 

Perennial, to 80 cm high; root thick, woody, coarsely fibrous, passing above into multi- 
cipital rhizome covered with brown squamiform leaves; stems erect or ascending, at base 
8 mm across, more or less densely covered with small sessile capitate glands intermixed 
with very long spreading simple hairs, mostly branching nearly from base; branches sterile 
or the upper and sometimes nearly all with small inflorescence at summit; cauline leaves 
generally 3.3-3.5 cm long, 2.4-3 cm wide, rarely to 4.8-5.5 cm long and 3.5-4 cm wide, 
ovate or broadly ovate (on axillary branches narrower and smaller), acute or obtuse, with 
cordate or deeply cordate base, at margin rather large-toothed or crenate, covered with 
flexuous simple hairs (these longer on veins) and small whitish sessile glands; punctate 
oil glands copious on lower side, sparse on the upper; petioles 1/4-1/3 (1/2), the upper 
ca. 1/8 as long as blade; lower floral leaves resembling upper cauline leaves, the upper 
narrower, mucronate, subbracteiform; terminal inflorescence subglobose to oblong-ovoid, 
with 1-3 (4) remote verticillasters below; semiverticels of lower verticillasters distinctly 
peduncled, the others subsessile; bracts lanceolate to hnear, green or violet, crisp-hairy at 
margin, reaching the middle of calyx or the base of its teeth; calyx 6.5-9 mm long, 2- 
2.5 mm wide, green or violet, densely covered with crisp hairs; upper teeth triangular or 
triangular-lanceolate, 1/3-1/2 as long as tube, the lower narrower, to linear-lanceolate, all 
terminating in a rather rigid but not spinous point; corolla 14-19 mm long, lilac-blue, 
pubescent outside, the narrow part of tube half as long again as calyx; upper lip cleft to 
1/3-1/2 into broadly ovate obtuse lobes; middle lobe of lower lip 3-4 mm long, 5-7 mm 

323 wide, broadly and rather deeply emarginate, the lateral lobes 2-2.5 mm long, 2-3 mm wide, 
triangular or ovate-triangular; stamens and style slightly longer than upper lip; nutlets 
elongate-eUipsoid, 1 .75-2.5 mm long, 0.75-1 mm wide, brown, dull. Fl. second half of 
June to first half of October; fr. from August. (Plate XX, Figure 1.) 



215 



Subalpine belt, in grass plots, banks of streams near waterfalls. - Centr. Asia: T. Sh. 
(Chatkal, Talass, Fergana, Uzun-Akhmat ranges), Pam.-Al. (Gissar, Peter the First ranges, 
Darvaz and surrounding area). Endemic. Described from mountain pass Sagyrdasht in 
Darvaz Range. Type (lectotype) in Leningrad. 

Notes. 1) Crosses between this species and N. formosa Kudr. (see Note to N. for- 
mosa) occur frequently. 

2) Apparently there is a disruption in the distribution area of N. bucharica between 
the Pamir-Alai section and the W. Tien Shan section. Painstaking comparison of the her- 
barium specimens from within these two sections has provided no basis for separating from 
N. bucharica the two species observed in the S.N. Kudryashev herbarium: N. ferganen- 
sis Kudr. (specimen from Uzun-Aklimat Range) and N. talassica Kudr. (mountain pass 
Karagoin ravine in the western part of Talass Ala-Tau). The shape of the calyx and corolla, 
character of indument, shape and color of the nutlets of these "species" are the same as in 
the typical N. bucharica. Also the shape of the inflorescence and leaves does not 
depart from variations within N. bucharica. All that can be pointed out is the consis- 
tent deep violet of the calyx and bracts in specimens from W. Tien Shan while, among the 
Pamir-Alai plants, specimens often occur with a weakly pigmented calyx which clearly 
does not provide a basis for establishing the Fergana plant as (one or more) species. For a 
definitive solution comparisons must be made of the plants in nature in both areas of N. 
bucharica. Characters that are lost in the herbarium, such as corolla color and scent, 
could then be determined. 

Economic importance. The plant yields a pale yellow essential oil with a strong odor. 
The properties of the oil have not yet been investigated (Kudryashev, 1932). 

15. N. maussarifii Lipsky in Tr. Bot. sada, XXIII (1904) 216; 0. and B. Fedch.Perech. 

rast.Turk. V, 140. 

Perennial, 45-75 cm high; root thick, to 2.5 cm across, woody, coarsely fibrous, passing 
into a thick multicipital rliizome; stems erect, 2-4 mm across, densely leafy, velutinous 
with very short simple hairs and farinaceous with minute sessile glands, branching nearly 
from base; branches mostly sterile, few terminating in inflorescence; basal leaves brown, 
squamiform; cauline leaves grayish on both sides with short flexuous hairs, ovate or tri- 
324 angular-ovate (often narrower on axillary branches), to 3.5-4 cm long, 2.5-3 cm wide, 
acute or obtuse, with cordate or some with subhastate-cordate base, mostly distinctly or 
obscurely crenate, partly entire or acutely toothed; petioles gray-tomentose-pubescent, 
the lower 2/5-1/2 as long as blade, the upper leaves 2/7; all verticillasters crowded in 
ovoid spicate terminal inflorescence or 1-2 (3) lower remote; lower floral leaves resem- 
bling the cauline, the upper much smaller, entire, sessile; bracts violet, narrowly lanceo- 
late, reaching the middle of calyx or the base of its teeth; leaves subtending semiverticels 
slightly broader and longer, beset outside with small whitish glands, at margin villous-hairy; 
calyx 14-15-(17) nerved, violet, 7-8 mm long, whitish with spreading crisp hairs, the upper 
teeth triangular-lanceolate or lanceolate, 1/3-1/2 as long as tube, the lower longer and nar- 
rower; corolla bluish, 12-16 mm long, pubescent, the narrow part of tube slightly exsert- 
ed, abruptly dilated into wide throat; upper lip 2.5-3.3 mm long, cleft to middle into 
broad obtuse lobes; middle lobe of lower lip 3-4 mm long, 4-5.5 mm wide, broadly emar- 
ginate, with slightly sinuate-dentate margin, the lateral lobes 1.3-1.5 mm long, 2-2.5 mm 



216 



wide, semiorbicular or triangular-semiorbicular; stamens and style nearly as long as upper 
lip; nutlets oblong-obovoid, dark brown. Fl. July-September; fr. from end of July. 

Central Asian juniper and alpine belt, banks of small rivers and streams. — Centr. Asia: 
Pam.-Al. (western part of Zeravshan Range and Darai-Nikhan Mountain). Endemic. De- 
scribed from vicinity of Mausarif village in western part of Zeravshan Range. Type in 
Leningrad. 

16. N. alatavica Lipsky in Tr. Bot. sada, XXIII (1904) 214; 0. and B. Fedch. Perech. 
rast. Turk. V, 140. - N. menthoides /3. virescens Rgl. in Tr. Bot. sada, VI, 2 (1880) 
360. - N. kokanica Franch. in Ann. Sc. Nat. VII ser. XVIII (1884) 230, non Rgl. 

Perennial, to 1 m high; stems in lower part to 4-5 mm across, branching from base, 
densely covered with short spreading hairs; branches slender, densely leafy; leaves dense- 
ly covered on both sides with crisp hairs, narrow, oblong-ovate to lanceolate, with cuneate 
or rarely subtruncate base, gradually acuminate, at margin with acute (rarely obtuse) teeth, 
to 4.5 cm long, 18 mm wide; verticillasters dense, 1 or 2 lower remote, others grouped in 
ovoid or oblong spicate terminal inflorescence; lateral axillary branches also usually ter- 
minating in inflorescence; floral leaves lanceolate or linear, often narrowly linear, shorter 
325 than calyx; calyx 5-6.5 (7) mm long, like bracts blue-violet, densely covered outside with 
long crisp implexed hairs, 15-nerved, strongly oblique, the upper teeth lanceolate-triangu- 
lar, 1/3-2/5 as long as tube, the lower lanceolate, all long-acuminate but not aristate; 
corolla bluish, pubescent outside, 10-13 mm long, twice the length of calyx, the narrow 
part of tube slightly exserted; upper lip ca. 2.5 mm long, cleft slightly beyond middle into 
obovate obtuse lobes; middle lobe of lower lip broad, ca. 2.5 mm long, 5 mm wide, shal- 
lowly emarginate, with slightly sinuate or coarsely crenate-undulate margin, the lateral 
lobes broad, orbicular-triangular, 1 mm long, 2 mm wide; lower stamens equaling upper 
lip, the upper 1 .2 mm longer; style as long as upper stamens; ripe nutlets dark brown, 
ellipsoid, 1 .3-1 .5 mm long, 0.6-0.7 mm wide. Fl. June-August; fr. from August. 

Subalpine and alpine mountain belts. - Centr. Asia: T. Sh. (Talass, Pskem and Ugam 
ranges). Endemic. Described from Talass Range. Type dectotype) in Leningrad. 

17. N. schugnanica Lipsky in Tr. Bot. sada, XXIII (1904) 220; 0. and B. Fedch. 
Perech. rast. Turk. V, 140. 

Perennial, to 75 cm high or slightly higher; stems to 6 mm across, ascending, brown in 
lower decumbent part, with few pairs of brown squamiform leaves, branching nearly from 
base, more or less viDous with long crisp white hairs intermixed with small sessile scabUke 
glandular hairs; leaves to 6 cm long, 5 cm wide, orbicular or broadly ovate, with deeply 
cordate base, short-acuminate or obtuse apex, coarsely crenate, grayish-velutinous on both 
sides, the hairs implexed, crisp, simple, dense at first, later partly disappearing, especially 
in lower leaves; leaves of axillary branches sometimes narrower than the cauHne to oblong- 
ovate; petioles villous-hairy, 1/3 as long as blade in lower leaves and 1/7 in upper; upper 
verticillasters crowded at apex of stem and of axillary branches in ovoid-spicate inflores- 
cence, the lower 1 or 2 remote; lower floral leaves resembhng the upper cauhne, some- 
times exceeding flowers, the upper narrower, always shorter than flowers; bracts linear- 
lanceolate or linear, green, reaching calyx-teeth, rarely shorter than calyx, sometimes 
nearly as long; calyx 13-toothed, green, densely covered, like bracts, with whitish scablike 



217 



glands and punctate aromatic glands, with long crisp hairs along nerves and margin, 7- 
8 mm long, 2-2.5 mm wide, the throat strongly oblique; upper teeth triangular or lanceo- 
late-triangular, approximately 1/3-2/5 as long as tube, the lower narrowly lanceolate, 
326 long-acuminate, all teeth without rigid mucro; corolla 13-15 mm long, bluish, covered 
outside with crisp hairs, the narrow part of tube exserted to 1/4, abruptly expanding into 
broad neck; upper Up approximately half as long as the lower, cleft slightly beyond mid- 
dle into 2 broadly obovate or sometimes nearly semiorbicular obtuse lobes; middle lobe 
of lower lip oblate, 2-3 mm long, 4-4.5 mm wide, emarginate, the lateral lobes much 
smaller, 1 .3-1.5 mm long, 2-2.5 mm wide, broadly triangular or nearly semiorbicular; 
upper stamens and style slightly exceeding upper lip. Fl. August. 

Stony mountain slopes in subalpine belt. - Centr. Asia: Pam.-Al. (Shugnan). Endemic. 
Known only from location: BakJian, between Nishus and Anderob villages on Pyandzh 
River. Type in Leningrad. 



Series 4. Supinae Pojark. - Verticillasters forming a globose or subglobose-ovoid 
capitate terminal intlorescence, lowermost whorl (very rarely 2) remote; tliroat of calyx 
strongly obUque or 2-lipped, the teeth much shorter than tube, lanceolate or triangular; 
middle lobe of lower lip of corolla with slightly sinuate or entire margin, bracts sliglitly 
shorter than calyx; cauline leaves broad, small, largest 1.5-2.5 mm long. Low plants of 
high mountains. 

In addition to the three Central Asian and two Caucasian species, N. eremocosmos 
Rech. til. (central Elburz in N. Iran) and N. pseudosupina Pojark. (Kaslimir) should be 
included in this series. 

18. N. supinaStev. inMem. Soc. Nat.Mosc. Ill (1812)265; M. B. Fl. taur.-cauc. Ill, 
391 ; Ldb. Fl. Ross. Ill, 373; Benth. Lab. gen. at sp. 473; Boiss. Fl. or. IV, 642; Grossg. 
Opred. rast. Kavk. 331. - N. fissa Benth. 1. c. 471 (non 737) quoad, descr. (excl. loco), 
non C. A.M. ~ Dracocephalum caucasicum Lipsky et Akinf. in Tr. Bot. sada, XIII 
(1894) 326. - Nepeta caucasica Somm. et Lev. in Nouv. Giorn. Bot. Ital. IV, 2 (1897) 
298. -Ic: Tr. Bot. sada, XVI, Plate XLI.-Exs.: GRF,No.223; Fl. Cauc. exc. No. 293. 

Perennial; root fibrous, more or less twisted, passing above into brandling rliizome, 
this Uke toot covered with blackish-brown bark and squamiform leaves; stems numerous, 
10-30 cm long, ascending or nearly decumbent, more or less densely covered with short 
jointed simple hairs and small whitish capitate glands, not branching or with few axillary 
branches, the lower branches often decumbent, long but only rarely developing inflores- 
cence; other branches short, with undeveloped internodes; leaves yellow-green, when 
i327 young often rather densely covered with shaggy simple hairs and very small glandular ones, 
also with punctate glands beneath, later often subglabrous except on veins; cauline leaves 
1-2 (2.5) cm long, 0.7-1 .8 (2) cm wide, ovate, with truncate or rounded or sometimes 
cuneate or shallowly cordate base, acute or obtuse; leaves on branches smaller and often 
narrower, the margins evenly large-toothed or almost crenate, the petioles 2-3 mm long; 
upper leaves sessile; semiverticels subsessile, forming a rather dense subglobose-ovoid 
capitate inflorescence, sometimes 1 verticillaster remote; floral leaves resembling upper 
cauline leaves, slightly shorter than calyx, lanceolate-linear or linear, covered like calyx 



218 



with very small glandular hairs, the margin with a dense fringe of long simple jointed hairs; 
calyx 7-8.5 mm long, slightly curved, blue in lower part, rarely all blue, 2-lipped or nearly 
so, the 3 upper teeth triangular, acute or more or less acuminate , (1 /6) 1 /4- 1 /3 as long as 
tube, the 2 lower lanceolate or narrowly lanceolate, long-acuminate or sometimes nearly 
subulate-acuminate, half as long again to twice as long as upper teeth; corolla bright blue, 
14-20 mm long, the tube exserted nearly half its length, expanding above into large neck, 
4.5-5.5 mm long, 5-6 mm wide; upper Up 2.5-3.3 mm long, cleft nearly to base into 
obtuse broadly ovate lobes; lower lip with subtetragonal middle lobe, 3-4 mm long, 6- 
7 mm wide, the semiorbicular lateral lobes 1-1 .3 mm long, 2-2.5 (3) mm wide; upper 
stamens as long as upper Hp; lower stamens distinctly shorter than the upper; nutlets 
dark brown, elongate-ellipsoid, 2 mm long, 0.75-1 mm wide. Fl. second half of July to 
September; fr. from August. 

Stony and gravelly places in alpine belt, moraines, pebbles, rock crevices. — Caucasus: 
Main Range, Dag., E. Transc. (Kuba area). Endemic. Described from Main Range, Khod- 
zhal River near foot of Tfan-Dag peak. Type in Helsinki, topotype in Leningrad. 

Note. The taxonomic position of this species has not been satisfactorily elucidated to 
date. In their systems, Boissier and Briquet placed it in section Cataria although, except 
for the smooth nutlets, N. supina has nothing in common with the other species in this 
section either in the structure of flowers and inflorescence or in habit. 

19. N. buschii Sosn. et Mand. in Zam. po sist. i geogr. rast. Bot. inst. AN Gruz. 16 
(1951) 10. -N. supina auct.: Grossg. Fl. Kavk. Ill (1932) 293; idem., Opred. rast. Kavk. 
331, p.p. non Stev. - N. citriodora Sosn. et Mand. in Bot. zhurn. SSSR, XXXIX (1949) 
287, non Dum. nee Baker. 

Perennial, 15-50 cm high; root as in N. supina; stems numerous, erect from ascend- 
ing base, in lower part to 3.5 mm across, more or less densely covered with long simple 
white jointed hairs and very small whitish glands, with internodes to 2.7 cm long (in lower 
part to 10 cm), usually branching; branches partly short, sterile, partly terminating in 
inflorescence and reaching 25 cm in length; leaves pale green, 1 .5-2.5 cm long, 1-2.2 cm 
wide, ovate, cuneate at base, mostly acutish, along margin with large obtuse teeth, rather 
densely covered between the veins above and mostly on the veins beneath, with appressed 
shaggy hairs and often rather copious minute sessile whitish glandular hairs as well as 
sparse yellow resinous punctate glands; petioles 1-2.5 mm long; verticillasters at ends of 
stems and of lateral branches forming an ovoid capitate inflorescence 2-2.5 cm long, some- 
times 1 verticillaster remote; floral leaves resembUng the cauUne leaves but distinctly 
smaller, lanceolate, long-acuminate, slightly shorter than calyx; calyx 9-1 1 mm long, 
2-lipped, usually sparsely (rarely densely) covered, like bracts, with minute glandular hairs, 
on nerves and margin with simple shaggy jointed hairs, these forming a long fringe at the 
margin of teeth; lower teeth lanceolate to lanceolate-subulate, 3.5-4.3 mm long, 2/5-1/2 
as long as tube; upper teeth narrowly triangular, acuminate, 2.5-2.8 mm long; corolla 
20-27 mm long, bright blue, the tube 10-16 mm long, curved, abruptly expanding above 
into large neck, 5-6 mm long, 5.5-6.5 mm wide; upper lip 3.5-4 mm long, cleft to 2/3 
into obtuse lobes, 2.5-2.8 mm wide; lateral lobes of lower lip broadly triangular, 2- 
2.5 mm long, 2.7-3 mm wide, the middle lobe 4-5 mm long, 7-8 mm wide; nutlets 
oblong-obovoid, brown, 2.5 mm long, 1 mm wide. Fl. July-August; fr. August. 



219 



Alpine belt, stony and gravelly placers. - Caucasus: S. Transc. (Aragats and Kapud- 
zhikh mountains). Gen.distr.: Arm.-Kurd. (Mount Ararat). Described from Aragats 
Mountain in Armenia. Type in Tbilisi. 

Note. A species very closely allied to N. supina Stev. from which it differs mainly 
in having much larger flowers and more erect, higher and thicker stems. 

Economic importance. The plant has a very strong lemon scent and may possibly be 
of value in the perfume industry. An ornamental. 

20. N. kokanica Rgl. in Izv. Obshch. lyub. est. antr. i etn. XXXIV, 2 (1882) 65; 0. 

329 and B. Fedch. Perech. rast. Turk. V, 141, p.p. - N. cephalotes var. brevipedunculata 
Rgl., op. cit. 65. - N. pamiroalaica Lipsky in Tr. Bot. sada, XXIII (1904) 230, p. p. 

Perennial, grayish or whitish with dense indument; root to 4 cm across, dark brown, 
twisted, strongly splitting, passing into rhizome, the numerous, dark brown, more or less 
woody stem bases hidden under rubble, covered, like the rhizome, with squamiform leaves; 
stems (5) 10-40 cm high (generally 20-30 cm), slender, ascending or erect, branching all 
the way up, more or less densely tomentose, often (var. hissarica (Lipsky) Fedtsch.) with 
loose white tomentum of long implexed hairs at nodes and particularly under inflores- 
cence; branches mostly short, rudimentary, seldom developed, terminating in inflores- 
cence; leaves rather thick, covered on both sides with dense tomentum; cauline leaves 
5-15 cm long and wide, orbicular or rhombic-ovate, at times subreniform, obtuse or short- 
acuminate, with entire cuneate base and toothed or crenate margins, all petiolate, the 
petioles 2/7-2/3 as long as blade; leaves of axillary branches smaller and often narrower; 
terminal head (2) 2.5-3.5 cm across, 1.7-3.5 cm long, sometimes a solitary remote verti- 
cillaster present below; floral leaves resembling the cauline but smaller and often narrower, 
shorter than calyx, narrowly linear, covered outside, like calyx, with small capitate glands, 
at margin with a dense fringe of hairs, these long, very fine, implexed, simple, jointed 
(consisting of 5-18, sometimes to 28 cells), usually unbranched, occasional hairs with 1-2 
rudimentary branches of 1 (2) cells; calyx 6-7.5 mm long, with strongly oblique throat, 
more or less violet, the teeth narrowly triangular or tubular-lanceolate, usually tapering 
from the middle to a narrow nearly subulate point, all nearly equal, the upper 2/3 as long 
as tube; corolla (pale?) blue, 15-1 8 mm long, slightly pubescent and glandular outside, the 
tube exserted 2-2.5 mm; upper hp 2.7-3.25 mm long, cleft to 1/3 into obtuse obovate 
lobes; middle lobe of lower lip 3.8-4.5 mm long, 6-6.5 mm wide, with broadly emargi- 
nate lateral lobes 1 mm long, 3 mm wide, nearly semiorbicular; nutlets dark brown, 
slightly attenuate toward base, trigonous, 2-2.3 mm long, 1-1.8 mm wide. Fl. second half 
of July to September; fr. second half of August (Plate XX, Figure 2.) 

Gravelly and stony placers, rock crevices in alpine belt. — Centr. Asia: Pam.-Al. (Alai, 
Turkestan, Peter the First, Zeravshan, Gissar and Darvaz ranges). Gen. distr.: Dzhu.-Kash. 
Described from the northern slope of Turkestan Range, Dzhiptyk pass (upper part of basin 
of Isfara River). Type in Leningrad. 

330 Notes. 1) Lipskii renamed this species N. pamiroalaica Lipsky because Regel', in 
describing N. kokanica, had erroneously referred some specimens to another species 
(under the name "N. cephalotes Boiss. var. brevipedunculata Rgl.")- The diagnosis 
of N. kokanica itself is not quite accurate, but these mistakes could hardly provide justi- 
fication for renaming N. kokanica Rgl. which was described in keeping with the rules of 



220 



(331) 




PLATE XX. 1 - Nepeta bucharica Lipsky, summit of plant, flower, nutlet; 2 - N. kokanica Rgl. 
general aspect, flower, nutlet; 3 - N. ladanolens Lipsky, summit of plant, flower, nutlet. 



221 



international botanical nomenclature, and in accordance with these rules the name N. ko- 
kanica Rgl. should be retained. In the process, Lipskii, on the assumption that he was 
merely renaming N. kokanica Rgl., extended the range of this species and included in it 
N. pamirensis Franch., a species that morphologically and geographically differs from 
N. kokanica Rgl. (see Note on N. pamirensis) and whose independent status is here- 
with restored. 

2) For some reason, Lipskii left the taxonomic position of N. kokanica unresolved, 
even though he legitimately included closely related species of the series Bucharicae in the 
section Spicatae. In "Perechen' rastenii Turkestana," 0. and B. Fedchenko placed this 
species in the section Capituliferae without any justification whatsoever. 

21. N. pseudokokanica Pojark. in Bot. mat. gerb. Bot. inst. AN SSSR, XV (1953) 291. 
Perennial, easily distinguished from N. kokanica Rgl. in outer appearance by the 

greenish or even green leaves, covered on both sides with short crisp 2-5-jointed hairs, 
these distinctly spreading on veins; stems, branches and petioles more or less densely 
covered with similar spreading hairs; stems ascending or often arcuately curved, branching 
from base; branches developed, with several pairs of leaves, branching and usually termi- 
nating in inflorescence; cauline leaves ovate, broadly ovate, with rounded-cuneate or cu- 
neate base, mostly acute, the margin with 4-5 rounded, often outward-curved teeth; in- 
florescence terminal, capitate; remote verticillasters 1 or occasionally 2; floral leaves 
resembling the cauline or bracteiform; bracts shorter than calyx, lanceolate or lanceolate- 
linear, long-acuminate, densely covered outside, like calyx, with very small capitate glands 
and simple, the nerves and margin rather densely covered with eglandular hairs, these 3-5 
(8)-jointed, distinctly thicker toward base; calyx 7.5-9 mm long, violet, with strongly 
oblique throat; teeth lanceolate, produced into nearly subulate mucro, the 3 upper 2/3 
333 as long as tube; corolla 17-18 (19) mm long, the tube distinctly exserted to 2-3 mm; 
upper lip cleft to 2/3 into obtuse lobes; middle lobe of lower Up oblate, the lateral lobes 
1 mm long, 3 mm wide, rounded; upper stamens as long as upper lip or barely exserted; 
nutlets brown, lustrous, 1 .75-2.25 mm long, 1-1 .25 mm wide, obovoid, markedly attenu- 
ate toward base, slightly truncate at the broad upper end. Fl. from July; fr. from first 
half of August. 

Rocky and stony places in alpine mountain belt. -- Centr. Asia: Pam.-Al. (Fergana 
Range). Endemic. Described from natural boundary Ak-Boguz in the southwestern part 
of the range. Type in Leningrad. 

Note. This species is easily distinguished from both Central Asian species of the series 
Supinae by its habitually scantier indument and greenish or green (not gray or whitish) 
leaves of thin consistency; it also differs from N. pamirensis in the nonbranching hairs 
and in the corolla-tube being relatively longer in relation to the calyx. 

22. N. pamirensis Franch. in Bull. Mus. Hist. Nat. II (1896) 345. - N. pamiro-alaica 
Lipsky in Tr. Bot. sada, XXIII (1904) 230, p. p. -N. kokanica 0. et B. Fedtsch.Perech. 
rast. Turk. V (1913) 141, p.p. non Rgl. - ? N. oxicola Franch. 1. c. 346. -N. supina 
Duthie in Alcock, Repert. nat. hist, results Pamir bound. Comm. (1898) 25, non Stev. 

Perennial; root and rhizome as in N. kokanica Rgl., which this species resembles in 
outward appearance; stems numerous, tightly crowded, woody in lower part, erect or 



222 



more rarely ascending, usually 15-20 cm high (rarely higher), densely covered with branch- 
ing jointed hairs; branches produced from all internodes but frequently sterile and partly 
rudimentary, only rarely in the largest specimens upper branches with small terminal inflo- 
rescence; leaves gray, with dense short tomentum on both sides, on stems 4-14 mm long, 
5-1 1 mm wide, flabelliform or rhombic-obovate, with entire cuneate base and evenly cre- 
nate margin; petioles tomentose, 1/4-1/3 (rarely 1/2-2/3) as long as blade; upper leaves 
subsessile; floral leaves resembling the cauline but smaller; inflorescence globular or sub- 
orbicular-obovoid, 1.5-3.5 cm long, rarely with 1 remote verticillaster below; bracts nar- 
rowly linear, shorter than calyx, resembling the cauline but smaller, densely covered out- 
side with small glandular hairs intermixed with punctate yellow glands; long, very fme, 
arachnoid, implexed, white, jointed, branching, partly 2-branched hairs forming a thick 
fringe at the margin and also covering the nerves and the margin of teeth; calyx suffused 
334 with blue-violet, 6.5-9 mm long, densely covered with small glandular hairs and punctate 
glands; throat strongly oblique; teeth lanceolate, long-acuminate, the 3 upper 1/2-3/4 as 
long as tube; corolla bluish-violet (?), 12.5-16.5 mm long, the tube not exserted from 
calyx or scarcely exserted to 1 (1.5) mm; upper lip cleft to 2/3-3/4, the obtuse obovate 
lobes 1.75-2.75 mm long, 2 mm wide; middle lobe of lower hp broad, 3.5-4.5 mm long, 
5.5-8.5 mm wide, emarginate and sometimes with coarsely sinuate margin, the lateral lobes 
1-1 .25 mm long, 2.3-3 mm wide, nearly semiorbicular; nutlets 2-2.5 mm long, 0.75-1 mm 
wide, elongate-obovoid to oblong-ellipsoid, brown. Fl. July-October; fr. from August. 

In alpine belt, mixed grass-sheep's fescue and wormwood-meadowgrass steppes, cobre- 
sia-sedge meadows, exposed stony and gravelly slopes, moraines, dry pebble-beds. — Centr. 
Asia: Pam.-Al. (W. and E. Pamir). Gen. distr.: Afghanistan, areas adjacent to Pamir and 
Ind.-Him. (Hindu Kush and NW Himalayas). Described from Bazai-Gumbez (upper part of 
basin of Vakhan-Darya River) in northeastern part of Afghanistan. Type in Paris. 

Note. In habit N. pamirensis Fr. does not differ from N. kokanica Rgl. but is 
clearly differentiated from this species by two characters: 1) strongly branching hairs, 
with branches consisting of several cells (joints) and branching in turn; and 2) the narrow 
part of corolla-tube scarcely exserted from the calyx. The distribution area of N. pami- 
rensis differs from that of N. kokanica. N. pamirensis has been unjustly neglected 
and has never appeared in the Russian literature. This is perhaps explained by the fact 
that Lipskii, in his time, having found no differences between N. pamirensis Fr. and 
N. kokanica Rgl., included it in the latter which he inaccurately named N. pamiro- 
alaica Lipsky (see Note to N. kokanica). 



Section 3. Capituliferae (Benth.) Pojark. comb. nov. - Sect. Pycnonepeta § Capitu- 
hferae Benth. in DC. Prodr. XII (1848) 379. - Sect. Eunepeta § Capituliferae Boiss. Fl. 
or. IV (1879) 637; Briq. in Pflanzenfam. IV, 3a, 236. - Calyx tubular, slightly curved, 
with more or less oblique or nearly 2-hpped throat and narrowly triangular teeth; corolla- 
tube curved and abruptly expanding into short neck; middle lobe of lower lip with swell- 
ing at base, concave; nutlets narrowly ellipsoid, smooth, dull; inflorescence consists of 
a headlike terminal verticillaster (or of several confluent in a head) and of many remote 
pairs of dense headlike semiverticels, these with elongate peduncles and forming a very 
335 lax racemiform or paniculate inflorescence or else sessile or partly subsessile and then 



223 



forming an interrupted spike. Perennial plants or sometimes subshrubs, mostly patent- 
branching at base. 

Many species in this section are distributed in W. Himalayas and in Hindu Kush; a num- 
ber of species grow in the south of Central Asia, and a few are known from Iran. 



Subsection 1 . Podocephalae Pojark. subsect. nov. in Addenda XIX, 354. - All semi- 
verticels( except sometimes the uppermost) with elongate peduncles; aggregate inflores- 
cence loosely racemiform or paniculate; throat of calyx strongly oblique, nearly 2-lipped. 



Series 1. Floccosae Pojark. — Leafy branches produced only on lower part of stem; 
leafless peduncles terminating in head-shaped semiverticel developing in the axils of upper 
leaves. Aggregate inflorescence a raceme. 

In addition to the two Central Asian species, this series also contains N. floccosa 
Benth. (Kashmir and E. Afghanistan), N. monticola Kudr. (SE Afghanistan) and N. 
cephalotes Boiss. (Iran). 

23. N. vakhanica Pojark. in Bot. mat. gerb. Bot. inst. AN SSSR, XV (1953) 296. - N. 
floccosa 0. et B. Fedtsch. Perech. rast. Turk. V (1913) 140; Kudryash. in Bot. mat. gerb. 
Bot.inst. Uz. fil. ANSSSR, I, 1, 10, p. p. non Benth. - Ic: Poyark., op. cit. Figure 3. 

Perennial; stems 25-30 cm long, ascending, 1.5-3.5 cm across, often reddish, leafy only 
in upper part, branching at base; branches as long and as tliick as stems and densely cover- 
ed, like stems, with very small capitate sessile or short-stipitate glands forming a yellowish 
bloom, when young also covered near base with long fine arachnoid simple jointed hairs; 
2-3 lower internodes 0.7-2.5 cm long, others except the upper 6-9.5 cm long; leaves in 
2-3 approximate nodes, often rosulate, with petioles \^/2-2^/i times as long as blade; leaves 
of a pair situated at tlie level of lower fourth or third of the stem short-petioled or subses- 
sile, blade cordate, 1 .5-3 .5 cm long and wide, obtuse, crenate at margin, thin, soft, greenish 
or grayish, more or less densely covered on each margin with thin and loose white tomen- 
tum, subglabrous along the veins and thus appearing rugose; semiverticels hemispherically 
head-shaped, disposed at ends of stem (and radical branches) and of very distant axillary 
peduncles, forming a very lax racemiform or in lower part rarely paniculate, inflorescence 
comprising 2/3-3/4 of the plant; axillary heads (8) 10-65-flowered, to 13 mm long, 
336 20mm wide, much larger than the 2-8-flowered terminal head; peduncles 3-5 cm long 
(the uppermost shorter), mostly arcuately curved, divergent, often at nearly a right angle; 
floral leaves and leaves subtending tlie heads narrowly linear, 4.5-9 mm long, 0.5-1 mm 
wide; bracts smaller, to subfiliform, violet, with a broad fringe of white or violet arachnoid 
implexed hairs, approximately 3/4 as long as calyx; calyx (7) 8-9 mm long, slightly curved 
above, with nearly 2-Upped throat, violet, densely covered outside with small capitate 
glands intermixed (often copiously) with punctate yellow glands, the nerves with fugacious 
lanate-arachnoid hairs; teeth narrowly triangular, long-acuminate, ciliate, the upper 2/7- 
1/4 as long as tube, the 2 lower broader at base than the upper; corolla 10-12 cm long, 
blue, hairy outside and covered with capitate sessile glands, the tube narrow, not exserted 
or exceeding calyx by 1-1 .5 mm, abruptly expanding into short neck; upper lip 1 .5-2 mm 



224 



long, deeply cleft into obtuse lobes; middle lobe of lower lip 2 mm long, 3.5-4 mm wide, 
coarsely crenate-incised, the lateral lobes triangular-semiorbicular, 1.3-1.5 mm long; nut- 
lets narrowly ellipsoid, pale brown, 2.5 mm long, 0.75 mm wide. Fl. July-August; fr. 
from first half of August. 

Sliding gravelly taluses and between stones in alpine belt. — Centr. Asia: Pam.-Al. 
(southern part of W. Pamir: Vakhan). Endemic. Described from Pyandzh River between 
mountain passes Kizil-Kirchin and Mats. Type in Leningrad. 

24. N. pseudofloccosa Pojark. in Bot. mat. gerb. Bot. inst. ANSSSR, XV (1953) 293.- 
N. floccosa Lipsky in Tr. Bot. sada, XVIII(1900j 100; 0. and B. Fedch. Perech. rast. 
Turk. V, 140, p. p. non Benth. 

Perennial; stems 15-35 cmlong, slightly curved in lower part, densely covered with 
very small sessile and short -stipitate capitate glands and, when young, with very short 
dense white tomentum, the lower and upper internodes short, the middle ones 6-9 mm; 
leaves at 3-4 lower nodes rosulate long-petioled, cordate, or rarely broad-ovate, crenulate; 
leaves of a remote pair higher up similar but smaller and short-petioled; blade 1 .5-2 cm 
long, 1 ;3 mm wide, firm, thick, densely covered on both sides with short whitish tomen- 
tum, the network of veins impressed above, prominent beneath, not tomentose; semiver- 
ticels capitate, hemispherical, borne at the end of stem and of distant axillary peduncles, 
337 forming a very sparse racemiform inflorescence; peduncles straight, at an acute angle, the 
lower to 8 cm long, the upper 1 .5-2 cm long; terminal verticillaster 20-70-flowered, glob- 
ular, much larger than axillary heads; floral leaves subtending semiverticels linear, to 
8 mm long, 1 mm wide, covered with small short-stipitate glands; bracts similarly glandu- 
lar and with a broad fringe of white hairs, shorter than calyx; calyx 6-7.5 cm long, curved, 
nearly 2-lipped, densely covered outside with short-stipitate and sessile glands and with 
fugacious tomentose-lanate indument, upper teeth narrowly triangular, 1/4 as long as tube; 
lower teeth broader, sometimes longer; corolla 9-1 1 mm long, densely glandular outside; 
tube exserted to 1 .5 mm, narrow, abruptly expanding into wide neck; upper lip cleft into 
oblong-elliptic lobes; middle lobe of lower Up ca. 2 mm long, 3.5 mm wide, coarsely undu- 
late-crenate, the lateral lobes broadly triangular, ca. 2mm long, 0.75 mm wide; upper 
stamens slightly longer than upper lip; nutlets oblong-ellipsoid, pale brown, 1 .75 mm 
long, 0.7 mm wide. August. 

Taluses in stibalpine belt. - Centr. Asia: Pam.-Al. (Darvazj. Endemic. Described from 
near Tavil-Dara village on Khingou River. Type in Leningrad. 

Note. In most characters, N. pseudofloccosa approaches the Himalayan N. flocco- 
sa Benth. rather than N. vakhanica. It is hardly distinguishable from N. floccosa, 
in general aspect, but it has a different indument, only one pair of developed cauline leaves, 
a sparser inflorescence, longer stamens, and less compact heads with fewer flowers. 



Series 2. Badachschanicae Pojark. — Stems leafy nearly all the way up, strongly 
branching; axillary branches many, long, leafy, fertile and developing flowering branches; 
aggregate inflorescence a strongly branching panicle. 

A monotypic series. 



225 



25. N. badachschanica Kudr. in Bot. mat. gerb. Inst, bot i zool. AN UzSSR, IX 
(1947) 18. 

Perennial; stems 30-50 cm long, robust, stout, 3-4 mm across, branching at base, gray- 
tomentulose; lower internodes 0.8-3 mm long, at middle of stem 5-6 cm, toward summit 
again shorter; axillary branches produced all along the stem, shorter toward summit, most- 
ly (except uppermost) branching and producing several pairs of short fertile branches; 
leaves on lower part of stem with petiole about the length of blade, firm, thick, 1 .5- 
4.5 cm long, 1 .5-3.5 cm wide, the upper side greenish or grayish above, pubescent or to- 

338 mentose, with impressed glabrous veins, the lower side gray, pubescent-tomentose, with 
indistinct main veins; petioles white-tomentose; lower floral leaves 7-8 mm long, 3-5 mm 
wide, elliptic, the upper Unear, 4-8 mm long, ca. 1 mm wide; semiverticels capitate, hemi- 
spherical.at the end of stem and axillary branches and of peduncles 0.4 mm (the upper) to 
1.5 (2)mm long, 8 -2 5 -flowered, 1-3 cm long, 1-2 cm wide, forming a compound panicle; 
leaves subtending semiverticels (heads) and bracts narrowly linear, acuminate, violet, cov- 
ered above and along the margin, like calyx, with long fine implexed arachnoid white hairs; 
bracts shorter than calyx; calyx 8-12 cm long, nearly 2-lipped, 15-nerved, curved in upper 
part; teeth lanceolate, acuminate, the upper 1/4-2/7 as long as tube; corolla 12-13 mm 
long, pubescent outside, with more copious and longer hairs on lobes of limb, the 
tube narrow, slightly exserted, abruptly expanding into short neck; lobes of upper lip 
oblong-elliptic, obtuse; middle lobe of lower lip oblate, the lateral lobes semiorbicular- 
triangular; upper stamens as long as upper lip of corolla or slightly longer; style more 
exserted; nutlets brown, smooth, oblong. End of flowering and fruit in September. 

Between stones on cliffs in the wood and scrub belt. - Centr. Asia: Pam.-Al. (Vanch 
Range in SE Tadzhikistan). Endemic. Described from area between Motraun and Khei- 
khak. Type in Leningrad. 

Subsection 2. Apodocephalae Pojark. in Addenda XIX, 355. — Semiverticels, with 
possible exception of the lowermost sessile or semisessile; aggregate inflorescence seem- 
ingly an interrupted spike; throat of calyx oblique. 

Series 3. Ladanolentes Pojark. in Addenda XIX, 519. — Semiverticels few-flowered, 
their subtending leaves lanceolate or ovate-lanceolate; bracts lanceolate; calyx with strongly 
obUque throat; corolla-tube not exserted; leaves green on both sides; all parts copiously 
glandular-hairy. 

One species. 

26. N. ladanolens Lipsky in Tr. Bot. sada, XXIIl (1904) 235; 0. and B. Fedch. Perech. 
rast.Turk.V, 144. 

Perennial; stems 60-90 cm long, from decumbent base arcuately ascending, branching 
nearly from base, with internodes 8-12 cm long, densely covered with long fine implexed 
simple hairs and short yellow-headed glandular hairs; lower axillary branches stout, nearly 
as long as main stem, leafy, terminating in inflorescence, the upper short, slender, often 
sterile; leaves grayish-green, covered on both sides with loose appressed gray tomentum 
(this absent on and near the veins and hence leaves apparently rugose) and yellow-glandu- 

339 lar, the lower with petioles about the length of blade, cordate, 1 .5-2.8 cm long, 1 .3-2.5 cm 
wide, crenate, the middle short-petioled or subsessile, smaller; all leaves ovate, with 



226 



orbicular or cuneate base ; lower floral leaves resembling the cauline leaves but smaller, the 
next above narrower to lanceolate, entire, sessile; semiverticels capitate, small, few-flowered, 
borne at the end of stem and axillary branches in 3-7 remote pairs, most sessile, only the 
lower on peduncles 0.7-2 cm long; terminal verticillaster much smaller than the others, 
4-6-flowered; leaves subtending semiverticels 5-8 mm long, 0.5-1 .4 mm wide, ovate-lan- 
ceolate or lanceolate, shorter than semiverticels; bracts lanceolate, usually not exceeding 
the base of upper calyx-teeth, covered (more densely at margin) with long woolly implex 
fine jointed eglandular hairs interspersed with short-stipitate glands; calyx 6-8 mm long, 
densely covered outside with fine short-stipitate glands and spreading hairs; teeth long- 
cihate, the upper narrowly lanceolate, 1/3-2/5 as long as tube, the lower narrower; corolla 
8-10 mm long, pubescent and glandular outside, tube not exserted, expanding into minute 
neck; upper lip 1 .2-1 .5 mm long, cleft into ovate lobes; lower lip nearly 3 times as long 
as upper, its middle lobe not clawed, 2.5 mm long, 4 mm wide, coarsely undulate-crenate, 
the lateral lobes semiorbicular, 0.5 mm long, 1-1.2 mm wide; nutlets 2.2-2.4 mm long, 
0.8-0.9 mm wide, oblong-eUipsoid, brown, with distinct rib on ventral side. End of flow- 
ering and fruit second half of August. (Plate XX, Figure 3.) 

Cliffs in upper part of wood and scrub belt. - Centr. Asia: Pam.-Al. (SE Tadzhikistan). 
Endemic. Described from western slope of Yazgulemskii Range, from ravine of Obi- 
Matraun River. Type in Leningrad. 

Note. The plant has a very strong, pleasant smell of resin. 



Series 4. Olgaenae Pojark. — Semiverticels mostly many-flowered (to 20-40), their 
subtending leaves and bracts lanceolate-linear to fiUform or subulate; calyx with slightly 
obhque throat; corolla-tube distinctly exserted from calyx; leaves white-tomentose 
beneath (rarely on both sides); glandular hairs absent. 

One species. 

27. N. olgae Rgl. in Izv. Obshch. lyub. est. antr. i etn. XXXIV, 2 (1882) 65 ; Lipsk. in 
Tr. Bot. sada, XVIII (1900) 99; 0. and B. Fedch. Perech. rast. Turk. V, 1 14. - Exs.: 
HFAM, No. 205. 

Perennial, sometimes almost suffrutescent, all parts covered with snow-white tomentum; 
root thick, woody, firm; stems many, with woody, perennial branching base to 6 cm 
across, ascending or erect, leafy, often reddish, especially in lower part, white villous- 
tomentose, rarely covered with short and dense tomentum, simple or branching, often 
from base; axillary branches elongate, bearing inflorescence of 2-6 semiverticels, usually 
much shorter than stem; leaves broadly ovate, obtuse, with cordate or rounded base, 
mostly green above, more or less densely covered with appressed hairs, rugose by impres- 
sed network of veins, white-tomentose beneath (sometimes on both sides), the lower 1.3- 
3.5 cm long and wide, with petiole about the length of blade, others with shorter petioles, 
the upper sessile; petioles loosely white villous-tomentose; floral leaves small, 0.5-1 cm 
long, the lower ovate, the others linear, bracteiform; inflorescence consisting of a terminal 
capitate verticillaster (or 2-3 spiciformly crowded) and 4-10 distant 8-40-flowered verti- 
cillasters; semiverticels sessile or subsessile, only 1-2 lower pairs with peduncles 1 cm long; 
leaves subtending semiverticels and bracts lanceolate-Unear to narrowly linear, subfiliform, 
2/5-2/3 as long as calyx and, like the calyx, white-tomentose with loose and long, rarely 
short and densely appressed hairs; calyx 6-8 mm long, tubular, slightly curved, attenuate 



227 



above, with slightly oblique throat; teeth subequal, narrowly triangular, acuminate or 
nearly linear, the 3 upper 1-1}/% times as long as tube; corolla 11-1 2 mm long, densely 
covered outside with nearly tomentose pubescence; tube narrow, curved, exserted to 
1.5-2 mm, abruptly expanding to 1 mm long neck; upper lip ca. 2 mm long, half as long 
as the lower, cleft to the middle into ovate lobes; middle lobe of lower lip with entire or 
undulant margin, 2.3-2.5 mm long, 3-3.3 mm wide; upper stamens not longer than upper 
lip; style distinctly exserted; nutlets oblong-ellipsoid, 2-2.5 mm long, 1 .75 mm wide, 
brownish-black, with sharp rib on ventral side. Fl. secondhalf of May-August; fr. from July. 

Foothills and dry mountains, semidesert belt, stony and gravelly slopes, mainly Hme- 
stone and shale, rarely granite or outcrops of variegated sandstone. — Centr. Asia: T. Sh. 
(Pskem River valley, near Namangan), Pam.-Al. (Mogol-Tau mountains, Turkestan and 
Zeravshan ranges, northern slope of Alai Range, Baba-Tag Range and apparently, after a 
gap, in Darvaz and W. Pamir: Khorog). Endemic. Described from vicinity of Ura-Tyub, 
foothills of Turkestan Range. Type in Leningrad. 

341 Note. The var. mogoltavica Lipsky, segregated by Lipskii and reported to have 
leaves tomentose on both sides and approximate semiverticels, is of no taxonomic signifi- 
cance since such forms also occur in other parts of the distribution area and are Unked 
with the typical form by numerous intermediate forms. 

Section 4. Macronepeta Benth. Lab.gen.etsp.(1834)467,482,p.p. (quoadtyp.). — 
Nepeta § Macronepetae Briq. in Pflanzenfam. IV, 3a (1897) 237, p. p. - Calyx tubular, 
curved , with oblique throat ; corolla medium-sized or large '( 1 . 4-3 . 5 cm long) , with narrow 
curved tube abruptly expanding into wide neck, the narrow part of tube half as long again as 
calyx; middle lobe of lower Up with flat drooping lobules; nutlets smooth. High perennial 
plants, distributed mainly in East and Middle Asia; two species occur in Central Asia. 

Series 1. Sibiricae Pojark. — Corolla large, (17) 20-35 mm long, the dilated part of 
tube 5-9 mm long and as wide, the narrow part twice as long as calyx; flowers in distant 
few-flowered verticillasters; peduncles shorter than flowers; cauline leaves narrow, short- 
petioled, obliquely ascending. 

In addition to the two Russian species, this series contains N. koreana Nakai (N. 
Korea), N. subsessilis Maxim. (Japan), N. przewalskii Pojark. (Hansu province in 
Central China) and N. erecta Benth. (W. Himalayas and Tibet). 

28. N. manchuriensis S. Moore in Journ. of Bot. XVIII (1880) 5; Kom. Fl. Man'chzh. 
Ill, 354; Kom. and Alls. Opred. rast. Dal'nevost. kraya, II, 899. 

■= . Perennial; root long, creeping, 5-7 mm across, woody, coarsely fibrous; rhizome not 
pronounced; stems to 1 m long, to 4-5 mm across, robust, erect, all or only in lower part 
dark purple, subglabrous below with sparse short simple hairs, in upper part densely cover- 
ed with very fine glandular hairs interspersed with sparse spreading longer eglandular hairs, 
mostly simple, rarely with 1-2 pairs of mostly fertile axillary branches; leaves thin (char- 
taceous when dry) dark green above, glabrous or with sparse simple hairs, pale glaucescent 
beneath, densely beset, more densely on veins, with yellow punctate glands and sparse 
jointed eglandular hairs; secondary veins thin but proninent beneath; lower cauline leaves 
ovate, acute, the others lanceolate, to 15-16 cm long, 6-7 cm wide or narrowly lanceolate, 
to 15 cm long, 3.5-4 cm wide, with rounded-truncate or cuneate base, rarely obtuse, usual- 
ly acute or acuminate, often narrowly apiculate, with acutely toothed margin, upper leaves 

342 sometimes entire; petioles of middle leaves 2-3 cm long (1/5-1/3 as long as blade), 3- 

228 



5 mm long in upper leaves; lower floral leaves resembling upper cauline leaves but much 
smaller, the others small, narrowly lanceolate, much shorter than semiverticels, differing 
little from bracts; inflorescence of 4-7 verticillasters; upper semiverticels sessile, their pairs 
sub remote to closely approximate, the lower on peduncles 0.4-3 cm long, 2-6 (9) cm apart; 
bracts narrowly Unear, 4-7 mm long; pedicels to 1.5 mm long; calyx 8.5-9.5 mm long, 
completely covered outside (hke pedicels and bracts) with very small glandular hairs, 15- 
nerved; teeth lanceolate, acuminate, the upper 2/5 times as long as tube, the lower narrow- 
er and longer; corolla 17 -20 mm long, more or less densely covered outside with short 
glandular hairs; narrow part of tube exserted to 1/3, expanding above into wide neck 5- 
7mm long; upper Up erect, 2.5-3.5 mm long, cleft to middle into ovate lobes; middle 
lobe of lower lip reniform, 3-4 mm long, 7-7.5 mm wide, broadly emarginate, with shal- 
lowly and coarsely sinuate-dentate margin, the lateral lobes triangular, 2-2.2 mm long, 
2.8-3 mm wide; upper stamens slightly shorter than upper lip; nutlets 2.5-2.6 mm long, 
1-1.2 mm wide, ellipsoid-obovoid, with a very faint edge on ventral side, brown. Fl. 
June-July; fr. August. 

Pebbles of rivers and streams in mixed forests. — Far East: Uss. (only in southern part). 
Endemic. Described from Maritime Territory, from the Sea of Japan between 44 and 
45°N. Type in London. 

29. N. sibirica L. Sp. pi. (1753) 572 (excl. syn. Buxb.); P. Smirnov in Byull. Mosk. 
obshch. ispyt. prir. Otd. biol. XL VI, 2, 96. - Dracocephalum sibiricum L. Syst. nat. 
ed. X(1759) 1 104; Sp. pi. ed. II, 830 (excl. syn. Buxb.); Bge. in Ldb. Fl. Alt. II 389. - 
Nepeta macrantha Fisch. Catal. hort. Gorenk. ed. 2 (1822) 22, nom. nud.; Benth. Lab. 
gen. et sp.482, diagn.; Benth. in DC.Prodr. XII, 387; Ldb. Fl. Ross. Ill, 1, 378; Kryl. 
Fl. Zap. Sib. IX, 2307; 0. and B. Fedch. Perech. rast. Turk. V, 143. - Moldavica elata 
Moench, Meth. (1794) 410. — M. sibirica Moench ex Steudel Nomenclat. bot. ed. 1 
(1821) 153. -Ic: PaU. Fl. Ross. tab. 1 13; Bot. Mag. tab. 2185. - Exs.: S mi r now, PI. 
alt. exs. No. 71. 

Perennial; 60-100 cm high; root long, creeping, woody, forming some suckers, split- 
ting at the end into coarse fibers; stems 40-100 cm long, slender, to 6 cm across at base, 
obtusely 4-angled, sparsely covered with hairs interspersed with small scablike sessile 
glands, usually densely glandular beneath inflorescence, often reddish in lower part, simple 
or branching all the way up from base; axillary branches long, fertile; leaves lanceolate or 
343 ovate-lanceolate, gradually acuminate above, with cordate or rarely truncate base; cauline 
leaves 5-15 cm long, 1-6 cm wide, evenly serrate or dentate or crenate-dentate, glabrous or 
sparsely covered above with small sessile capitate glands, with numerous punctate yellow 
glands beneath, the lower with petioles 1/7-1/4 as long as blade, the upper with shorter 
petioles; lower leaves subtending inflorescence resembling cauline leaves, the upper Hnear- 
lanceolate, 7-15 mm long, entire; inflorescence consisting of 3-8 remote verticillasters 
forming a loose raceme; lower peduncles to 2 cm, the upper 2-3 mm long; pedicels 1.5- 
2.5 mm long; bracts linear to nearly subulate, slightly longer than pedicels; all parts of 
inflorescence except corolla completely covered with sessile or short-stipitate glands inter- 
mixed with simple villous spreading hairs; calyx 9.5-14 (15) mm long, usually blue, the 
throat strongly obHque, the 3 upper teeth triangular or lanceolate-triangular, 2/7-1/3 as 
long as tube, the 2 lower narrower and longer, 2/3 as long as tube, all acuminate; corolla 
2.5-3.5 (4) cm long, azure-blue, with dark spots on lower lip, sparsely pubescent outside; 
tube exserted nearly to the middle, narrow, gradually and slightly dilated above and then 

229 



abruptly passing into neck, (5.5) 7-9 mm long, (6) 8-10 mm wide; upper lip cleft beyond 
middle into elliptic obtuse lobes; middle lobe of lower lip reniform, 4-7.5 mm long, 6- 

8.5 mm wide, deeply emarginate, with coarsely crenate-incised margin, the lateral lobes 
triangular or ovate -triangular or ovate, 2.5-3.5 mm long, 3.5-5.5 mm wide; upper stamens 
slightly shorter than upper lip; style as long as upper lip; nutlets elongate-obovoid, 2- 

2.6 mm long, 1-1.75 mm wide, grayish-ocherous, with prominent edge on ventral side. 
Fl. July-August; fr. from August. 

A mountain plant growing at altitudes of 500-1200 m along banks of mountain rivers 
and streams, steppe meadows, wastelands, meadow and stony slopes, rarely broadleaved 
forests, usually a weed growing near habitations. - West Siberia: Alt.; East Siberia: Ang.- 
Say., western part (near Kansk, Western Sayans, Tannu-Ola Range); Centr. Asia: Dzhu.- 
Tarb. (Saur and Tarbagatai mountains). Gen. distr.: Dzhu.-Kash. (northeastern part), 
Mong. (Mongolian Altai, Khangai, Gobi Altai). Described from cultivated specimens of 
Siberian origin from the Botanical Garden in Upsala. Type in London. 

Notes. 1) This species is known in the botanical literature under the name N. macran- 
tha Benth. or N. macrantha Fisch. ex Benth. All authors, commencing with Bentham 
and Ledebour, included Dracocephalum sibiricum L. in the synonymy of this name, 
citing the second edition of "Species plantarum" (page 830), although in the first edition 
344 of this work (page 572), Linnaeus described Nepeta sibirica L. and later in the 10th 
edition of "Systema naturae" and in the second edition of "Species plantarum" merely 
transferred his species to the genus Dracocephalum. He also included N. sibirica L. 
in the synonymy of Dracocephalum sibiricum, preserving word for word the diagno- 
sis of N. sibirica and references to the Uterature. Hence the name N. sibirica L., restor- 
ed by Smirnov, should be considered as valid. 

2) N. sibirica occurs as an introduced plant in Lad.-Ilm. (vicinity of Leningrad and 
Tikhvin). Earlier authors (Pallas, Falk and Claus), as well as Ledebour, reported N. ma- 
crantha Benth. for many parts of southeastern Russia where apparently it also occurred 
as introduction and escape. 

Economic importance. The aerial part of the plant yields about 0.15% essential oil 
that may be used in the perfume industry. An ornamental, flowering profusely for a long 
period. It has been known as a cuhivated plant since 1750 and is easily naturalized. 



Series 2. Formosae Pojark. — Corolla medium-sized, 14-18 (20) mm long; narrow 
part of tube exserted to about 1/3, the dilated part 3-5 mm long and wide; flowers in 
remote, more or less many-flowered, compound cymes; most peduncles much longer than 
cymes. Bright green plants, faintly covered with simple and glandular hairs. 

One Central Asian species in this series. 

30. N. formosa Kudr. in Bot. mat. gerb. Inst. bot. i zool. ANUzSSR, IX (1947) 
15. -N. grandiflora auct. fl. turk.: O. and B. Fedch.Perech. rast. Turk. V, 146, non M. B. 

Perennial; root creeping, woody, dense, passing above into branching rhizome; stems 
to 100 cm long, to 8 mm across, with internodes 5-13 cm long, green, covered, densely in 
upper part and sparsely below, with very short simple hairs and small sessile glands, usually 
branching nearly all the way up; lower axillary branches sterile, short, the others 5-40 cm 



230 



long, terminated by inflorescence; leaves thin (chartaceous when dry), bright green, much 
paler beneath; upper side rather densely covered with very small appressed hairs; lower 
side with similar hairs (longer on veins) and punctate yellow glands; lower and middle cau- 
line leaves 4.5-14 cm long, 2.5-8 cm wide, ovate, cordate or nearly triangular-cordate, 
acute or acuminate, with subcordate or rarely slightly notched base, evenly crenate or 
crenate-dentate from base; upper cauline leaves and leaves of axillary shoots commonly 
345 narrower to ovate-lanceolate and frequently with rounded base; petioles puberulent, the 
lower leaves 2/3 as long as blade, the upper much shorter; cymes mostly compound, 
many-flowered, the upper subsessile, crowded at summit of stem and of axillary branches, 
others on peduncles 1-3 cm long in remote pairs; aggregate inflorescence paniculate; lower 
floral leaves resembling cauline leaves, the upper bracteiform; bracts narrowly lanceolate 
to hnear-subulate, 1/3 as long as calyx; pedicels 1-1 .5 mm long; calyx 6.5-9 mm long, 
green or violet, densely covered with very small, sessile and short-stipitate glands and, 
mainly on nerves and margin of teeth, with short spreading simple hairs; teeth triangular, 
rarely lanceolate, tapering to a point, the upper slightly longer than lower, 1/4 -2/5 as long 
as tube; corolla 14-18 (20) mm long, lilac, pubescent and glandular outside, the narrow 
part of tube exserted to 2.5-3 mm; upper lip 3-3.5 mm long, erect, cleft to 1/3-1/2 into 
obtuse lobes; middle lobe of lower lip 3-3.5 mm long, 4.5-5 mm wide, coarsely crenate, 
emarginate, the lateral lobes ovate-triangular or triangular-ovate, 1.75-2.5 mm long and as 
wide; upper stamens with filaments as long as upper Up; style slightly longer; nutlets 
oblong-ob ovoid, 1 .75-2 mm long, 0.75-1 mm wide, dark brown, obtusely 3-angled. Fl. 
first half of June-August; fr. July-September. 

Mountain forests, mainly walnut and walnut-maple, fine earth enriched with humus, 
often in great profusion. - Centr. Asia: T. Sh. (Fergana, Chatkal, Kara-Tau ranges), Pam.- 
Al. (Alai,Zeravshan, Gissar,Vakhsh and Peter the First ranges). Endemic. Described 
from Fergana Range, between Charvak and Arslanbob villages. Type in Leningrad. 

Notes. 1) In habit N. formosa is very much Hke the Caucasian N. grandiflora M.B. 
with which it was identified for many years, but the structure of the middle lobe of upper 
coroUa-Up and the smooth nutlets indicate that there is no close relationship between 
these two species and thatN. formosa is related to the E. Asian representatives of the 
section Macronepeta. 

2) Apparently crossings of N. formosa Kudr. with N. bucharica Lipsky are very 
common. In habit they are intermediate between the two species, at times inclining more 
toward one or the other. Flowers in dense many-flowered compound cymes often resem- 
bling semiverticels or in semiverticels, these all in distant pairs or the upper forming a 
compact spike; corolla violet-bluish, the tube long-exserted; leaves of a thin consistency, 
sometimes recalling the leaves of N. formosa, sometimes more like those of N. bucha- 
rica, the lower surface puberulent or glabrous. 



Series 3. Gontscharovianae Pojark. — Corolla large, 22-27 mm long; corolla-tube 
long, strongly curved, the narrow part exserted to 1/3 or 1/2, the dilated part 4-5 mm long 
and as wide; flowers in remote few-flowered but compact cymes, with short peduncles. 
Plants covered with gray hairs and densely beset with sessile capitate whitish glands. 

Two Central Asian species in the southwestern part of Pamir-Alai. 



231 



3 1 . N.gontscharoviiKudr. inBot. mat. gerb. Inst. bot. i zool. AN UzSSR, IX (1947) 16. 
Perennial; rhizome thick, woody, dark brown, beset with obtuse small scales; stems 

erect or ascending, 30-100 cm long, ca. 3 cm across, robust, with internodes 4-5 cm long, 
densely covered with small short-stipitate capitate glands forming a kind of a powdery 
bloom, in lower part also with loose whitish arachnoid-tomentose hairs, simple or in axils 
of middle leaves with short rudimentary leafy sterile branches and in upper part sometimes 
with 1-2 pairs of short slender leafy branches bearing inflorescence; lower leaves squami- 
form, brown, the others pale green, short-petioled (petioles 1/9-1/6 as long as blade), 
erect, partly parallel to stem, broadly ovate or the uppermost to oblong-ovate, all mth 
deeply cordate or subhastate base, obtuse at apex, crenate or crenate-dentate along the 
whole margin, 3-4.5 cm long, 1 .7-3 cm wide, densely covered above with short glandular 
hairs interspersed with long simple ones, rugose by impressed veins, gray-tomentose beneath, 
the tomentum nearly masking numerous short glandular hairs and round orange sessile 
glands, the veins thin but prominent; lower bracts resembling upper cauline leaves but 
slightly smaller, others much smaller, the uppermost bracteiform, narrowly lanceolate, ca. 
5 mm long; inflorescence of 3-4 pairs of remote, few-flowered (3-8 flowers) but compact 
cymes with peduncles from 3 mm (upper) to 10 mm (lower); pedicels to 2 mm long; bracts 
lanceolate-subulate, 2-4 mm long, much shorter than calyx, densely covered with short 
glandular hairs, at margin with long simple hairs; calyx slightly curved, 8-10 mm long, 
with 15 thick prominent nerves, suffused with violet, very densely covered with small 
short-stipitate glands, the throat strongly oblique, nearly 2-lipped; upper teeth oblong- 
triangular, short-acuminate, 1/3 as long as tube, the lower narrowly lanceolate, long- 
347 acuminate, 2/3 as long as tube, all teeth with coarse, sparse ciUa; corolla 22-27 mm long 
(blue), sparsely hairy outside, tube slender, curved, exserted to half its length, abruptly 
expanding into neck, ca. 4 mm long, 5 mm wide; upper lip 4-5 mm long, cleft into broad 
lobes; lateral lobes of lower hp obHquely triangular-orbicular, 2 mm long, 3 mm wide, the 
middle lobe 3-4 mm long, 5.5-6 mm wide, broadly emarginate; upper stamens slightly 
exceeding upper Up; lower stamens and style slightly shorter than upper lip; nutlets ellip- 
soid, dark brown, 2.5 nmi long, 1 mm wide. Fl. July. (Plate XXI, Figure 1 .) 

Subalpine belt, in meadows. — Centr. Asia: Pam.-Al. Endemic. Described from south- 
ern slope of Gissar Range, from Tuvish River (Tadzhik SSSR, Romitskii region) - thus far 
the only location recorded for this species. Type in Leningrad. 

32. N. consanguinea Pojark. sp. nov. in Addenda XIX, 355. 

Perennial, with thick woody root; rhizome undeveloped; stems erect, 65-100 cm long, 
ca. 5 mm across, strongly branching, densely covered, especially in upper part, with small 
sessile capitate whitish glands and spreading simple white hairs, these usually very dense 
and long in lower part; axillary branches long, fertile, producing in turn short fertile 
branches; several pairs of lower leaves brown, sessile, squamiform, others on petioles 
1/5-2/7 as long as blade, spreading at an acute to nearly right angle, gray beneath with 
dense soft subvelutinous indument of short fme subappressed hairs, on both sides densely 
covered with capitate whitish sessile glands, and fewer round flat yellow glands, triangular- 
ovate, with deeply cordate base, gradually attenuate above, acute, rather coarsely crenate, 
rarely dentate-crenate, the middle largest, ca. 5-7 cm long, 3.5-4.5 cm wide, gradually 
smaller and sometimes narrower toward summit; floral leaves mostly resembling upper 



232 



cauline leaves, the uppermost bracteiform; flowers at ends of stem and axillary branches 
in remote 4-10 (12)-flowered compact cymes disposed in (3) 4-6 pairs and forming a 
narrow sparse raceme; upper cymes on peduncles to 5 mm long, the lower on slender 
straight peduncles to 2-2.5 cm long; leaves subtending cymes lanceolate-linear, 4-6 mm 
long; bracts similar but shorter, 3-4 mm; calyx 9-10 cm long, tubular, faintly curved, 
with 15 prominent nerves, very densely covered with capitate sessile glands, on nerves 
348 with sparse simple spreading thick hairs; throat strongly oblique; teeth triangular, acumi- 
nate, the upper ca. 1/3 as long as tube, the lower much shorter; corolla 19-24 mm long, 
with slender tube exserted from calyx to 1/3-1/2, abruptly expanding into neck ca. 
3.5 mm long, 5 mm wide; upper lip ca. 4 mm long, cleft to middle into broad lobes; 
middle lobe of lower lip 3.5-4 mm long, 6-7 mm wide, the lateral lobes obliquely ovate, 
3 mm long, 3.5 mm wide; upper stamens as long as upper lip; nutlets unknown. 

Stony, fine earth taluses, upper zone of wood and scrub belt. — Centr. Asia: Pam.-Al. 
Endemic. Described from Vakhsh Range, Sufan-Mir-Tau Mountain. Type in Leningrad. 

Note. Very close to the preceding species from which, however, it is readily distin- 
guished by the strongly branching stems, differently shaped patent leaves (not parallel to 
stem), with developed petioles; also by the shorter corolla-tube. 



Section 5. Cataria Benth. Lab. gen. et sp. (1834) 466, 476, p. p. - Nepeta § Steno- 
stegiae Boiss. Fl. or. IV (1879) 638, p.p. — Nepeta § Macronepetae Briq. in Pflanzenfam. 
IV, 3a (1897) 237, p. p. non Benth. — Calyx in flower narrowly tubular, in fruit broader 
to ovate, with more or less oblique throat; corolla small or medium-sized (7.5-10 mm or 
10-22 mm long), included in calyx to dilated part of tube or to limb, rarely the narrow 
part of tube slightly longer than calyx; middle lobe of lower Up markedly concave, the 
involute margin with several large teeth; nutlets more or less distinctly tuberculate or 
smooth; bracts narrow, much shorter than calyx; flowers in cymes or verticillasters form- 
ing racemiform or subpaniculate or spicate inflorescences. Perennials. 

The distribution area of the section stretches from the Himalayas across Central Asia 
and encompasses SW Siberia, Middle Asia, nearly the entire Mediterranean region and the 
most southerly areas of Central and Eastern Europe. 



Subsection 1 . Leiocarpae Pojark. subsect. nov. in Addenda XIX, 356. — Sect. Cataria 
Benth. Lab. gen. et sp. (1834) 466, p.p. (quoad typ.) sensu Briq. in Pflanzenfam. IV, 3a 
(1897) 237. 

Type of subsection: N. cataria L. 

Most species in this subsection occur in W. Himalayas and Afghanistan, very few in the 
Mediterranean region; N. cataria L. is distributed from the Himalayas to Western Europe. 



Series 1 . Catariae Pojark. — Calyx not strongly curved, ovoid in fruit, its upper teeth 
not differing in size from the rest; corolla small, to 10mm long; flowers in many-flowered 
dense cymes at ends of stem and axillary branches forming oblong narrow racemes, these 
dense in upper part, often nearly spicate. 



233 



349 Among members of this series, N. cataria L. has apparently only one close relation — 
N. atlantica J. Ball. (Morocco). 

33. N. cataria L. Sp. pi. (1753) 570; M. B. Fl. taur.-cauc. II (1808) 39; Benth. Lab. 
gen. et sp. 477; Ldb. Fl. Ross. Ill, 374; Boiss. Fl. or. IV, 643; Shmal'g. Fl. II, 312; 
Grossg. Fl. Kav. Ill, 293; idem,Opred. rast. Kavk. 331; Kryl. Fl. Zap. Sib. IX, 2309. - 
N. minor Mill. Diet. ed. VIII (1768) No. 2. - Cataria tomentosa Gilib. Fl. Lith.(1781) 
78. - C. vulgaris Moench, Meth. pi. (1794) 387. - N. vulgaris Lam. Fl. Fr. II (1778) 
398. - N. macrura Fisch. ex Spreng. Syst. veg. II (1825) 729; Benth. in DC. Prodr. XII, 
395.- Glechoma cataria 0. Ktze. Rev. gen. (1891) 518. - Ic: Rchb. Ic. Fl. germ. 
XVIII, tab. 1242; Fedch. and Fler. Fl. Evrop. Ross. (1910) 807; Syreishch. Fl. Mosk. 
gub.III,87; Fl.Yugo-vost. VI, Figure 604. -Exs.: GRF,No.27; Fl. pol. exs. No.668; 
Sintenis, It. trans.-casp. an. 1900-1901, No.668 and No. 812. 

Perennial; root woody, branching; rhizome not evident; stems 40-100 cm long, robust, 
erect or ascending at base, covered with short soft pubescence of simple white reclinate 
hairs, branching; branches leafy, fertile, mostly short; leaves triangular-ovate, with cor- 
date base, acute, large-toothed, covered on both sides with short hairs, much more densely 
so and hence grayish beneath, later sometimes glabrescent; cauline leaves 2-8 cm long, 
1.2-4 (5) cm wide, all petiolate; petioles pubescent, the lower 2/3 as long as to slightly 
shorter than blade, the upper much shorter; lowermost (1-2 pairs) floral leaves resembling 
upper cauline leaves, others much shorter, entire, lanceolate, the upper small, bracteiform; 
flowers in dense compound cymes at ends of stem and branches forming a raceme; upper 
cymes few-flowered, dense, subsessile, generally crowded, the lower on peduncles to 1.5- 
2 cm long, more or less remote, many-flowered, looser; bracts linear-subulate, shorter than 
calyx; pedicels 1-1 .5 mm long; calyx 5.5-7.5 mm long, slightly curved, densely grayish- 
puberulent, green or in upper part more or less violet, in flower narrowly tubular, in fruit 
ovoid, with 15 ribbed nerves; throat oblique; teeth lanceolate at base, abruptly tapering 
from middle to long subulate point, the upper (1/2) 2/3 as long as to rarely equaling tube, 
the lower about equaling the upper but longer in relation to tube; corolla 7.5-10 mm long, 
dingy white, with purple or violet spots on lower hp, rarely without them (f. immaculata 
Syr.); tube 3-4 mm long, slightly curved, not exserted from calyx, rather gently expanding 
above into neck 2-3 mm long, 2.5-3 mm wide; upper lip ca. 2 mm long, cleft to middle 

350 into ovate lobes; lower lip twice as long as the upper, the middle lobe oblate, large-toothed, 
2.5-3.5 mm long, 4-4.5 mm wide, involute-margined, the lateral lobes semiorbicular, 0.7- 
1 mm long, 1.3-1.5 mm wide; nutlets brown, ellipsoid, 1.3-1.5 mm long, 0.8-1 mm wide, 
smooth. Fl. last half of June-July; fr. from July. 

Meadows, forest margins, forests, thickets, riverbanks, mainly weed-infested places, fields, 
orchards, gardens, ditches, meadow slopes of mountains in central mountain and subalpine 
belts. - European part: Bait., Lad.-Ilm., U. V., V.-Kama, U. Dnp., M. Dnp., V.-Don, 
Transv., U. Dns., Bes., Bl., Crim., L. Don, L. V.; Caucasus: all regions; West Siberia: U. 
Tob.,Irt.; Far East: (Uss.) sometimes introduced; Centr. Asia: Ar.-Casp. (northern part 
- Mangyshlak, Ust Urt and outlier mountains), Balkh. (mountains), Dzu.-Tarb., Mtn. 
Turkm. (Kopet-Dagh), Pam.-Al., T. Sh. Gen. distr.: nearly all Europe but, except for 
east and south, only escaped or introduced; Med., Bal.-As. Min., Arm. -Kurd., Ind.-Him., 
Jap. (introduced), N. Am. (introduced). Described from Europe. Type in London. 



234 



(351) 




PLATE XXI. 1 - N. gontscharovii Kudr., general aspect, flower; 2 - N. longituba Pojark., general 
aspect, flower, nutlet; 3 - N. velutina Pojark., lower and upper parts of inflorescence, leaf, flower, 
nutlet. 



235 



Note. Gynodioecism and gynomonoecism are observed at times in N. cataria. The 
female flowers are smaller than the bisexual and have four staminodes hidden in the dilated 
part of the tube. 

Economic importance. The plant contains up to 3% essential oil that is used in the 
perfume industry; a form with a strong lemon odor, f. citriodora Dam., has been culti- 
vated for years as a nectariferous plant. In Tashkent conditions (Tsukervanik, 1934) the 
aerial parts of this form yielded 0.15% essential oil which consisted of 13% citral, 8% 
limonene and dipentene, and about 80% alcohol. The alcohol fraction contained about 
17% geraniol, 25% citronellol and about 38% innerol [?]. Decoctions of N. cataria are 
used in popular medicine in the treatment of colds, catarrhal gastritis, anemia and other 
ailments. There are indications that the roots of N. cataria contain substances that have 
a strong stimulating effect on the nervous system. 



Subsection 2. Tuberculatae Pojark. subsect. nov. in Addenda XIX, 356.- Sect. 
Cataria Benth. Lab. gen. et sp. (1834) 466, pro min. parte. — Sect. Macronepetae Benth. 
1. c. 467, pro min. parte. — Nutlets tuberculate, sometimes very small. 
553 Type of subsection: N. grandiflora M. B. 

The habitats of the overwhelming number of species in the section lie in the countries 
of Southwest Asia and the Caucasus; several species occur in S. Europe. 

Cycle 1 . Heterodontae Pojark. — Calyx strongly curved, with strongly oblique near- 
ly 2-lipped throat, oblong or tubular-ovoid in fruit, the uppermost tooth distinctly longer 
than adjoining teeth; corolla 10-15 mm long; nutlets with very small and very flat, pellu- 
cid tubercles; flowers in compound cymes forming a racemiform or paniculate, more or 
less pyramidal inflorescence. 

The closest relations of the Caucasian species in this cycle are the Western Mediterra- 
nean N. nepetella L.s.l. (S. Europe), N. amethystina Desf. (SE Spain and N. .-^"Urci- 
ca Guir. (Spain —Granada) which consist, however, of the phyletic series Nepetellae Pojark. 
that, in some characters, occupies a position between the cycle Heterodontae and the 
series Cataria and thus links the two subsections of the section Cataria in the range adopt- 
ed here. N. nepetella L. has been reported for the south of the European part of the 
Soviet Union (partly under the synonymous name N. graveolens Vill.) but, if these 
reports are true, they can only refer to introduced plants. 

34. N.cyaneaStev. inMem.Soc.Nat. Mosc. 111(1812) 265; Benth. Lab. gen. et sp. 
478, p. p.; inDC.Prodr. XII,383,p.p.; Ldb. Fl. Ross. Ill, 375, p.p.; N. Pop. in Tr. Bot. 
sada Yur'evsk. univ. XIV, 234, p. p.; Grossg. Opred. rast. Kavk. 332, p. p. - N. incana 
var. cyanea M.B. Fl. taur.-cauc. Ill (1819) 392.- ? N. mollissima Tausch in Flora, I 
(1831)219. -N. cyanea var. steveniana Trautv. in Tr. Bot. sada, V (1877) 469; Lipsk. 
Fl. Kavk. 421; Grossg. Fl. Kavk. Ill, 295; Sosnovsk. Fl. Gruz. VII, 297. 

Perennial, grayish or green, with dark brown woody fibrous root; stems 20-50 cm long, 
2-4 mm across in lower part, densely covered in lower part with short appressed hairs, in 
upper part usually with longer spreading hairs, often downy-villous under inflorescence, 
branching or rarely nearly simple; middle internodes to 6-8 mm long; branches simple 



236 



(rarely with rudimentary secondary branches in axils of leaves), upright, often all short, 
the lower sterile, the upper terminating in inflorescence, rarely most of them long and 
fertile; leaves green, mostly not rugose, rarely rugulose, usually rather densely covered on 
both sides with coarse hairs, these appressed or subpatent beneath, interspersed, densely 
beneath and sparsely above, with punctate yellow glands; cauHne leaves broadly ovate to 
oblong or lanceolate, 2-4.5 cm long, 0.8-2.5 cm wide, with truncate or cuneate base, ob- 

354 tuse or acute, the margin with distinct acute or obtuse teeth; petioles of lower leaves 1/3- 
1/2 as long as blade, those of upper leaves 1/8-1/6 as long; lower floral leaves resembUng 
the upper cauline; upper floral leaves long-acuminate, entire, sessile; leaves subtending 
cymes linear-subulate ; flowers at ends of stems and axillary branches forming a narrowly 
pyramidal or cyUndrical panicle; upper part of panicle very dense, nearly spicate, consist- 
ing of several pairs of crowded semiverticels or cymes, the lower part composed of 2-4 
pairs (on stem) or 1-2 (on branches) of remote cymes with peduncles 0.6-2 cm long; 
cymes compound, dense, outwardly resembling semiverticels; axes and pedicels short; 
bracts linear or subfiliform, 2-3 mm long; calyx 9-12 mm long, downy- or tomentose- 
villous with spreading blue (or white in lower part) long jointed (6-10 cells) hairs and 
beset with short-stipitate and sessile whitish glands, very strongly curved, the uppermost 
tooth narrowly lanceolate, long-acuminate, 2.8-4.2 mm long, reaching the base of upper 
lip of corolla, 2/5-1/2 as long as tube, the middle and lower teeth abruptly attenuate from 
broad base to porrect point, the middle 0.3-0.7 mm shorter than the uppermost, the two 
lower 2/3 as long; corolla (12.5) 13-15 mm long, included in calyx nearly to the limb, the 
neck 2.3-2.5 mm long, 3.5-4 mm wide; upper lip 3-3.5 mm long, cleft to middle or 2/3 
into distally enlarged obtuse lobes 2.1-2.4 mm wide; lower Up with middle lobe 3-4 mm 
long, 5-6 mm wide, the lateral lobes semiorbicular-triangular, 1.8-2 mm long, 2.5 mm 
wide; upper stamens 1-1 .5 mm shorter than upper lip of corolla; nutlets ellipsoid, 2.2- 
2.4 mm long, 1.1-1.2 mm wide, brownish-black in maturity, covered on both sides with 
rather large, flat, round, irregularly papilliform tubercles. Fl. June-first half of September; 
fr. from August. (Plate XXII, Figure 1.) 

Central and subalpine, sometimes alpine, belts, 1250-3300 (4200) m, bluffs and taluses. 
— Caucasus: Main Range (eastern part). Dag. Endemic. Described from S. Dagestan, near 
Khinalug village. Type in Helsinki, topotypes in Leningrad. 

Note. The taxonomic significance of N. ruprechtiana Boiss.(Fl. or. IV, 1879, 661), 
described from N. Dagestan (between Golotl' and Avarskoe Koisu River at an altitude of 
about 1 50 m), remains obscure. The authentic specimen of this species has many features 
in common with N. cyanea Stev. (shape and dentation of leaves, character of vesture, 
shape of calyx and its teeth, dense semiverticel-like cymes, sculpture of nutlets). The most 
significant feature distinguishing this species from N. cyanea is the corolla included in 
the calyx only to the base of the dilated part of the tube (not up to the limb). It must 
also be noted that N. ruprechtiana was collected in the lower mountain belt where 

355 N. cyanea does not occur. There are specimens near Gunib, at altitudes of 1200-1500 m, 
that resemble N. ruprechtiana in essential features. More material and investigations of 
this species in nature are required. 

35. N. kubanica Pojark. in Addenda XIX, 356. - N. incana C. A. M. Verzeichn. Pfl. 
Cauc. kasp. Meer. (1831)92, p.p. non Sol. nee M. B. - N. cyanea auct. fl. cauc. non 



237 



Stev. — N. cyanea var. steveniana Lipsky in Sched. herb. Fl. Ross. No. 934, non Trautv. 
-Exs.: GRF,No.934. 

Perennial; stems 30-55 cm long, robust, erect, densely covered all over with very short 
appressed crisp hairs, purple in lower part, 2.5-4.5 mm thick, branching from base; inter- 
nodes 15-17, the lower and those in inflorescence short, others 4-8 cm long; lower branches 
often short, sterile, the upper bearing inflorescence of 3-5 pairs of cymes, these subequal, 
usually much shorter than stem; leaves yellowish-green, concolor, covered with very short 
and fme (magnifying glass!) appressed hairs and beset, very densely beneath, more sparsely 
above, with punctate glands, the thick nerves prominent beneath; cauline leaves 2-5.5 cm 
long, 0.5-1.2 cm wide, oblong, lanceolate or narrowly lanceolate, cuneate at base, gradual- 
ly tapering to acute apex, the margin with acute teeth; upper cauhne leaves and leaves of 
axillary branches often entire; petioles ca. 2/9 as long as blade, the upper shorter; lower- 
most floral leaves resembling the cauline, others small, ovate, entire, the uppermost and 
those subtending cymes bracteiform; cymes 5-13-flowered, 5-9 pairs at the end of stem 
and axillary branches forming lax cylindrical or sometimes subpyramidal panicles, the up- 
per peduncles 3-7 mm long, the lower 2.5-3 cm long; most cymes twice loosely dichoto- 
mously branching, the axes of first order sometimes forming a short 3-5-flowered bostryx; 
pedicels 0.3-0.8 mm long, only in bifurcation of cyme 2-2.5 mm long; bracts linear, 
1 .5-2.5 mm long; calyx 6-7.5 (8) mm long, more or less blue-tinged, puberulent and beset 
with scattered whitish and yellow sessile glands; uppermost tooth narrowly lanceolate, 
1 .5-2 (2.5) mm long, usually barely reaching the base of neck of corolla-tube or even 
shorter, the middle and lower teeth abruptly attenuate from broad base to a porrect point, 
the middle slightly shorter than the uppermost, the lower 2/3 as long; corolla 1 1-12 mm 
long, in dry condition with bluish spots on upper hp and on lateral lobes of lower lip, 
356 the middle lobe purple-blue or blue, the narrow part of tube often exserted to 0.2-1 mm, 
the neck ca. 2 mm long, 3 mm wide; upper hp 2-2.5 mm long, cleft to middle or beyond 
into broad, sometimes nearly semiorbicular lobes; middle lobe of lower lip 2.5-4 mm 
long, 4-6 mm wide, the lateral lobes broadly triangular, obtuse, 1 .2-1 .8 mm long, 2-2.5 
2.5 mm wide; upper stamens as long as upper hp; nutlets ellipsoid, with an obscure edge 
on convex ventral side, blackish-brown in maturity, covered with small flat tubercles, 2- 
2.2 mm long, 1-1.2 mm wide. Fl. and fr. July-August. 

Central mountain belt, 1 200-2000 m, exposed stony slopes, taluses, limestone outcrops, 
thorny thickets and pine forests. — Caucasus: W. Transc. and western part of Main Range. 
Endemic. Described from near Madniskhevi village at upper reaches of Kuban' River. 
Type in Leningrad. 

Note. N. kubanica differs outwardly from N. cyanea Stev., with which it used to 
be confused, by its long, virgate axillary branches, narrower leaves, different indument, 
smaller flowers, and the comparatively lax, few-flowered, remote cymes. 

36. N. czegemensis Pojark. sp. nov. in Addenda XIX, 357. 

Perennial, with coarsely fibrous woody root; stems 30-50 cm long, 2-3 mm thick, as- 
cending at base, often more or less curved, densely grayish-pubescent with very short ap- 
pressed rechnate hairs, branching mostly from base, the middle internodes 4-7 cm long; 
branches varying in length without apparent pattern, all fertile or the lower sterile; leaves 
thin, grayish, densely covered with very short (magnifying glass!) appressed hairs, with 



238 



punctate yellow glands (more copious beneath), smooth, the thin veins not prominent 
beneath; cauline leaves 1.8-3 cm long, 0.9-1.5 cm wide, oblong-ovate to lanceolate, with 
truncate or rarely cuneate base, obtuse or acuminate at apex, each margin with 5-10 in- 
conspicuous teeth, these acute or obtuse, sometimes distant; petioles 1/4-1/3 as long as 
blade; lower floral leaves resembUng cauline leaves or broadly ovate, equaling or exceed- 
ing peduncles, abruptly decreasing in size and bracteiform above; leaves subtending cymes 
and bracts linear or lanceolate-linear, 2-3 mm long; cymes on stem 5-8 (12) pairs, on lat- 
eral branches fewer (2-6), the upper 2-6-flowered on peduncles 5-8 mm long, loosely 

357 approximate, the lower 2-4 cm distant, many -flowered, dichotomous or trichotomous, on 
peduncles to 4 cm long; aggregate inflorescence a loose oblong panicle; calyx (6.5) 7- 
8.5 mm long, densely villous with long blue, rarely white implexed hairs, oblong-ovoid in 
fruit, the median tooth slightly larger than lateral teeth, 1.5-2.5 mm long, lanceolate, acu- 
minate, 1/4-1/3 (2/5) as long as tube, lateral and lower teeth subequal, with broad base, 
abruptly attenuate to a long point; corolla 1 1-1 2 mm long, with whitish tube included in 
calyx to base of dilated part (neck); upper lip 2-3 mm long, cleft to middle into obtuse 
lobes 1 .7-2 mm wide; lateral lobes of lower Up obliquely semiorbicular, 1 .2-1 .5 mm long, 
1.8-2 mm wide, like upper lip purple-spotted, the middle lobe in dry condition blue, 4.5- 

5 mm long, 6-7 mm wide; nutlets brown, 2-2.3 mm long, 0.8-1 mm wide, oblong-eUipsoid, 
3-angled, rounded above, pointed at base, dull, finely punctate with rather sparse flat tu- 
bercles. Fl. July-August; fr. from second half of August. 

Central mountain belt 1700-2500 m, mountain-steppe plant communities, pine forests, 
stony taluses. — Caucasus: Main Range, upper part of western tributaries of Terek River 
(Baksan, Chegem and others). Endemic. Described from Chegem River north of Bulungu 
village. Type in Leningrad. 

37. N.biebersteiniana(Trautv.) Pojark. sp. nov. - N. incana M. B. Fl. taur.-cauc. II 
(1808) 60, non Soland in Ait. - N. incana var. acinifolia M. B. Fl. taur.-cauc. Ill (1819) 
392 (non N. acknifolia Spreng.). — N. incana var. cyanea C. A. M. Verzeichn. Pfl. 
Cauc. kasp. Meer. (1831) 92, p.p. non Sol. necM. B. — N. cyanea auct.: Benth. Lab. 
gen. et sp. (1834) 478, p. p.; in DC. Prodr. XII, 384, p. p.; Ldb. Fl. Ross. Ill, 375, p. p.; 
Boiss. Fl. or. IV, 656, p. p.; N. Pop. in Tr. Bot. sada Yur'evsk. univ. XIV, 234, p. p.; 
Grossg. Opred. rast. Kavk. 332, p. p. non Stev. — N. cyanea var. biebersteiniana Trautv. 
inTr. Bot. sada, V (1877) 470; Lipsk. Fl. Kavk. 421; Grossg. Fl. Kavk. Ill, 295; Sos- 
novsk. Fl. Gruz. VII, 297. - Ic: Rchb. Ic. bot. tab. CCCV, fig. 478. 

Perennial, strongly branching, grayish or whitish all over with dense short hairs; root 
woody, fibrous, more or less twisted, to 2 cm thick, branching; stems 20-50 cm long, 
1 .5-3 mm thick, densely covered with very short crisp appressed hairs (sometimes sUghtly 
spreading in upper part), with numerous (to 20) internodes 1.3-3.3 (4) cm long, branch- 
ing from base; axillary branches leafy; all branches fertile, more or less branching, arcu- 
ately curved, the lower long, often as long as stem, others gradually shorter toward stem 

358 summit, hence the ends of axillary branches apparently equidistant from stem; leaves 
covered on both sides with very short yellowish or grayish hairs, often nearly tomentose 
and on both sides, more densely beneath, with yellow punctate glands, usually rugose with 
veins impressed above and prominent beneath; cauline leaves 9-30 (40) mm long, 6-25 
(35) mm wide, oblong, ovate-oblong or lanceolate, obtuse or acuminate, with cuneate or 



239 



rarely truncate base, the margin with small appressed teeth; lower petioles 2/5-2/3 and of 
upper about 2/7 as long as blade; leaves on axillary branches smaller and often narrower; 
lower floral leaves resembUng cauline leaves, others smaller, acuminate, entire, the upper 
(also leaves subtending cymes) bracteiform; all floral leaves, except the uppermost, longer 
than peduncles; flowers in 3-9-flowered cymes at ends of stem and axillary branches form- 
ing sparse racemiform or paniculate inflorescences; peduncles from 0.3-1.5 cm (the upper) 
to 1 .5-2.5 cm long; most cymes dichotomous or trichotomous, the axes of the first order 
mostly forming a short bostryx; pedicels 0.3-1.3 mm long; bracts ovate -lanceolate or 
lanceolate, 1 .5-2.5 mm long, acuminate; calyx 8-1 1 mm long, densely and uninterrupted- 
ly covered with whitish or bluish appressed antrorse hairs, rarely puberulent, beset with 
sessile or short -stipitate whitish glands; teeth tapering to a long nonspinescent point, the 
uppermost 2-3 mm long, 1/2-2/3 as long as tube, the two lower shortest, 0.8-1 (2) mm 
long; corolla 10-13 mm long, whitish with violet-reddish lower lip, puberulent, included 
in calyx nearly to the Hmb; tube abruptly expanding into neck 2.2-2.7 mm long, 2.7- 
3.5 mm wide; upper Hp 2.8-3 mm long, cleft to middle into obtuse distally enlarged lobes; 
middle lobe of lower lip 2.5-3.5 mm long, 3.5-5.5 mm wide, the lateral lobes broadly tri- 
angular, 0.6-1 .2 mm long, 1 .8-2.7 mm wide, all with dark purple spots; nutlets elUpsoid- 
obovoid, brown, 2.2-2.5 mm long, 1-1.2 mm wide, convex on ventral side but without rib, 
obscurely tuberculate, the flat tubercles minute, almost punctate. Fl. June-July; fr. sec- 
ond half of June to August. 

Middle and lower mountain zone, 1 50-2000 m, exposed gravelly and stony slopes, and 
mountain-steppe plant communities. — Caucasus: Cisc. (vicinity of Stavropol', basins of 
Kuma, Terek and Ardon rivers). Dag. Endemic. Described from S. Dagestan, Akhta vil- 
lage. Type in Leningrad. 

Note. N. biebersteiniana has so far been considered a variety of N. cyanea. There 
are, however, good reasons for classifying it as a species. Morphologically, it is readily dis- 
359 tinguishable from N. cyanea by a number of characters (habit, character of branching, 
shape of leaves, indument, shape of inflorescence, color of corolla, sculptural details of 
nutlets). The occurrence of specimens from Dagestan (apparently of hybrid origin) that 
gravitate toward N. cyanea, mainly as regards indument, does not alter the fact. More- 
over, the distribution areas of the two species overlap to some extent only in Dagestan, 
and this incompletely since N. biebersteiniana is associated with the lower mountain 
zone, ascending to the middle zone, whereas N. cyanea occurs mainly in the high-moun- 
tain zone and does not descend below the middle zone. 



Series 2. Grandiflorae Pojark. — Calyx curved, with distinctly but moderately obUque 
throat; lower teeth slightly smaller than the upper; corolla (14) 15-1 8 mm long, included 
in calyx to the base of dilated part of tube; nutlets inconspicuously and unevenly foveo- 
late-tuberculate, the tubercles small and flat; flowers in many-flowered cymes forming 
loose racemes at the ends of stem and axillary branches. High, strongly branching plants, 
with cauHne leaves more or less horizontally spreading. 

Monotypic series. 

38. N. grandiflora M. B. Fl. taur.-cauc. II (1808) 42; Ldb. Fl. Ross. Ill, 376; Boiss. Fl. 
or. IV, 659; Lipskii, Fl. Kavk. 421; N. Pop. in Tr. Bot. sada Yur'evsk. univ. XIV, 234; 

240 



Grossg. Fl. Kavk. Ill, 295; idem,Opred. rast. Kavk. 331. - N. grandiflora var. glabrata 
N. Kusn. in Uch. zap. Yur'evsk. univ. 2 (1899)46, p.p. — N. melissaefolia Pers. Synops. 
2(1807) 115,nonLam. - N. colorata Willd. Enum. pi. II (1809) 601. - Ic: Fl. Gruz. 
Ill, Fig. 320; Hegi, III. Fl. V, 4, fig. 3249, a-b. - Exs.: GRF, No. 1083. 

A high perennial; root oblique, slender, woody, tough, without tuberiform swellings; 
stems few or solitary, 50-1 50 cm long, (1.5) 3-6.5 mm thick, erect, strongly 3-angled, 
rather densely puberulent below inflorescence, subglabrous or with few hairs below, 
branching from base or only in upper part, rarely simple; lower axillary branches short, 
sterile, the others fertile, (3) 5-35 cm long, some branching; leaves thin, the cauHne to 
8 cm long, 4.5 cm wide, ovate or oblong-ovate to sublanceolate, mostly with cordate base, 
acute, crenate-dentate, dark green above, rather sparsely covered with short hairs, glauces- 
cent beneath, more densely puberulent; lower petioles to 15-20 mm long, the upper 2- 
3 mm long; cymes mostly compound, of fewer flowers at ends of stem and upper axillary 
branches, in loose elongate racemes forming a paniculate aggregate inflorescence; pedun- 
360 cles 2-3 (5) cm long, the upper approximate; lower floral leaves resembling the cauline; 
upper floral leaves and leaves subtending cymes lanceolate-linear, bracteiform; bracts 
linear or nearly subulate, 1/4-1/3 as long as calyx, bluish-violet, ciliolate, covered with 
small glands; calyx (8.5) 9.5-10.5 mm long, violet-blue, densely covered with very fine 
capitate glands, with very short spreading or more or less appressed simple hairs along 
nerves; teeth narrowly triangular or triangular, acuminate or acute, 1/3-1/2 as long as 
tube; corolla (14) 16-18 mm long, violet-blue, pubescent, the narrow part of tube not 
exserted, the neck elongate; upper lip 2.5-3.5 mm long, cleft to 1/3-1/2, the lobes obtuse, 
distally enlarged; middle lobe of lower lip 4-7 mm long, 7-8.5 mm wide, coarsely toothed, 
the lateral lobes broadly triangular, 2.5-3 mm long, 1 .3-1 .5 mm wide; upper stamens and 
style slightly longer than upper hp; nutlets broadly eUipsoid, 3-angled, 1 .75-2 mm long, 
1-1 .25 mm wide, dark brown, with sparse irregular flat tubercles. Fl. last half of June to 
August; fr. from first half of July. 

Subalpine meadows and tall-grass stands, often very abundant; rarely wood margins 
and glades in the forest belt, meadow-steppe slopes. — Caucasus: Cisc. (W. and E.), Dag., 
W., E. and S. Transc. Endemic. Occurring as an escape in many parts of the RSFSR and 
in the Ukraine, also Centr. Europe. Described from the Caucasus. Type in Leningrad. 

Note. N. zangezura Grossh. (in Beihefte Bot. Centralbl. XLIV, 2, 1928, 234), which 
was described from Migry, at the southern end of the Zangezur Range in Armenia, is very 
closely allied to N. grandiflora. A comparison of one of the original herbarium speci- 
mens with a plant collected in Migry and cultivated in the Armenian botanical garden 
(Erevan) revealed that, morphologically, N. zangezura can just be distinguished from 
N. grandiflora by the following characters: smaller calyx covered with shorter, coarser 
hairs, the broadly triangular teeth 1/5-1/4 as long as the tube (similar shape and size ratio 
may be occasionally observed in N. grandiflora); racemes more contracted. A reported 
difference in color of corolla (pale blue) has not been confirmed: in nature the Migry 
plant has a violet-blue corolla. It should be borne in mind that N. zangezura occurs 
under completely different ecological conditions — open xerophilous forests of Quercus 
araxina in the lower mountain belt — and also that its distribution area is apparently dis- 
junct from that of N. grandiflora. It is therefore important to determine the taxonomic 
position of N. zangezura. There is need for more material and study in natural conditions. 



241 



Economic importance. Sometimes cultivated as an ornamental; easily naturalized. The 
essential oil of tliis plant is apparently of no value. 



361 Series 3. Betonicifoliae Pojark. — Calyx slightly curved, with slightly oblique throat 
and subequal teetli; corolla 15-23 mm long, the narrow part of tube exceeding calyx by 
1.5-2.5 mm; nutlets with sparse flat tubercles; verticillasters dense, crowded at summit 
of stem in a dense spicate inflorescence (the lower 1 or 2 sometimes distant); stems few, 
simple; leaves short-petioled, upright. 

This series consists of the three species described below. 

39. N. betonicifolia C. A. M. Verzeichn. Pfl. Cauc. kasp. Meer. (1831) 92; Ldb. Fl. 
Ross. Ill, 374; Boiss. Fl. or. IV, 658; Lipsk. Fl. Kavk. 421 ; Grossg. Fl. Kavk. Ill, 294, 
p. p.; idem, Opred. rast. Kavk. 332, p. p. 

Perennial, gray-pubescent; rhizome creeping, slender, woody, tough, with internodes 
ca. 1 cm long, at nodes bearing small dark brown squamiform leaves; stems 25-40 cm 
lonj,, .„j^.xily ascending, 3-5 mm thick, simple, densely leafy, sharply 4-angled, thickened 
along ribs, densely covered all over with fine crisp hairs, almost tomentose in upper part; 
leaves upright, firm, gray-pubescent on both sides, with fine flexuous 3-5-jointed hairs, 
densely beset beneath with very small round brownish glands; cauline leaves to 3.5 cm 
long, 2 cm wide, the lower sometimes exceeding internodes, mostly oblong-ovate (the 
lower sometimes ovate), with cordate base and obtuse apex, the upper narrower, ovate- 
lanceolate, acute; all leaves evenly toothed from base, rarely crenate, with 14-17 (20) 
teeth on each margin, the lower leaves with petioles 5-12 cm long, others with petioles 
2-3 mm long or sessile; verticillasters dense, crowded at summit of stems in a firm ovoid 
or slightly elongate cylindrical spicate inflorescence 3.5-6 cm long and 2.5-4 cm wide, 
1 lower verticillaster sometimes remote; semiverticels sessile or sometimes 1 or 2 lower 
pairs on short peduncles 2.5-3.5 mm long; lower floral leaves lanceolate, sessile, with cu- 
neate base, acutely toothed to nearly entire, 12-22 mm long, 5-1 1 mm wide, the upper 
bracteiform, linear, 10-12 mm long, 1-1.5 mm wide; bracts narrowly Unear, the outer 
5-8 mm long, the inner 3-4 mm long, 2/5-1/2 as long as calyx, long-hairy, sometimes 
violet; calyx 8.5-10 mm long, slightly curved, with scarcely oblique throat, 15-nerved, 
densely arachnoid-tomentose, with very fine implexed long 4-6-cellular white hairs, violet 
in upper part; teeth lanceolate or oblong-triangular from middle, rarely long-acuminate, 

362 subequal, 1/3-2/5 as long as tube; corolla 18-20 mm long, densely white-pubescent out- 
side, the slender curved tube distinctly (2-2.5 mm) exserted from calyx, abruptly expand- 
ing into neck 3-3.5 mm long, 4 mm wide; upper lip ca. 3 mm long, cleft to the middle 
into obtuse lobes; lower lip twice as long as the upper, the middle lobe 3.5 mm long, 

5 mm wide, the lateral lobes broadly triangular, asymmetrical, 1 .25-1.5 mm long, 2.5- 
3 mm wide; upper stamens with filaments exceeding upper lip of corolla by 0.5-1 mm, 
filaments of lower stamens equaling it; style slightly longer than upper lip of corolla; 
nutlets (immature) oblong-obovoid, with sparse flat tubercles. Fl. and fr. July. 

Subalpine meadows. — Caucasus: Tal. Gen. distr.: likely to occur in the part of 
N. Iran bordering on Talysh. Described from Talysh mountains. Type in Leningrad. 



242 



40. N. somkhetica Kapell. in Vestn. Tifl. bot. sada, nov. ser. 5 (1931) 35; Grossg. Fl. 
Kavk. Ill, 295; idem, Op red. rast. Kavk. 331. — N. grandiflora var. glabrata N. Kusn. 
in Uch. zap. Yur'evsk. univ. 2 (1899) 46, p. p. - N. grandiflora f. simplex Lipsky in 
Gerb. russk. fl. (1902) No. 1082. - N. grandiflora auct.: N. Pop. in Tr. Bot. sada 
Yur'evsk. univ. XIV (1914) 234, p. p. non M. B. - Exs.: GRF, No. 1082. 

Perennial; root woody, coarsely fibrous, with numerous distant, often fusiform swell- 
ings; rhizome short, horizontal; stems erect or ascending, 35-100 cm long, 2-3.5 mm 
thick, simple, with 8-10 internodes (not counting inflorescence), more or less densely 
covered with very short simple 1-3-jointed hairs, these slightly longer at nodes and on 
lower internodes; upper internodes (4) 7-10 cm long; leaves thin, not rugose, upright, 
the upper parallel to stem, oblong-ovate, with cordate base (the upper often with nearly 
truncate base, narrower to lanceolate), acute, rarely obtuse, with 1 5-17 large obtuse or 
acute teeth on each margin, dark or bright green above, paler beneath, sparsely covered 
on both sides with short simple hairs, these appressed above, more or less spreading on 
lower side, especially along veins, interspersed with punctate yellow glands; largest leaves 
at middle part of stem 4-9 cm long, 1 .5-4 cm wide ; lower petioles 1 /3-1 /2 as long as 
blade, the middle and upper shorter; uppermost leaves sessile; lower floral leaves resem- 
bling the cauline but smaller, long-acuminate, few-toothed, the upper lanceolate-linear to 
linear-subulate, bracteiform; leaves subtending semiverticels and bracts narrowly linear, 
363 mostly conduplicate and then subaristate, green or lilac, hairy, 10 mm and 3-5 mm long, 
respectively; pedicels 0.3-1 mm long; verticillasters very dense, many-flowered, all or the 
upper tightly crowded in ovoid, rarely cylindrical, spicate inflorescence 4-7 cm long, 2- 
4.3 cm wide; 1-2 (3) lower pairs of semiverticels sometimes remote, with peduncles 3-10 
(15) mm long; calyx 8-10 mm long, slightly curved and dilated in upper part, very promi- 
nently 15-nerved, densely covered with antrorse hairs, these short, appressed, in lower part 
longer and spreading; teeth oblong-triangular, long-acuminate, 1/4-1/3 (2/5) as long as 
tube; corolla 18-23 mm long, bluish-lilac, whitish-hairy outside, the tube exserted to 2- 

3 mm, abruptly expanding into large neck, 4-5 mm long and as wide; upper lip cleft to 
1/3-1/2 into orbicular-obovate lobes; middle lobe of upper lip 4.3-5 mm long, 6.5-7 mm 
wide, rounded-truncate, the lateral lobes semiorbicular-triangular, 1.3-1. 7 mm long, 2.7- 

4 mm wide; upper stamens mostly somewhat shorter than upper lip; nutlets oblong-ellip- 
soid, 2-2.5 mm long, 1-1.3 mm wide, brown, obscurely foveolate-tuberculate, nearly 
smooth in places, with papiUiform apical protuberances sometimes coalescent into a 
coriaceous appendage and with 3-5 oblong dark ribs on the back. Fl. June to first half 
of August; fr. from second half of July. 

From upper forest to alpine belt, meadows, tall-grass ranges, forest margins. — Caucasus: 
W. Transc. (upper reaches of basin of Kubana River), E. and S. Transc. Endemic. Described 
from vicinity of Sanain in Georgia. Type in Tbilisi. 

41. N. grossheimii Pojark. in Bot. mat. gerb. Bot. inst. AN SSSR, XV (1953) 310. - 
N. betonicaefolia Grossh. Opred. rast. Kavk. (1949) 330, p.p. non C. A.M. 

Perennial; root woody, twisted, splitting into coarse fibers, passing above into short 
rhizome; stems few, 15-40 cm long, 1.7-3.5 mm across, erect or slightly ascending, un- 
branched, green or slightly violet-tinged, rather densely covered with short flexuous sub- 
accumbent hairs, with 7-9 internodes (to inflorescence) of which the longest (the upper) 



243 



4-7.5 cm long; leaves upright (in herbarium often all or most parallel and appressed to 
stem), bright green, almost concolor, sparingly or rather densely covered with very fine 
short 3-5-jointed simple hairs and beset beneath with punctate glands, triangular-ovate 
or oblong-ovate (the upper sublanceolate), with deeply cordate base, acute (rarely some 
364 obtuse), each margin with (11) 13-15 large teeth, these mostly acute, rarely subobtuse; 
middle leaves largest, 2.5-4.8 cm long, 1 .8-2.7 cm wide; lower petioles ca. 2/3 as long to 
as long as blade, the middle short, the upper sessile; lower floral leaves resembling the 
cauline but smaller and narrower, lanceolate, acuminate, with fewer more acute teeth or 
partly entire, the upper lanceolate-linear, narrow, completely resembling bracts; inflores- 
cence 4.6-1 1 cm long; semiverticels dense, many-flowered, the lower 1 or 2 on peduncles 
0.5-1 cm long, 1-3 cm apart, the others subsessile, tightly crowded in a dense ovoid or 
cylindrical spike 2-3 cm wide; bracts narrowly hnear, nearly setiform, 1/2-2/3 as long as 
calyx, violet, densely covered outside and along margin with long fine soft (5) 8-10- 
jointed white hairs; pedicels 0.5-1 mm long; calyx 7-8.5 mm long, violet, with narrow 
distally dilated tube and slightly oblique throat, densely covered in lower part vvith nearly 
straight long spreading white hairs, in upper part with shorter antrorse hairs, also beset 
with sparse minute sessile white glands; teeth subequal (the lower sometimes narrower), 
triangular, gradually or abruptly acuminate, corolla 15-17 mm long, bright, bluish-violet, 
densely white-puberulent outside, the tube exserted to 1.5-2 mm, abruptly expanding into 
neck 3-4 mm long and as wide; upper lip 3-3.5 mm long, cleft slightly beyond middle in- 
to suborbicular lobes; middle lobe of lower lip twice as wide as long, its margin with sev- 
eral large triangular teeth, the lateral lobes triangular-semiorbicular, 1 .5-2 mm long, 2.8- 
3.3 mm wide; style slightly longer than upper Up, the upper stamens slightly shorter; 
nutlets unknown. Fl. end of May to first half of July. (Plate XXIII, Figure 1.) 

Meadow slopes in upper belt of xerophilous open woodland. — Caucasus: S. Transc. 
(Zangezur Range). Endemic. Described from western slope of Zangezur Range below 
Bichenakh mountain pass. Type in Leningrad. 



Series 4. Strictifoliae Pojark. — Calyx scarcely curved, with slightly obUque throat 
and subequal teeth; corolla 13-18 mm long; narrow part of tube barely exserted; nutlets 
with dense but isolated distinct tubercles; bracts linear-lanceolate to setiform, conduph- 
cate and curved; verticillasters remote, the upper largest, loosely approximate, not con- 
fluent; stems solitary or few, nearly always simple; leaves upright, more or less parallel 
to them. 
365 In addition to the Transcaucasian species, this series contains N. elbursensis Rech. 
fil. (N. Iran - Elburz Range), N. speciosa Boiss. et Noe (S. Kurdistan), N. glabrescens 
Boiss. (Arm.-Kurd. - southern spurs of Bimgel mountains) and N. concolor Boiss. et 
Heldr. (southeastern part of Asia Minor - Tavr Isaurskii). 

42. N. strictifolia Pojark. in Bot. mat. gerb. Bot. inst. AN SSSR, XV (1953) 303. - 
N. betonicaefolia Grossh. Fl. Kavk. Ill (1932) 294, p. p. non C. A.M. - N. reiche- 
bachiana Grossh. in Dokl. AN Azerb. SSR (1949) 70; idem, Opred. rast. Kavk. 332, 
p. p. non Fisch. et Mey. 

Perennial; root woody, branching, rather slender; stems few (1-3), straight, sturdy, 



244 



upright, simple, very rarely with 1 or 2 pairs of leafy fertile branches in upper part, 
35-80 cm long, 2-3.5 mm across, with 14-16 (18) internodes (including inflorescence) 
5-10 cm long, at first densely covered with very short slightly crisp white hairs; leaves 
thin, not rugose, with prominent, upright or (especially the upper) parallel to stem, both 
sides densely covered with small (magnifying glass!) simple 2-3 (4)-jointed hairs and scat- 
tered punctate pale yellow glands, usually only the thin secondary veins prominent; mid- 
dle leaves largest, 2.5-4.5 cm long, 0.8-2.3 cm wide, oblong-ovate or lanceolate, acuminate, 
rarely obtuse, with shallowly cordate or nearly truncate base, each margin with 10-13 large 
acute or subacute unequal teeth; lower petioles mostly short, rarely the longest 1/3 the 
length of blade, others still shorter; uppermost leaves sessile; only lowermost floral leaves 
resembling the cauline though smaller, the others Hnear-lanceolate, the upper almost seta- 
ceous, bracteiform; inflorescence lax, oblong, 5-18 cm long, consisting of 5-10 pairs of 
few-flowered semiverticels, the 2-4 lower 2.5-4 cm apart, with peduncles 4-10 mm long, 
the others loosely approximate (not confluent), sessile or subsessile; leaves subtending 
semiverticels and bracts violet, linear-setaceous, the first 5-8 mm, the second 3-5 mm long; 
pedicels 0.5-0.7 mm long; calyx 7-9 mm long, tubular, straight, with slightly oblique 
throat, with 15 prominent thick nerves, violet-blue, densely covered with short antrorse 
fine slightly crisp white hairs, subappressed on tube, spreading on teeth; teeth triangular- 
lanceolate, mostly gradually acuminate, 2/7-2/5 as long as tube, all equal or the 2 lower 
slightly narrower; corolla 14-16 mm long, bright violet-blue, densely covered outside with 
366 mostly spreading fine hairs, these longer at outer side of hmb; tube slender, exserted to 
ca. 1 mm, abruptly expanding into elongate neck, 3-3.2 mm long; upper lip cleft to mid- 
dle into broad suborbicular lobes 2-2.5 mm wide; lower lip approximately twice as long 
as upper, the middle lobe 3.5-4.5 mm long, 5.5-7 mm wide, the lateral lobes triangular- 
semiorbicular, 1 .3-1 .5 mm long, 2-2.5 mm wide; upper stamens as long as upper Up of 
corolla; in a once encountered female specimen corolla with less developed lower lip, 
longer style and staminodes included in the enlarged part of tube; nutlets pale brown, 
oblong-obovoid, obtusely 3-angled, 1 .8-2.3 mm long, 0.8-1 .3 mm wide, with small remote 
obtuse tubercles. Fl. second half of May to June; fr. from first half of June. (Plate XXII, 
Figure 2.) 

In the belt of xerophilous open woodland, on meadow and steppe slopes of northern 
exposure and in light forests. - Caucasus: S. Transc. (Nakhichevan ASSR, southern part 
of Armenia). Gen. distr.: Arm.-Kurd. (southern part of Kars). Described from Nakhi- 
chevan ASSR, Almabulak River. Type in Leningrad. 

Notes. l)Crosses N. strictifolia Pojark. X transcaucasica Grossh. encountered in 
the Nakhichevan ASSR comprise different combinations of features characterizing these 
well differentiated species, gravitating toward one or the other. 

2) All the three Caucasian species which we have segregated in the series Strictifoliae, 
were erroneously referred by Grossgeim to N. betonicifolia C.A.M. in which, judging 
by his herbarium determinafions, he also included N. somkhetica Kapell., although 
the latter species had at one time been nominally acknowledged by him. In 1949 he clas- 
sified a complex of S. Transcaucasian forms as a species to which he quite inappropriately 
assigned the name N. reichenbachiana Fisch. et Mey. This species belongs to the cycle 
of forms allied to N. mussinii Spreng. and has Uttle in common with species of the series 
Strictifoliae. 



245 



43. N. alaghezi Pojark. in Bot. mat. gerb. Bot. inst. AN SSSR, XV (1953) 307. - N. 
speciosa E. Busch in Tr. Tifl. bot. inst. I (1933) 211, non Boiss. et Noe. — N. reichen- 
bachiana Grossh. in Dokl. AN Azerb. SSR, V (1949) 70; idem,Opred. rast. Kavk. 332, 
p.p. non Fisch. et Mey. 

Perennial, with woody branching root passing into short rhizome; stems few, erect, 
simple, 25-70 cm long, 1 .5-2.5 cm across, covered with minute (magnifying glass!) 1-2- 
jointed thickish flexuous appressed simple hairs (in lower part longer, spreading); inter- 
nodes 6-7 to inflorescence (including inflorescence 12-16), the upper 4-5 cm long; leaves 

367 20-32 mm long, 0.9-2 mm wide, thin, not rugose, at least the upper upright, oblong-ovate, 
with cordate or subtruncate base (the lower often broadly ovate, with deeply cordate 
sinus), obtuse or sometimes acuminate at apex, each margin with 12-14 large obtuse teeth, 
both sides covered (like stem) with fine appressed hairs, the lower side with longer, spread- 
ing hairs and densely punctate -glandular; lower petioles 2/5-2/3 as long as blade; upper 
cauhne and lower floral leaves with petiole 1.5-2 mm long; lower floral and cauUne leaves 
longer than semiverticels; upper floral leaves small, lanceolate-hnear, colored, bracteiform; 
inflorescence 6-25 cm long, lax, oblong, consisting of 6-9 verticillasters; upper verticilla- 
sters 4-10-flowered, subsessile, approximate but not confluent, the lower 12-16-flowered, 
dense, 2.5-7 cm apart; peduncles 5-15 mm long; leaves subtending semiverticels and 
bracts blue, narrowly linear or linear-subulate, the first 3-5 mm, the second 1-2.5 mm 
long; pedicels 0.5-1 mm long; calyx 6-7.5 mm long, straight, with slightly oblique throat, 
blue, densely covered with yellowish hairs (these nearly straight, spreading, longer in lower 
part, short, antrorse in upper part) and rather densely beset with round yellow glands; 
teeth with broad base, triangular, gradually or abruptly tapering to a point, the margin 
long-cihate; corolla 14-15 mm long, pubescent outside; tube curved, exserted to 1- 

1.5 mm, abruptly expanding into neck, 3-3.5 mm long; upper Up 3-3.5 mm long, cleft to 
middle or slightly less into obovate or suborbicular lobes 2-2.5 mm wide; middle lobe of 
lower lip 4 mm long, 7 mm wide, with large teeth, the lateral lobes 1 .2-1 .3 mm long, 3- 
3.5 mm wide; upper stamens as long as upper lip; style slightly exserted from corolla; 
nutlets brown, ellipsoid, 2.5 mm long, 1 .3-1 .4 mm wide, mostly truncate and toothed at 
apex by acutish tubercles, covered on both sides with small obtuse tubercles, the dorsal 
side with 3 rather prominent ribs. Fl. July -August; fr. from first half of August. 

Subalpine meadow coenoses on stony and pebbly slopes. — Caucasus: S. Transc. (NW 
Armenia, Mount Aragats and Mount Alibek). Endemic. Described from Mount Aragats 
(Alagez), Karamlykh ravine. Type in Leningrad. 

44. N. buhsei Pojark. in Bot. mat. gerb. Bot. inst. AN SSSR, XV (1953) 299. - N. 
racemosa Benth. in DC. Prodr. XII (1848) 385, p. p. (quoad pi. pers.) non 1834; Boiss. 
et Buhse in Nouv. Mem. Soc. Nat. Mosc. XII, 175; Bge. in Mem. Acad. Sc. Petersb. VII, 

368 ser. XXI, 1, 54 (excl. pi. Szovits.); Boiss. Fl. or. IV, 660, p. p. (quoad pi. pers.) non 
Lam. - ? N. racemosa var. ghilanica Benth. in DC. Prodr. XII (1848) 385. - N. rei- 
chenbachiana Grossh. in Dokl. AN Azerb. SSR, V( 1949) 70; idem,Opred. rast. Kavk. 
332, p. p. non Fisch. et Mey. 

Perennial; root woody, more or less twisted, branching, passing into short horizontal 
rhizome; stems few, erect, 30-70cm long, 1.5-2. 5mm across, covered with short soft 3-4 
(5)-jointed spreading hairs; internodes6-7 to inflorescence (13-16 including inflorescence). 



246 



the lower short, the upper 5-7 cm long; lower leaves obliquely upright, the upper parallel 
to stem, not rugose, with prominent larger veins, yellowish-green, broadly ovate, with cor- 
date base (only upper leaves oblong-ovate, with nearly truncate base), each margin with 
9-19 large obtuse or acute teeth, the upper side with very slender short spreading 3-5 
(7)-jointed hairs, the lower side with longer (especially on veins) subappressed hairs and 
densely covered with punctate glands; cauline leaves 1 .9-3 cm long, 1 .3-2.3 cm wide; 
petioles short, the lower 1/6-1/4 as long as blade, the others shorter; upper leaves sessile; 
bracts shorter than semiverticels, the 2 or 3 lower pairs resembling cauline leaves but 
smaller, the others narrowly lanceolate, small, colored, bracteiform; inflorescence 10- 
25 cm long, lax, consisting of 6-9 verticillasters; upper semiverticels 2-4-flowered, sub- 
sessile, in subremote pairs, the others 6-12-flowered, dense, peduncles 10-1 5 mm long, 
very distant; leaves subtending semiverticels and bracts hnear-subulate or rarely lanceo- 
late-linear, the first 4-5 mm long, the second 1 .5-2.3 mm long, blue, covered with long 
soft hairs; pedicels 0.6-1 mm long; calyx 6.5-7.5 mm long, nearly straight, with slightly 
obhque throat, blue, 15-nerved, densely tomentose-villous with implexed white or blue 
hairs; teeth subequal, triangular, with broad base, in upper part abruptly (rarely gradual- 
ly) acuminate, 2/7-1/3 as long as tube; corolla 13-14 mm long, pubescent outside, with 
slender tube exserted to 1 .5 cm [sic], abruptly expanding into neck 3-4 mm long; upper 
hp cleft slightly beyond middle into broad obtuse lobes; middle lobe of lower lip 4 mm 
long, 6-7 mm wide, with large triangular marginal teeth, the lateral lobes nearly semiorbi- 
cular-triangular, 1 .2-1 .3 mm long, 2.1-2.6 mm wide; upper stamens with filaments slightly 
longer than upper lip of corolla, lower stamens and style slightly shorter; nutlets brownish- 
black, ellipsoid, 2 mm long, 1-1.2 mm wide, unevenly obtusely tuberculate, the 3 longitu- 
dinal dorsal ribs not very prominent, the ventral side with a sharp edge. Fl. June; fr. from 
first half of July. 

Exposed gravelly and stony slopes in upper part of central mountain belt and in subal- 
pine belt. — Caucasus: Tal. Gen.distr.: Iran, (northern part adjacent to Tal., Iranian 
Azerbaidzhan, Gilyan). Described from vicinity of Karabak-Yurdinskii in Tal. Type in 
Leningrad. 

Cycle 2. Mussinianae Pojark. - Calyx markedly curved, with oblique throat; lower 
teeth smaller than the upper; corolla (10) 12-20 (22) mm long, the narrow part of tube 
slightly exserted; nutlets pitted and obtusely tuberculate. Inflorescence a more or less 
one-sided raceme consisting of remote or distally more or less crowded few-flowered ver- 
ticillasters; stems many; leaves subhorizontally patent. 

This series contains species growing in Transcaucasia and Southwest Asia. In addition 
to species described below, it should also include N. crassifolia Boiss. et Buhse (Elburz 
Range), N. longiflora Vent. (N. Iran), N. mariifolia Boiss. et Huet (Erzerum) and N. 
daflersiana Schfth. (mountains of Yemen in the Arabian Peninsula). 

45. N. mussinii Spreng. in Henckel, Adumbr. pi. (1806) 15; M. B. Fl. taur.-cauc. II, 
39, III, 390; Benth. in DC. Prodr. XII, 385, p. p. quoad pi. iber.; Ldb. Fl. Ross. Ill, 376, 
p.p.; Boiss. Fl. or. IV, 660, p. p.; N. Pop. in Tr. Bot. sada Yur'evsk. univ. XIV (1914) 
234, p.p.; Grossg. Fl. Kavk. Ill, 295 p.p.; idem,Opred. rast. Kavk. 332, p.p. - N. longi- 
folia Sims in Curtis Bot. Mag. XXIII, I (1806) 23, non Vent. - N. salviaefolia Pers.Syn. 



247 



II (1807) 125. - N. willdenoviana Adams in Hoffm. Hort. mosc. (1808) 25 (excl. 
synon. Vent.). — N. grandiflora 7. mussini N. Kuzn. in Uch. zap. Yur'evsk. univ. 2 
(1899) 48. - Ic: Rchb. Ic. bot. VI, tab. 587, fig. 806; Sims, 1. c. tab. 923 (sub. N. lon- 
gifolia). — Exs.: PI. or. exs. No. 270. 

Perennial; root woody, slender, more or less knotty, coarsely fibrous, passing into 
woody short rhizome, sometimes branching; stems many, 13-40 cm long, slender, 0.7- 
1 .5 mm across, ascending, with woody sometimes perennial bases, with 12-16 internodes 
(including inflorescence), at first unbranched or with few short mostly sterile axillary 
branches, later often developing rather numerous branches, densely grayish-hairy, the hairs 
very short, fine, crisp, appressed (sometimes longer and spreading in lower part of stem); 
leaves thin, the upper side with network of nerves and thus often rugose, green or grayish- 
green, densely covered with very short appressed hairs, the lower side with some prominent 
veins, usually grayish tomentose-velutinous or velutinous all over with very fine short im- 

370 plexed hairs and with numerous punctate yellow glands; cauUne leaves (7) 10-24 mm long, 
(6) 7-10 mm wide, oblong-ovate, rarely ovate, obtuse, more or less deeply cordate, crenate 
or dentate-crenate from base (with 11-17 teeth on each margin); lower petioles ca. 2/3 as 
long as blade, the upper 1/4-1/2 as long; leaves of axillary branches narrower, usually lan- 
ceolate, often acute; lower floral leaves resembhng the cauline but subsessile or sessile, the 
upper bracteiform; inflorescence of 4-8 (10) verticillasters forming a more or less one- 
sided raceme (2) 3-10 (14) cm long; upper semiverticels subsessile, crowded at summit, 
often confluent, the lower on peduncles 0.5-2 cm long, in remote pairs; bracts narrowly 
lanceolate, usually conduplicate, 1.5 -2. 5 mm long; leaves subtending semiverticels slightly 
longer; calyx (5) 6-7.5 mm long, usually bluish, softly tomentose-arachnoid with implex- 
ed long patent jointed blue and white hairs; teeth triangular to lanceolate, (1/3) 1/4-2/7 
as long as tube, the 2 lower sUghtly narrower and often slightly shorter than the upper; 
corolla (9) 10-13 mm long, densely short-hairy outside, lilac-blue, with darker violet-blue 
spots at throat; tube exserted to 1-1 .5 mm, abruptly expanding into neck, ca. 3 mm long; 
upper lip cleft to middle into obtuse lobes; middle lobe of lower lip 3-4 mm long, 4.5- 

6 mm wide, strongly concave, coarsely crenate, the lateral lobes obliquely semiorbicular- 
triangular, 2-2.5 mm wide, 1-1 .5 mm long; style and upper stamens as long as upper lip 
of corolla; nutlets blackish-brown, broadly elUpsoid, 1 .75-2 mm long, ca. 1 mm wide, 
densely covered with small obtuse tubercles. Fl. first half of April to July; fr. second 
half of June to September. 

Dry gravelly and stony slopes, from lower mountain belt to subalpine. — Caucasus: E. 
Transc.(Tushetia, near Tbilisi, Trialet Range). Gen.distr.: Arm. -Kurd. (Kars). Describ- 
ed from live cultivated plants, apparently originating from near Tbilisi. Type unknown. 

Note. N. mussinii belongs to a very polymorphous cycle and is clearly an aggregate 
species even after separation of N. transcaucasica Grossh. Determination of the geo- 
graphical races presents considerable difficulties and cannot be achieved without careful 
observations in nature. A serious obstacle in the way of systematic treatment of this cycle 
is the great plasticity of its forms that is revealed as conditions of habitat change. Kuznet- 
sov (1899) reported that, when grown under northern conditions, N. mussinii changes 
radically in habit in the course of one vegetative period: high, strongly branching stems 
are produced, indument is lost, leaves become large and broader. This is also observed in 

371 nature, although not in such drastic form: individuals growing on exposed dry slopes 



248 



differ markedly from those growing in the shade or near a source of water, and stems pro- 
duced at the onset of summer differ from those developing and flowering in July-Septem- 
ber. All the same, it would hardly be justified at present to accept N. mussinii in the 
customary wide scope. Some races may be segregated although their differentiation in 
the herbarium presents difficulties. It is relatively easier to delimit N. mussinii Spreng. 
s. str. (as conceived within the above-outlined geographical limits). A reliable character 
for recognizing N. mussinii Spreng. s. str. is its indument which is made up of hairs that 
are much finer and more implexed than in other forms of the cycle (the difference be- 
comes very clear when the lower side of the leaves is viewed under high magnification). N. 
mussinii s. str. is distinguishable from the outwardly most similar W. Armenian race 
N. reichenbachiana F. et M. also by the usually smaller flowers, more compact inflo- 
rescences, more floriferous cymes and the corolla (judging by Reichenbach's drawing), 
which is apparently not quite the same color. 

46. N. reichenbachiana Fisch. et Mey. in Ind. sem. hort. Petrop. VIII (1841) 68. - ? 
N. racemosa Lam. Encycl. meth. Bot. I (1785) 711, non Benth. (1848) necBoiss. 
(1879). - N. racemosa i3. reichenbachiana Benth. in DC Prodr. XII (1848) 385; Ldb. 
Fl. Ross. Ill, 376. 

Perennial; root woody, coarsely fibrous, more or less twisted; stems many, ascending, 
30-65 cm long, slender, grayish with short fine reclinate crisp hairs, with 13-20 (22) inter- 
nodes, at onset of flowering simple or with few axillary branches; stems vegetating at end 
of summer, usually strongly branching; branches slender, leafy, all or the upper sterile; 
leaves more or less rugose above by impressed network of veins, pale green, rather densely 
covered with short fine crisp appressed hairs, the lower surface grayish or whitish, with 
longer hairs (sparser and longer still on the veins) and often with small capitate glands; 
cauline leaves ovate-lanceolate or lanceolate, rarely oblong-ovate, with shallowly cordate 
or subtruncate base, obtuse or rarely acuminate, finely crenate from base (with 12-16 
crenae on each margin), (8) 12-23 mm long, 7-12 mm wide; lower petioles 1/3-1/2 as 
long as blade, the upper shorter, verticillasters 5-8 (10) pairs, forming a lax, more or less 
one-sided raceme; semiverticels of 2-4, the lower of 6 flowers, the upper subsessile, the 
372 pairs loosely approximate, not confluent, the lower 1.5-3 cm apart, on peduncles 5-1 6 mm 
long; lower floral leaves resembling upper cauline leaves but sessile, the upper small, 2- 
5 mm long, lanceolate, entire; leaves subtending cymes 1.5-2 mm long; bracts 1-1.5 mm 
long, narrowly lanceolate or linear-subulate, arcuately curved, violet, densely pubescent; 
pedicels 1-2 mm long; calyx 6.5-9 mm long, violet, arachnoid- or downy -tomentose, with 
violet and white implexed hairs; teeth triangular or narrowly triangular, 1/5-2/7 as long 
as tube; corolla 1 1-14 mm long, faintly violet-blue, densely white-hairy outside; tube 
slender, exserted to 1-1 .5 mm; upper lip cleft to middle into obtuse lobes; middle lobe 
oflowerlip3mmlong, 5mm wide, undulate-crenate, the lateral lobes semiorbicular-trian- 
gular, 1-1 .5 mm long, 2-2.5 mm wide; nutlets dark brown, obovoid, 3-angled, obtusely 
tuberculate-foveolate, with 3 more or less prominent longitudinal ribs on dorsal side. Fl. 
May-August (September); fr. from June. (Plate XXII, Figure 3.) 

Rocky and stony slopes in middle mountain and subalpine belts. — Caucasus: S. Transc, 
W. Armenia, and, apparently, Nakhichevan ASSR. Gen. distr.: Arm. -Kurd. Described 
from Mount Ararat. Type in Leningrad. 



249 



Notes. 1) Bentham considered N. reichenbachiana Fisch. et Mey. as a variety of 
N. racemosa Lam. and Boissier regarded them as identical. Neither of these authors saw 
either the type of N. racemosa Lam. (described from material by Tournefort) or that of 
N. reichenbachiana Fisch. et Mey. and obviously drew their conclusions from the diag- 
noses of these species which are, in fact, very similar. It cannot be ruled out that these 
two species are not actually identical since Tournefort's route passed through that part of 
Armenia which includes Mount Ararat, the area of N. reichenbachiana. It should also 
be noted that the plant described by Reichenbach (Rchb. Ic. bot. tab. DLXXXV, fig. 804) 
under the name N. racemosa Lam. is very similar to N. reichenbachiana. Proceeding 
from the impossibility of proving the identity of N. racemosa Lam. with N. reichen- 
bachiana Fisch. et Mey., we have chosen the second epithet for the Armenian race as 
quite reliable, all the more so since the name N. racemosa was applied in the past to a 
wide array of completely unrelated species. Even Bentham and Boissier identified N. 
racemosa Lam., not just nominally but also in practice, with species remote from N. rei- 
chenbachiana, listing under this name mainly specimens from N. Iran (Buhse, Bunge, 
Oshe, Kotschy) that belong to species now described under the name N. elbursensis 
Rech. fil. and N. buhsei Pojark. Grossgeim (1944) made it clear that he had earlier 
reported N. transcaucasica Grossh. from the Caucasus under the name N. racemosa. 
373 2) In his last works, Grossgeim (1949), being unacquainted with the type of N. rei- 
chenbachiana, erroneously included under this name the Caucasian species which have 
here been separated in the series Strictifoliae. 

3) In habit, N. reichenbachiana resembles most closely N. mussinii Spreng., from 
which it differs in the characters indicated in the Note to the latter species. Fully con- 
forming with typical specimens of N. reichenbachiana are plants from the vicinity of 
Erevan (spurs of Akhmagan Range), where I was able to observe and collect this species, 
and also specimens from Mount Aragats. From the material available it is difficult to as- 
certain the eastern limit of this species (Nakhichevan ASSR) and differentiate it from the 
very polymorphous and, apparently, aggregate species N. transcaucasica Grossh. 

47. N. hajastana Grossh. in Izv. Azerb. fil. AN SSSR, 10 (1944) 88, p. p. quoad typ.; 
idem, Opred. rast. Kavk. 332, p. p. 

Perennial, very close to N. reichenbachiana Fisch. et Mey. and distinguished from it 
by the erect, longer (30-90 cm) and thicker (2.5-3 mm) stems, mostly branching, grayish 
with very fine appressed pubescence; internodes to 5-7.5 cm long; branches slender, fer- 
tile; subsidiary stems arising from rhizome in varying number, densely leafy, short, also 
erect, partly fertile, but mostly sterile, with smaller and narrower leaves; leaves on main 
stems triangular-ovate, rarely oblong-ovate, the middle largest, 1.5-3.3 cm long, 0.8- 
2.3 cm wide, obtuse, with cordate base; inflorescence to 30 cm long, of 8-1 1 verticilla- 
sters, the upper rather loosely approximate or 1-2 cm apart, the lower 4 or 5 3-6 cm 
apart; semiverticels 2-6-flowered, the upper subsessile, the others on peduncles 1-2.5 cm 
long; lower floral leaves large, foliaceous, as long as peduncles, the upper bracteiform; 
bracts, calyx and corolla resembling those of N. reichenbachiana in size, shape, color 
and vesture; nutlets also completely alike. Fl. April-June; fr. from end of April. 

Stony and silty-stony slopes, ca. 1000 m. — Caucasus: S. Transc. Described from the 
Erevan suburb of Nork, the only location from which it is known. Type (lectotype) in 
Leningrad. 



250 



(375) 




PLATE XXII. 1 - N cvanea <;tPv fi 

nutlet in dorsal view; 3 N relhe'nbirhT ~\''''''''-'^^ Pojark., general aspect, flower, 

in ventral view. reichenbach.ana Fisch. et Mey., upper part of plant, flower, nutlet 



251 



Notes. 1) A critical species, in need of further study, especially in nature. As already 
reported, it is very close to N. reichenbachiana Fisch. et Mey. from which it differs, 
practically speaking, only in the high, erect stems, larger and broader leaves, and sparser 
374 and longer inflorescences. I was able to observe both species in flower (N. hajastana in 
its classical location and N. reichenbachiana near the Gekhart monastery at the spurs 
of Akhmagan Range) and must note that the above indicated distinctions between them 
still hold good. Consequently, I have decided to retain the separate identity of the two 
species. 

2) N. hajastana Grossh. has been described from herbarium specimens collected in the 
vicinity of Nork (near Erevan) and Bicheny (Nakhichevan ASSR). Careful comparison of 
authentic specimens showed differences between them and precluded their inclusion in 
one species. According to the author's diagnosis, it can be concluded that the species was 
described from a plant collected at Nork, and I have therefore accepted it as the type. The 
plant from Bicheny, which differs from N. hajastana in having a straight calyx with 
slightly obUque throat, coarser indument, and a short inflorescence consisting of a few pairs 
of short-peduncled semiverticels, should be included among crosses N. strictifolia X 
transcaucasica which are rather common in this part of S. Transcaucasia. 

48. N. transcaucasica Grossh. in Izv. Azerb. fil. AN SSSR, 10 (1944) 38; idem,Opred. 
rast. Kavk. 332. - N. longiflora C. A. M. Verzeichn. Pfl. Cauc. kasp. Meer. (1 831) 92, 
non Vent. - N. mussini auct.: Benth. in DC Prodr. XII (1848) 385, p. p. (excl. pi. iber.); 
Ldb. Fl. Ross. Ill, 376, p. p.; Boiss. Fl. or. Ill, 660, p. p.; Grossg. Fl. Kavk. 292, p. p. non 
Spreng. — N. racemosa Grossh. Fl. Kavk. Ill (1932) 296, non Lam. 

Perennial; root and rhizome as in N. reichenbachiana; stems many, ascending, 10- 
40 (50) cm long, 1-2 mm across, with internodes 3-6 (7) cm long, simple or branching 
mainly in lower part, densely patent-hairy, the hairs fairly long, especially in lower part, 
crisp, often shorter and subappressed in upper part; branches usually short, sterile, rarely 
fertile; leaves bright green above with copious short appressed hairs and impressed network 
of veins, grayish beneath, more densely covered with longer hairs; cauhne leaves much 
shorter than internodes, cordate-ovate, rarely oblong-ovate to lanceolate, 10-32 mm long, 
7-20 (22) mm wide (smaller and narrower on axillary branches), obtuse, evenly and often 
rather deeply crenate with (9) 11-16 crenae on each margin; lower petioles about the 
length of blade, the upper shorter, 1/6-2/5 as long as blade; lower floral leaves resembhng 
377 the cauline, usually as long as semiverticels without corollas, the upper bracteiform, narrow, 
entire; inflorescence 5-10 cm long, rarely slightly longer, consisting of small number of 
dense verticillasters, the upper 3 or 4 (often all) crowded at summit, 1 to 3 remote; semi- 
verticels 4-8-flowered, the upper sessile or subsessile, the lower on peduncles 3-10 (15)mm 
long; leaves subtending semiverticels narrowly lanceolate, 3-5 mm long; bracts 1.5-2 mm 
long, linear-oblong; pedicels 0.5-2 mm long; calyx 8-11.5 mm long, arachnoid-tomentose 
or tomentose-pilose outside (like bracts), the long soft hairs blue and white, of 4-9 cells; 
throat strongly obUque; teeth lanceolate, acuminate, the upper (1/4) 2/7-1/3 as long as 
tube; corolla 16-20 (22) mm long, violet-blue, white-pubescent outside, the narrow part 
of tube exserted 2-3 (3.5) mm, expanding into neck 3-4 mm long; upper lip cleft to 2/3- 
3/5 into ovate obtuse lobes; lower Hp twice as long as the upper, the large middle cordate 
lobe to 4-5 mm long, 8-9 mm wide, the lateral lobes obhquely semiorbicular, ( 1) 1 .3-1 .6 mm 



252 



long, 2.5-3.5 mm wide; upper stamens and style as long as upper lip; nutlets dark 
brown or blackish-brown, obtusely foveolate-tuberculate, broadly ellipsoid or nearly 
obovoid, 1 .75-2 mm long, 1 .25-1 .3 mm wide, with 3 prominent longitudinal ribs on the 
back. Fl. May to first half of August; fr. from end of June. 

Dry stony and pebbly slopes, pebbles in central and subalpine mountain belts. - Cauca- 
sus: E. and S. Transc, Dag. (extreme south), Tal. Gen. distr.: western parts of Iran. 
Described from vicinity of Shush. Type in Baku. 

Notes. 1) In describing N. transcaucasica, Grossgeim contrasted it with N. mussinii, 
indicating its higher growth, larger leaves and calyx, and distribution in S. Transcaucasia 
and Talysh. In accordance with this characterization of his species, Grossgeim consistently 
regarded the lower and more narrow-leaved forms growing in the above-named regions as N. 
mussinii, a position with which it is impossible to agree. N. mussinii can be reliably dis- 
tinguished from N. transcaucasica only by the very fine indument of the leaves and 
stems although, on the whole, it tends to be a lower plant with narrow leaves, while its 
flowers do not attain the large size often observed in N. transcaucasica. 

2) It is far more difficult to distinguish N. transcaucasica from N. reichenbachia- 
na which, like N. mussinii, is characterized by narrow leaves and small flowers. Even 
though one may discern certain characteristic features of N. transcaucasica, such as a 
slight tendency toward branching of the stems, terminal part of its inflorescence usually 
compact, nearly spicate, peduncles always short, and corolla-tube more exserted from the 
calyx, it is often impossible to differentiate with certainty between herbarium specimens 
378 of N. transcaucasica and N. reichenbachiana, especially in the western part of the 
range, where these species are linked by intermediate forms, possibly hybrids. However, we 
have decided to acknowledge N. transcaucasica Grossh. on the following grounds. In 
the more eastern regions (Karabakh, Gandzha, Talysh) there is a definite predominance of 
specimens with typical characters of N. transcaucasica, while such forms are unknown 
within the range of N. reichenbachiana. The specific status of N. transcaucasica, as 
well as its extent and contiguity, still have to be determined. Only by careful observation 
in nature will it be possible to sort out the diversity of forms revealed by herbarium mate- 
rial from Southern and Southeastern Transcaucasia. 

Among this diversity, plants from Talysh attract attention by more pronounced charac- 
ters: always densely hairy leaves, corolla-tube exceeding calyx by 3-3.5 mm, a feature 
which allies them to N. longiflora Vent, and probably accounts for their erroneous inclu- 
sion in that species by C. A. Meyer. 

49. N. noraschenica Grossh. in Dokl. AN Azerb. SSR, V (1949) 71; idem, Opred. rast. 
Kavk. 332. 

Perennial; root slender, woody, branching, passing into short rhizome, stems numerous, 
25-45 cm long, 1 .2-2 mm across, ascending, more or less arched, simple or with 1-2 pairs 
of short branches, green, with long and soft spreading simple 4-6-jointed hairs, tomentose 
under inflorescence; internodes 5-6 (not counting inflorescence), 3-6.5 cm long; lower- 
most leaves small, 0.5 cm long, 0.35 cm wide, brown, entire or undulate-dentate, the long- 
hairy petioles about the length of blade; other cauline leaves grayish-green, slightly paler 
beneath, thin, not rugose, the upper side densely covered with short appressed hairs, the 
lower side with much longer (especially along nerves) thinner softer hairs and punctate 



253 



yellow glands with a small admixture of sessile white capitate glands; upper leaves oblong- 
ovate, others ovate, all with deeply cordate base, rounded at apex, with a total of (12) 
14-16 large obtuse or acute teeth; middle leaves largest, 3-4 cm long, 2.3-3 cm wide; 
lower petioles 2/5-1/2 as long as blade, those of middle leaves short; upper cauUne leaves 
sessile; lower floral leaves resembling the upper cauline, usually exceeding semiverticels; 
upper floral leaves lanceolate-Unear, bracteiform; inflorescence 4-9 cm long, consisting of 

379 few sessile verticillasters crowded at summit and 1 or 2 pairs of remote semiverticels with 
short straight peduncles 7-10 mm long; semiverticels dense, 4-8-flowered; leaves subtend- 
ing semiverticels and bracts Hnear-lanceolate or linear, respectively 3-4 mm and 2-3 mm 
long, all densely covered with long hairs; pedicels ca. 1 mm long; flowers bisexual or gy- 
nomonoecious, the latter slightly smaller; calyx 7-9.5 mm long, tubular in flower, ovoid- 
urceolate in fruit, curved, with oblique throat; teeth subequal, triangular or triangular- 
lanceolate, 1/3-2/5 as long as tube, green or in upper part violet-tinged, very densely 
covered with long straight spreading 5-10-jointed soft white hairs; corolla, according to 
Grossgeim, "sky-blue" (violet-bluish when dry), grayish outside with dense pubescence, 
11-13 mm long in female flowers and to 17 mm in bisexual, the slender curved tube bare- 
ly exserted from calyx; upper lip cleft to middle into obtuse broad lobes; lower lip half 
as long as the upper, the middle lobe in bisexual flowers 2.5-3 mm long, 3.8-5.5 mm wide 
(in female flowers relatively smaller), the lateral lobes broadly triangular, 1.3 mm long, 
2-2.75 mm wide; stamens in bisexual flowers with elliptic cells convergent at an obtuse 
angle, as long as upper lip of corolla, in female flowers reduced to staminodes, included in 
dilated part of tube, with small rounded submembranous rudiments of anthers and with 
slender filaments; nutlets dark brown, densely covered with unequal obtuse tubercles, 
broadly eUipsoid or obovoid, 2-2.5 mm long, 1.3-1.5 mm wide. Fl. April-May; fr. from 
first half of May. 

Rocky slopes in lower mountain belt. - Caucasus: S. Transc. Described from Nakhi- 
chevan ASSR, Tandera Mountain near Arpachai, vicinity of Norashen (so far the only 
location). Type in Leningrad. 

Note. It is impossible to agree with Grossgeim, who relates N. noraschenica to N. 
persica Boiss. Similarity between these two species is restricted to the shape and denta- 
tion of the leaves and the long, spreading hairs. Differences between them are far greater and 
evidence of lack of close affinity. Among the features which set N. persica apart are: 
straight campanulate calyx with regular throat; corolla reddish, with white stripes; nut- 
lets ovoid, with strongly convex, tapering papilliform processes. N. persica is also char- 
acterized by abundant indument of stipitate, viscous, glandular hairs. In its aggregate of 
characters, N. noraschenica undoubtedly approaches species of N. mussinii Spreng. 
affiliation, especially N. transcaucasica Grossh. from which it differs in the long, spread- 

380 ing indument, paler corolla and, to a lesser degree, the shape of the leaves, since some large- 
leaved specimens of N. transcaucasica have leaves very similar in shape and dentation to 
those of N. noraschenica. 



Section 6. Micranthae (Boiss.) Pojark. comb. nov. (emend.); Boiss. Fl. or. IV, 639 
(pro subsect.); Briq. in Pflanzenfam. IV, 3a, 238 (pro subsect.). - Calyx tubular, in fruit 
oblong-ovoid or campanulate, curved or straight, with oblique or 2-lipped throat; corolla 



254 



mostly small, 3.8-7 (8.5) mm long or medium-sized, 1 1-1 5 mm long; middle lobe of lower 
lip subhorizontal, spreading or reclinate, concave, without basal swelling; surface of nut- 
lets coarsely tuberculatc to finely granular; flowers in cymes or scmiverticels, these all in 
remcjtc pairs or the upper crowded in a terminal head, their subtending leaves small, brac- 
teiform. Annual, sometimes biennial plants. 

These species are predominantly Iranian, some of them penetrating into Transcaucasia 
and Central Asia. They are distributed in the foothill and low mountain belts. 

Note. The scope of the section Micranthae has been extended here as compared with 
that generally accepted, by inclusion of a number of species formerly incorrectly referred 
to the section Micronepeta. Included in the transfer are N. ispahanica Boiss. and close- 
ly related species which compose the natural phyletic series Ispahanicae Pojark., whose 
characteristics and composition are given below; also a group of species forming the series 
Wettsteinianae Pojark. (not represented in the USSR), which is allied to the series Micran- 
thae and comprises the Iranian species N. wettsteinii H. Braun, N. petraea Benth., N. 
hymenodonta Boiss. and N. kurdica Hausskn. et Bornm. as well as one Afghan species, 
N. afghanica Pojark. 



Series 1. /I moe/iae Pojark. — Flowers medium-sized; calyx 6.5-9 mm long, corolla 
1 1-15 mm long; calyx curved, in flower narrowly tubular, in fruit oblong-ovoid with 2- 
lipped throat; upper teeth ovate, 1/4-1/3 as long as tube; lower teeth triangular-lanceolate, 
incisions between them not exceeding 1/3 the length of calyx; narrow part of corolla-tube 
distinctly exserted from calyx; nutlets coarsely tuberculate; flowers in cymes; lower 
cymes remote, peduncled, the upper sessile, more or less crowded, sometimes forming a 
capitate-spicate terminal inflorescence; leaves subtending cymes and bracts linear-oblong, 
linear, flat, not conduplicate. 

This series contains the two species described below. 

50. N. amoena Stapf in Denkschr. Akad. Wissensch. Wien, 1 ( 1885) 45; Grossg. Fl. 
Kavk. Ill, 291, p. p.; idem, Opred. rast. Kavk. 330, p.p. 
381 Annual or biennial; root slender, tough, woody, vertical or descending; stems simple 
or with few pairs of slender fertile axillary branches, sometimes branching at base with 2- 
8 branches nearly as long as stem, erect or often ascending, sometimes with woody or 
radicant base, 7-60 cm high, 0.6-4 mm across, with sparse or dense (especially in lower 
part) lanate spreading white jointed slightly crisp hairs and more or less densely covered 
with small white sessile glands; leaves thin, bright green above, unevenly covered between 
veins with white short appressed hairs, somewhat paler beneath, with similar though more 
evenly distributed indument, the veins more densely covered with longer lanate hairs; 
cauline leaves 0.8-3.5 cm long, 0.5-3.5 mm wide, broadly ovate, all cordate or the upper 
with truncate base, obtuse or rounded at apex, each margin with 7-12 large obtuse or 
rarely acute teeth; lower petioles as long to 1 ^i-l times as long as blade, the others short; 
upper leaves often subsessile; floral leaves often resembling the upper cauline in size and 
shape, the upper narrower, sparingly toothed to entire, the uppermost oblong, partly suf- 
fused with violet, bracteiform; cymes 3-8-flowered, forming elongate racemes or panicles, 
the upper 2 or 3 pairs of cymes dense, with short axis, resembUng semiverticels on peduncles 



255 



3-8 mm long, loosely approximate to subremote; other cymes on peduncles 1 .3-6 cm 
(the lowermost to 9 cm) long, 2.8 cm apart, sparser, usually loosely dichotomous, with 
axes of first order to 1 .5-2 cm long; bracts 2-5 mm long, linear-spatulate or oblong-linear; 
pedicels 0.8-2 mm long, in cyme bifurcations 2-6 mm long; calyx 6.5-9 mm long, promi- 
nently 15-nerved, violet, rather densely covered with spreading simple 4-6-jointed coarse 
hairs and sparse capitate stipitate glands intermixed with round sessile yellow resinous 
glands; upper teeth 1/4-1/3 as long as tube, ovate, short-pointed or more or less acuminate 
(especially the lateral), the middle tooth distinctly larger than lateral, the two lower teeth 
shorter, narrow, lanceolate, acuminate; corolla pale, bluish-lilac, with darker spots on low- 
er Hp, pubescent outside, 1 1-15 mm long, tube slender, exserted to 1.5-2.5 mm, abruptly 
expanding into neck 2.5-3 mm long, 3-4 mm wdde; upper hp ca. 3 mm long, erect, nearly 
flat, cleft to middle or slightly less into obovate lobes; lower lip twice as long as the upper, 
382 its middle lobe ca. 3.5 cm long, 5.5 mm v^de, strongly concave, with crenate-incised mar- 
gin, the lateral lobes semiorbicular-triangular, 1-1.3 mm long, 2 mm wide; upper stamens 
as long as upper hp of corolla; nutlets 1.5-1.75 mm long, 1-1.2 mm wide, broadly ellip- 
soid, 3-angled, strongly convex on ventral side, dark brown, coarsely tuberculate. Fl. 
April-May; fr. from beginning of May. (Plate XXIII, Figure 3.) 

Foothills and lower mountain belt, steppes, exposed stony pebbly and sandy slopes, 
taluses. — Caucasus: E. Transc. (vicinity of Khudoferinsk in Dzhebrail'skii region of 
Azerbaidzhan). Gen. distr,: N.Iran. Described from near Rudbar in Gilyan (N. Iran). 
Type in Vienna. 

51. N. troitzkii Sosn. in Fl. Gruz. VII (1952) 303 (in Georgian). - N. amoena 
auct. fl. Cauc: Grossg. Fl. Kavk. Ill, 291; idem, Opred. rast. Kavk. 330, p.p. non 
Stapf. — N. amoena auct. fl. Cauc. p.p. non Stapf. - N. micrantha auct. fl. Cauc. p.p. 
non Bge. 

Perennial, very closely related to the preceding from which it is distinguished by the 
following features: stems not lanate in lower part; leaves usually truncate, rarely shghtly 
notched at base; inflorescence much more compact, consisting of (2) 4-5 (7) pairs of 
cymes; upper cymes with up to 8 flowers, very dense, outwardly scarcely differing from 
semiverticels, sessile, the 2 or 3 upper pairs closely crowded at summit of stem in a spicate, 
cylindrical or ovoid inflorescence (sometimes all cymes clustered in a head), lower cymes 
on peduncles 1-4 cm long, less remote than in N. amoena, usually all compact, rarely 
the lowermost distinctly dichotomous, but axes of first order not more than 1 cm long; 
bracts 2-3 mm long, narrowly linear. Otherwise like N. amoena Stapf. Fl. April-May; 
fr. from end of May. 

Stony and gravelly slopes in foothills and lower mountain belt. — Caucasus: E. Transc: 
Georgia (Karayazy) and Azerbaidzhan (Kazakh region, Shirvanskaya steppe, Shemakhin- 
skii region, Apsheron). Endemic. Described from near Karayazy in Georgia. Type in 
Tbilisi. 



Series 2. Micranthae Pojark. - Flowers small, calyx 3.5-7 (8.2) mm long, curved, 2- 
lipped, narrowly tubular in flower, oblong-ovoid in fruit; corolla 3.8-8.5 (9.5) mm long; 
upper teeth 1/4-1/2 as long as tube; lower teeth narrowly lanceolate, incisions between 



256 



them 1/3-1/2 length of calyx; corolla included in calyx to the limb (sometimes even up 
to the middle of upper lip) or only to base of dilated part of tube (neck); nutlets pitted, 
with flat tubercles; flowers in remote peduncled cymes; leaves subtending cymes and 
bracts linear-subulate or setiform, usually condupHcate. 

383 In addition to species distributed in the USSR, the Iranian species N. brachyodonta 
Pojark. belongs to this series. 

52. N. micrantha Bge. in Ldb. Fl. Alt. II (1830) 401; id. Fl. Ross. Ill, 371; Benth. 
Lab. gen. et sp. 476; in DC. Prodr. XII, 382; Kar. et Kir, Enum. pi. song. 638; and B. 
Fedch. Perech. rast. Turk. V, 146 (excl. syn. Boiss.); Kryl. Fl. Zap. Sib. IX, 2312. - Ic: 
Ldb.Ic.pl. fl. Ross. V, tab. 412. 

Annual, (2.5) 5-30 (35) cm high, with slender vertical root; stems erect, simple or 
branching, with inflorescences in the axils of most leaves, more or less densely covered 
with very short reclinate slightly crisp hairs, these simple in lower part, intermixed with 
glandular, predominantly glandular in upper part, sometimes the stem subglabrous, espe- 
cially below; leaves 0.8-4 cm long, 0.7-3.5 cm wide, thin, smooth, with thin, distinct 
(but not prominent) midrib and secondary veins, the upper side pale green, with sparse 
short hairs (these all simple or intermixed with glandular) and with scattered yellow punc- 
tate glands; the lower side glaucescent, with sparse or (especially on veins) profuse glandu- 
lar spreading hairs interspersed with simple 3-4-celled hairs and numerous punctate yellow 
glands, often both sides subglabrous, lower and middle leaves ovate or lanceolate or oblong- 
elliptic, with cuneate or rarely rounded base, obtuse, the margin with 3-5 more or less 
remote obtuse or acute teeth, these often obscure; fairly often all leaves entire; upper 
leaves smaller and narrower, lanceolate or oblong to narrowly linear, mostly acute or acu- 
minate, often equaling but mostly somewhat shorter than peduncles; petioles densely 
glandular-hairy, in lower leaves as long as or slightly longer than blade; cymes few-flower- 
ed, forming a lax narrow raceme or panicle; upper cymes (2) 3-5 (8)-flowered, rather 
compact, with peduncles (1 .5) 3-12 mm long, in pairs 6-10 mm apart but not confluent; 
other cymes 5-8-flowered, sparser, 2-4.5 cm apart, on peduncles 2-3 cm long, often (especial- 
ly the lower) dichotomously branching, with axes of first order up to 8-13 mm long, in large 
individuals sometimes repeatedly branching and then containing up to 12(16) flowers; 
leaves subtending cymes and bracts narrowly linear or setiform, longer than pedicels, hairy; 
pedicels 1 .5-5 mm long, in bifurcation to 10-13 mm, densely covered (hke axes of cymes 
and peduncles) with glandular hairs; calyx usually 5 (4.5-6.5) mm long, green or violet, 
pubescent with fine straight spreading hairs and more or less densely covered with sessile 

384 or stipitate capitate white glands interspersed with sessile round yellow glands; upper 
teeth triangular or oblong-triangular, acute or acuminate, 0.9-1.3 mm long, 1/4-1/3 as 
long as tube, the lower 2 slightly longer (1-1.25 mm) and narrower, narrowly lanceolate, 
long-acuminate; corolla (5) 5.5-7 mm long, blue or white, only slightly exserted (the up- 
permost calyx-tooth reaching the middle and sometimes even the apex of upper Up), tube 
slender, curved; upper lip erect, 1-1 .2 mm long, cut to middle into semiorbicular lobes; 
lower lip nearly twice as long as the upper, the large-toothed middle lobe ca. 1.5 mm long, 
2-2.5 mm wide, the lateral lobes broadly triangular, 0.4-0.5 mm long, 1 mm wide; upper 
stamens as long as upper Up of corolla; nutlets oblong-ovoid, 1.5-1.7 mm long, 0.6 mm 
wide, with an obtuse edge on ventral side, dark brown, foveolate-tuberculate. Fl. May- 
June; fr. from second half of May. 



257 



Dry steppes and edgesofdeserts,mainly on sandy soils, sandy hillocks, barkhans, saline 
sands, saxaul associations, foothills, rarely in lower mountain belt. - West Siberia: Irt. 
(Ulu-Tau mountains); Centr. Asia: Ar.-Casp. (except for southwestern part), Balkh., Kyz. 
K. (eastern part), Syr D., T. Sh.: foothills and in spruce belt along Naryn River, intro- 
duced?, Pam.-Al. (Zeravshan valley). Gen. distr.: Dzu.-Kash. Described from Arkaul 
mountains in E. Kazakhstan. Type (lectotype) in Leningrad. 

53. N. meyeri Benth. Lab. gen. et sp. (1834) 478; in Dc. Prodr. XII (1848) 384; Hohe- 
nack. Enum. pi. talysch, 69; Ldb. Fl. Ross. Ill, 375. - N. micrantha auct.: C. A. M. 
Verzeichn. Pfl. Cauc. kasp. Meer. (1831) 92; Bge. Lab. pers. 56, p.min. p.; Boiss. Fl. or. 
IV, 664 (excl. syn. C. Koch); N. Pop. in Tr. Bot. sada Yur'evsk. univ. XIV, 234; Grossg. 
Fl. Kavk. Ill, 291, p.p.; idem,Opred. rast. Kavk. 330, p. p. non Bge. (1830). - Exs.: 
Herb. fl. Cauc. No. 186. 

Annual, with slender vertical root; stems (3) 10-45 cm long, densely covered with sim- 
ple white hairs, sometimes interspersed in upper part with subsessile whitish glands, simple 
or with 1 or 2 pairs of leafy inflorescence-bearing branches in axils of lower leaves and 
cymes in all other axils; leaves tougher than in N. micrantha Bge., the secondary and 
often even tertiary veins thicker, impressed above, prominent beneath, the upper side pale 
green, with sparse, rarely dense, appressed hairs, the lower side glaucescent to whitish, with 
more copious indument of jointed simple hairs interspersed with very small glandular hairs 
and numerous punctate glands; lower and middle leaves broad-ovate to oblong-ovate, 0.8- 
385 3.5 cm long, 0.4-1 .5 cm wide, with cordate, truncate or cuneate base, obtuse or acute, 
always dentate (teeth 4-8 pairs, obtuse or acute), the upper lanceolate to lanceolate-linear, 
usually with few teeth, rarely entire, often as long as peduncles of cymes or slightly shorter; 
petioles grayish, with profuse spreading simple hairs, the lower longer than blade, the up- 
per shorter; cymes compact, forming a narrow, usually rather dense raceme; upper cymes 
3 (the uppermost)-6-8-flowered, sessile or on peduncles 3-10 mm long, in pairs (2) 3- 
10mm apart, the others 6-12 (16)-flowered, with longer peduncles (2-3.5 cm), 2-5 cm 
apart, dense, with axes of first order to 5 mm, rarely the lower distinctly dichotomous, 
with axes 10-1 5 mm long; peduncles, axes and pedicels densely white-pubescent, with crisp 
hairs sometimes interspersed with very small whitish sessile or short-stipitate glands; pedi- 
cels 0.8-2.5 (3) mm long, to 10(15) mm long in bifurcation of cymes; bracts linear-subu- 
late, long-cihate, 2-5 mm long; calyx (5.5) 6-8.5 mm long, narrowly tubular in flower, 
oblong-ovoid in fruit, green with white corolla or more or less violet to dark violet with 
blue corolla, densely covered with long spreading hairs and with numerous minute capitate 
whitish glands and sparse punctate yellow glands; upper teeth narrowly lanceolate, acumi- 
nate; middle tooth slightly longer than lateral teeth, 1/3-2/5 as long as tube; lower teeth 
narrower, lanceolate-subulate, long-acuminate; corolla 7-8.5 (9.5) mm long, blue, rarely 
white or pink, the slender curved tube abruptly expanding into neck 2 mm long and ca. 
2.5 mm wide; tube (and partly neck) concealed in calyx, the limb completely exserted 
(the largest uppermost calyx-tooth reaching the base of upper Hp); upper lip 1 .5-2.5 mm 
long, cut nearly to the middle into semiorbicular lobes; lower lip with large-toothed mid- 
dle lobe 2-4 mm long and 4-4.5 mm wide, the lateral lobes semiorbicular-triangular, 0.6- 
1 mm long, 1 .5-2.3 mm wide; nutlets oblong-ellipsoid, 1 .3-1 .5 mm long, 0.75-0.85 mm 
wide, dark brown, with an obtuse edge on ventral side, foveolate, with flat tubercles. Fl. 
(March?) April to second half of July; fr. from second half of April. 



258 



Desert lowlands in Kura and Araks river valleys, also in lower and central mountain 
belts on exposed southern slopes. — Caucasus: E. Transc. (Kura valley), S. Transc, Tal. 
Gen. distr,: Arm.-Kurd. Described from Zuvant in Tal. Type in London; type duplicate 
in Leningrad. 

Note. Bentham first distinguished the Caucasian Nepeta, included by C. Meyer in 
his list of plants of the Caucasus (1831) under the name "M. micrantha," from specimens 

386 of the true N. micrantha Bge. known to him from the western foothills of the Altai, 
and he described it as a separate species. However, later authorities, including Bunge, did 
not recognize the specific status of N. meyeri Benth. and again combined it with N. mi- 
crantha Bge. The herbarium material that has since accumulated for both species fully 
confirms the need to separate N. meyeri Benth. It is readily distinguishable from N. mi- 
crantha Bge. by its larger flowers, the partly or fully exserted neck of the corolla, the more 
compact cymes with a larger number of flowers, the relatively tough and nearly always 
toothed leaves with more prominent veins, and glandular hairs on peduncles, inflorescence 
axes and petioles less developed or often absent. 

54. N. pallida C. Koch in Linnaea, V (1848) 675. - N. micrantha auct.: Boiss. Fl. or. 
IV (1879) 664, p. p.; Grossg. Fl. Kavk. Ill, 291, p. p.; idem, Opred. rast. Kavk. 330, p. p. 
non Bge. nee C. A.M. — N. bordzilovskii Tarn, et An. Fed. in Grossg. Opred. rast. Kavk. 
(1949) 330, ross. 

Annual, branching, 3-25 cm high; stems scaberulous with very short simple white hairs 
sometimes intermixed with sessile or subsessile whitish glands; branches in axils of 2 or 
3 lower pairs of leaves, usually nearly as long as stem, fertile; leaves pale, grayish-green, 
the rather thick veins impressed above, prominent beneath, the lower side rather densely 
covered with short capitate glandular hairs interspersed on the veins with longer simple 
jointed hairs and with numerous sessile round yellow glands, the upper side with similar 
but sparser indument; lower leaves 7-18 mm long, 5-12 mm wide, ovate, with rounded 
or cuneate or sometimes cordate base, acute or obtuse, the margin with obtuse or acute 
triangular teeth; middle and upper leaves usually narrower, oblanceolate or oblong-eUiptic, 
acuminate, gradually smaller and more bracteiform toward summit; floral leaves subtend- 
ing lower cymes usually slightly shorter than peduncles, the middle usually longer and 
sometimes exceeding the whole axillary inflorescence; cymes dense, compact, 6 (the up- 
permost)-8-26-flowered, in contracted raceme, the upper sessile or subsessile, in closely 
approximate pairs, the others 1-2 (3) cm apart, on 3-13 mm long peduncles; axes of first 
order very short, hence cymes very compact, often resembling semiverticels; pedicels 1.5- 
3 mm, in bifurcation 3-5 mm long; bracts narrowly linear or subulate, 2-3 mm long; in 

387 addition to individuals with bisexual flowers, there was one specimen with female flowers; 
calyx 3.6-5 (6) mm long, in flower narrowly tubular, in fruit oblong-ovoid; green (in white- 
flowered form — N. bordzilovskii Tarn, et An. Fed.) or violet (in typical form with blue 
flowers), densely covered outside with small sessile and subsessile whitish glands and spread- 
ing simple hairs; upper teeth oblong-triangular to lanceolate, acuminate, the middle tooth 
distinctly larger than lateral teeth, 1/3-1/2 as long as tube, the lower teeth with broad 
base, sharply tapering to a long point; corolla pubescent outside, very hairy on lips, 3.8- 
5.2 (6.2) mm long, nearly concealed in calyx (uppermost tooth reaching the middle or 
nearly the apex of upper lip, others reaching the base of lower lip); tube slender, curved, 



259 



abruptly expanding into neck; upper lip shallowly cut into semiorbicular lobes; lower lip 
wider than long, concave, with large-toothed margin, the lateral lobes triangular, obtuse; 
upper stamens in bisexual flowers as long as upper hp of corolla; style as long as or shorter 
than upper stamens; female flowers with 4 staminodes in dilated part of tube, these con- 
sisting of very short filament and rounded scarious sterile rudiment of anther; style longer 
than upper lip of corolla; lobes of stigma large, more than 1 mm long, thick, twisted; nut- 
lets eUipsoid, ca. 1 mm long, 0.8 mm wide, brown, tuberculate-foveolate. Fl. May-July; 
fr. June. 

Steppe and semisteppe plant communities in lower and central mountain belts. - Cau- 
casus: S. Transc. (W. Armenia: vicinity of Echmiadzin, Leninakan). Endemic? Described 
from a basalt outcrop in former Erevan province. Type was in Berlin. 

Notes. 1) Specimens from the vicinity of Echmiadzin and Leninakan differ markedly 
from N. meyeri in having much smaller flowers, almost completely concealed in the 
calyx, very compact many-flowered cymes, much more contracted inflorescence, an alto- 
gether distinctive habit, and profuse glandular hairs on all parts, including those that are 
often eglandular in N. meyeri (petioles, peduncles, axial parts of cymes). From the area 
of distribution of this unique Nepeta, C. Koch described N. pallida C. Koch, closely 
allied to N. meyeri. He note some features of N. pallida, e.g. the very dense, many- 
flowered cymes, which also characterize the above-mentioned specimens from the western 
part of the Armenian SSR, presented here under the name N. pallida Koch. 

2) N. bordzilovskii Tam. et An. Fed. is a white -flowered form of N. pallida C. Koch; 
other distinctions have not been found. White flowers are fairly often observed also in 
other species of the series Micranthae. 

3) Worthy of note is the occurrence of gynodioecism in N. pallida, as it has never be- 
388 fore been recorded for the section Micranthae. The female flowers are smaller than the 

bisexual, with scarcely exserted corolla; they form denser, subglobose lower whorls. 

55. N. saccharata Bge. Lab. pers. (1873) 56; Boiss. Fl. or. IV, 664. - N. meyeri var. 
laxiflora Kotschy. PI. Pers. bor. exs. No. 175. 

Annual, with slender vertical root; stem 20-70 cm long, erect, sparsely covered with 
very short glandular hairs, simple or in lower part with 1-3 pairs of mostly short fertile 
branches; leaves thin, pale green, slightly glaucescent beneath, covered (somewhat more 
densely beneath) with scattered short appressed simple hairs and sessile capitate whitish 
glands as well as a few punctate yellow glands, the veins not prominent; lower and middle 
leaves ovate, with cordate or truncate base, obtuse at apex, with 6-8 pairs of large obtuse 
teeth, 1.5-3.3 cm long, 0.8-1.8 (2.3) cm wide; upper leaves lanceolate, acuminate, usually 
with acute teeth, gradually smaller and narrower toward summit, subUnear and often en- 
tire; lower petioles half as long again to twice as long as blade, the middle short; upper 
leaves sessile; petioles densely covered with fine capitate glands, in upper part also with 
long spreading simple hairs; cymes loose, few-flowered, forming a lax raceme; upper 
cymes 3-5-flowered, unbranched, in subremote pairs, the peduncles 3-10 mm long, usual- 
ly shorter than bracts; other cymes 5-1 1 (13)-flowered in pairs 3-9 cm apart, on pedun- 
cles to 4.5 cm long, loosely dichotomous or trichotomous, with axes of first order 5- 
15 mm long, subtended by narrowly linear floral leaves 6-12 mm long; pedicels in termi- 
nal cymes 0.6-1 .5 mm long, in lower cymes to 5 mm, in bifurcation to 12 mm; all axial 



260 



parts of inflorescence finely glandular-hairy; bracts linear-subulate, 3-5 mm long, with 
sparsely ciliate margin; calyx (4) 5.5-7 mm long, narrowly tubular in flower, ovoid in 
fruit, covered outside with fme (usually sparse) whitish capitate glands interspersed with a 
few round yellow glands and with spreading simple white hairs, these sometimes very dense 
in lower part; calyx incised in front to middle or beyond, rarely less; upper teeth ovate- 
triangular or triangular, abruptly attenuate to a rather long point, the middle tooth larger 
than lateral teeth, 1 .5-1 .8 (1 .9) mm long, the lower teeth narrowly lanceolate, gradually 
long-acuminate, exceeding the upper, 2-2.5 mm long; corolla 7-8.5 mm long (pinkish-lilac 
389 or lilac-blue, with dark purple spots on lower lip?), the slender tube gradually expanding 
into neck 1-2 mm long and 2-2.5 mm wide, the limb completely exserted, the neck enclos- 
ed by calyx-teeth to about the middle; upper lip 1.2-1.5 mm long, shallowly cut into ovate 
lobes; middle lobe of lower hp ca. 3 mm long, 5 mm wide, the lateral lobes rounded, ca. 
0.3 mm long, 1 .5 mm wide; nutlets ellipsoid-obovoid, 1.8 mm long, 1 mm wide, pale 
brown, coarsely tuberculate, the tubercles flat, yellow. Fl. May -June; fr. June-August. 
(Plate XXIII, Figure 2.) 

Lower and central mountain belts; gravelly and sandy slopes; pebble beds of mountain 
streams. — Centr. Asia: Kopet Dagh, apparently only in its central part (ChuH, Saratovka). 
Gen.distr.: Iran, (northern part): Elburz, Siakukh, Kokhrud mountains. Described from 
N. Iran, Paskalekh near Teheran. Type in Leningrad. 



Series 3. Ispahanicae Pojark. — Flowers small; calyx 4-6 mm long, straight, with 
frontal incision \ 12-213 its length, the upper teeth 2/5-2/3 as long as tube, the lower 
teeth subulate; corolla 4.8-7.5 mm long, included mostly to limb (in one species to 
dilated part of tube); nutlets with finely tuberculate or granular surface; verticillasters 
subsessile, crowded at the end of elongate terminal internode of stem and axillary branches, 
forming a small but many-flowered inflorescence, this sometimes interrupted below; bracts 
lanceolate, flat, as long as calyx. 

Besides the two Central Asian species, this series should also comprise N. schirasiana 
Boiss. (SW Iran), N. eremophila Hausskn. et Bornm. (SE Iran), and N. kojeana Rech. 
fil. (western part of central Iran). 

56. N. ispahanica Boiss. Diagn. Ser. 1, 5 (1844) 23; id Fl. or. IV, 666; Benth. in DC. 
Prodr. XII, 394; Bobrov in Tr. Bot. sada, XLIV(1931) 72. 

Annual, 5-15 cm high; stem erect or ascending, more or less densely covered with long 
fine crisp spreading white hairs and densely beset with very fine short-stipitate or sessile 
capitate glands (sometimes only glandular), simple or branching at base, branches all with 
inflorescences, often nearly equaling the stem; cauline leaves mostly on lower part of 
stem, 1-2 (2.8) cm long, 0.5-2.4 (2.8) cm wide, ovate or cordate, obtuse to rounded at 
apex, coarsely crenate, densely grayish-lanate beneath, less densely so to glabrescent above; 
petioles as long as blade, patent-hairy, short-glandular, lower floral leaves as long as semi- 
390 verticels, subsessile, ovate, acute, the upper oblong-eUiptic to linear-lanceolate, bracteiform; 
verticillasters few-flowered, subsessile, the upper forming a terminal inflorescence, this 
dense, capitate, subglobular, rarely cyUndrical, 1 .8-2.3 cm long, 1-2 mm wide; stem and 
stronger axillary branches with 1 or 2 remote pairs of looser few-flowered semiverticels 



261 



below terminal head; bracts strict, linear, acuminate, equaling calyx, densely ciliate, with 
long fine implexed hairs; pedicels 1.5-3 mm long, elongating in fruit to 5-8 mm; calyx 
4-5 (5.5) mm long, cylindrical in flower, campanulate in fruit, violet, glandular and thinly 
patent-hairy outside, the throat slightly oblique; teeth erect, long-ciliate, covered inside 
with long appressed hairs, the upper 1.3-1.7 mm long, oblong-triangular, acuminate, (2/5) 
1/2 as long as tube, the lower 2.2-2.8 mm long, much narrower, hnear-lanceolate, the inci- 
sion between them reaching the middle of calyx; corolla 4.8-6 (6.5) mm long, included 
up to the Hmb, hairy outside; tube slightly curved, 2.8-4 mm long, rather abruptly ex- 
panding into infundibular neck 1.0-1.3 mm long and 1.2-1.5 mm wide; upper lip 1- 
1.25 mm long, erect, deeply cut into ovate concave lobes; lateral lobes of lower lip hori- 
zontally spreading, very small (ca. 0.3 mm long, 0.7 mm wide), obliquely semiorbicular, 
the middle lobe much larger, 1-1.5 mm long, 1.5-2 mm wide, with cordate base, entire or 
undulate-margined, concave, shallowly emarginate; upper stamens shorter than upper lip 
of corolla; nutlets 1.2-1.3 mm long, 0.6-0.7 mm wide, ellipsoid, brown, with numerous 
very small and nearly flat tubercles, the ventral side with an obtuse edge. Fl. (April?) 
May-June; fr. from first half of June. 

Foothill and low mountain belt on fine earth (including saline) and gravelly slopes. — 
Centr. Asia: Mtn. Turkm. (Greater Balkhan Range). Gen. distr.: Iran. Described from 
vicinity of Isfalian in Iran. Type in Geneva, type duplicate in Leningrad. 

57. N. santoana M. Pop. in Tr. Turkest. Gos. univ. 4 (1922) 57, 64. 

Annual, with slender vertical root; stems 5-20 cm long, erect, robust, stout, sometimes 
simple but mostly branching at base, grayish-pubescent with rather long fine jointed simple 
hairs; branches arched-ascending, terminating in inflorescence, often equaling the stem in 
length and diameter; cauHne leaves 2 or 3 pairs, very thin, the upper side smooth, green- 
ish, rather densely covered with appressed simple hairs and with sparse round yellow resi- 
391 nous glands, the lower side with prominent main veins, grayish with similar but denser 
indument; lower and middle leaves 1-2 cm long and as wide, broadly ovate, with cordate 
base slightly decurrent on petiole, obtuse or rounded at apex, coarsdy crenate, petioles of 
lower leaves as long as blade, those of middle leaves shorter; flowers in subsessile few- 
flowered semiverticels, these in 3-4 pairs at the ends of stem and branches; inflorescence 
dense, spicate-capitate, slightly one-sided, 10-25-flowered, 2-2.3 cm long, 1-1.5 cm wide, 
cylindrical or ovoid, 1 (2) remote verticillasters sometimes borne below inflorescence; 
floral leaves at base of the head ovate, resembling cauline but with acute teeth, rounded 
at base, smaller, to 1 .4 cm long and as wide; floral leaves subtending semiverticels bractei- 
form, 5-7 mm long, Unear-subulate, acuminate, curved, the margin sparsely long-ciliate; 
bracts usually slightly shorter than calyx; pedicels 1-1.5 (2) mm long, sturdy, stoutish; 
calyx 4.5-5.2 mm long, with 13 thick nerves, with yellow, sessile, resinous glands and with 
jointed hairs mainly on nerves, straight, cyUndrical in flower, ovoid in fruit, with slightly 
obhque throat, the 3 upper teeth distinctly shorter than the lower, 1-1.5 mm long, erect, 
oblong-triangular to sublanceolate, 1/4-1/3 as long as tube, the 2 lower teeth slightly 
curved, narrowly lanceolate, 1 .3-1 .8 mm long; calyx-tube with triangular acute incision 
2/5-1/2 its length; all teeth acuminate, glabrous inside, cili?te; corolla white, pubescent 
outside, 7-7.5 mm long, included to middle or nearly to limb, the curved tube gradually 
expanding into narrow neck; shape and position of lips and their lobes as in the preceding 



262 



species but the middle lobe of lower lip large-toothed at margin; nutlets 1 .5-1 .7 mm long, 
0.8-0.9 mm wide, oblong-ellipsoid to obovoid, brown, densely tuberculate, the tubercles 
very short and almost completely flat. Fl. and fr. in first half of June. (Plate XXV, 
Figure 1.) 

Desert hills with outcrops of gypsiferous limestone. - Centr. Asia: Pam.-Al. Described 
from the Santo oil fields near Sary-Tau Range in Uzbekistan (the only location). Type 
(or its duplicate?) in Leningrad. 



Section 7. Schizocalyx Pojark. sect. nov. in Addenda XIX, 358. — Sect. Macronepeta 
Benth. Lab. gen. ex sp. (1834)467,482; in DC Prodr. XII, 387, p.p. -Sect. Eunepeta § 
Longiflorae Boiss. p. p. (quoad descr. sed excl. N. longiflora Vent.); Briq. in Pflanzen- 
fam. IV, 3a, 236, p. p. — Calyx narrowly tubular, curved, with strongly oblique or 2-lipped 
throat, deep and broad frontal incision and short teeth; corolla-tube long, abruptly ex- 
392 panding into short and broad neck; narrow part of tube IVz-S times as long as calyx; mid- 
dle lobe of lower lip concave, coarsely crenate; bracts much shorter than calyx; nutlets 
smooth or with minute smooth tubercles (strong magnifying glass!); flowers in remote 
cymes or verticillasters, the upper of which sometimes crowded in capitate inflorescence. 
Perennial, mostly strongly branching plants; leaves cordate or triangular, small, 2-3 (4) cm 
long. Type of the section: N. fissa C. A.M. 

About twenty species distributed in the Caucasus, Iran and the eastern part of Asia 
Minor. 

Note. This section was accepted here in the same scope as the subsection Longiflorae 
Boiss. but, as N. longiflora Vent. (i.e. type species) had to be excluded, the rise in rank 
necessitated a change of name. The drawing accompanying the first description of N. lon- 
giflora Vent. (Hort. Cels. 66, tab. 66) leaves no doubt that this species has very Httle in 
common with any other species included in the section bearing its name (cf. inflorescence, 
shape of calyx and leaves!). The herbarium specimen of this species in the Botanical Insti- 
tute of the Academy of Sciences of the USSR, which was obtained from Olivier in Paris in 
1804 (i.e. presumably a type duplicate), fully supports this approach and provides evidence 
that this species is very closely related to the cycle N. mussinii Henck.— N. transcauca- 
sica Grossh. 



Series 1. Longitubae Pojark. — Flowers in few-flowered, simple cymes (the upper 
sometimes in semiverticels) of which the upper 2-4 pairs sessile, crowded in a head, the 
others remote with peduncles 2 cm long; corolla 20-33 mm long, the strongly curved tube 
2-3 times as long as calyx; nutlets scabrous or with minute flat tubercles; stems simple or 
slightly branching. Plants of lower and middle mountain belts. 

In addition to the two Caucasian species, this series should also include N. macrosi- 
phon Boiss. (SW Iran), N. pulchra Pojark. (S. Kurdistan) and N. pycnantha Benth. 
(Taurus Mountains in Asia Minor). 

58. N. longituba Pojark. in Bot. mat. gerb. Bot. inst. ANSSSR, XV (1953) 318. 
Perennial; stems arched-ascending, 30-60 cm long, 2.5-3.5 mm across (at base), simple 
or branching, rather densely covered with sessile or short-stipitate whitish glands and. 



263 



mainly in lower part, with spreading simple jointed hairs, leafy in lower part where nodes 
2-3 cm apart; middle internodes 7-10 cm long; cauline leaves broadly cordate, rounded 

393 at apex, with 6-9 large obtuse teeth per margin, 1-2.4 cm long, 1-2.8 cm wide, pale green, 
unicolored, densely covered above with fine appressed hairs, the lower side with scattered 
minute sessile whitish glands, the veins or the whole surface villous with 1-3-jointed simple 
hairs; lower petioles 234 times as long as blade, the upper leaves 1/2-2/3 as long; floral 
leaves, except the lowermost, bracteiform, 3-5 mm long, ovate-lanceolate or lanceolate, 
the upper usually violet-tinged; cymes 3-5-flowered, compact, not dichotomizing, the 
upper sessile or on very short peduncles, forming a (6) 10-12-flowered capitate intlores- 
cence at summit of stem, the lower on peduncles 1.5-1.8 mm long, remote; bracts ovate- 
lanceolate, rather abruptly attenuate to a short point, the margin beset with white jointed 
hairs; pedicels 0.5-2 mm long; calyx 7-9 mm long, violet, 15-nerved; tube densely cover- 
ed with sessile yellowish glands and on nerves with spreading white 2-3-jointed hairs, the 
frontal incision 1/2-2/3 its length; teeth broadly scarious-margined, the 3 upper broadly 
ovate, 1-1.6 mm long, abruptly short-pointed, 1/6-2/9 as long as tube, the 2 lower longer 
and narrower, ovate-lanceolate or ovate, 1 .7-1 .8 mm long; corolla (20) 22-27 mm long, 
blue, the slender tube 3-3^/d as long as the limb and 2]6. times as long as calyx; upper Up 
ca. 2 mm long, cleft to 2/3, the middle lobe dilated, the lateral lobes 0.7-0.8 mm long, 
2.5-3 mm wide; nutlets elUpsoid, 2.3-2.5 mm long, 1.3-1.5 mm wide, 3-angled. brown, 
with minute flat tubercles visible under high magnification. Fl. and fr. first half of 
August. (Plate XXI, Figure 2.) 

SHding taluses in central mountain belt. — Caucasus: E. Transc. (Nukhi; Geok-Chai 
River; Ismailly). Endemic. Described from northern part of Azerbaidzhan near Upper 
Dashagil village, not far from Nukhi River. Type in Leningrad. 

59. N. sosnovskyi Asker. in Bot. mat. gerb. Bot. inst. AN SSSR, XVI (1954). 

Perennial; root obhque, fibrous, imperceptibly passing above into a very short rhizome, 
this bearing a few closely approximate brown squamiform leaves ; stem 25-3 5 cm long, soh- 
tary, ascending, straight or sUghtly curved, with horizontal brown base buried in rubble, 
violet-tinged above, the lower internodes 1.5-2 cm, the middle 5-8 mm long; branches in 
axils of lower leaves sterile, leafy; 2-3 pairs of axillary branches fertile, with 1 or 2 pairs 

394 of remote, small leaves and inflorescence at the end; stems, branches and petioles beset 
with very short sessile or short-stipitate whitish capitate glands and short white spreading 
simple proximally thickened hairs; leaves firm, glaucescent, with prominent main veins 
on lower side, sometimes violet, densely covered on both sides with simple jointed hairs, 
these fine and appressed above, spreading and coarser beneath; lower cauhne leaves 1 .3- 
2.3 cm long, 1 .8-2.3 cm wide, cordate, with deeply incised base and rounded apex, the 
margin with 5-6 large rounded crenae, the petioles approximately as long as blade; upper 
leaves subsessile, ovate, more finely crenate; floral leaves sessile, ovate, mostly entire, 
acute, the uppermost bracteiform; inflorescence consisting of a dense terminal head 2- 
3.5 cm long, 3-3.5 cm wide, and sometimes a remote pair of 3-5-flowered short-peduncled 
cymes; inflorescences on axillary branches terminal heads not smaller than on stem or 
more often composed of 1 or 2 verticillasters; bracts narrowly eUiptic to Unear-lanceolate, 
4-7 mm long, long-acuminate, terminating in a rigid point; calyx 9-12 mm long, violet, 
densely covered with yellow sessile glands, the nerves with scattered spreading 2-3-jointed 



264 



white simple hairs, the frontal incision reaching the middle; upper teeth ovate or ovate- 
triangular, 1.5-2.5 mm long, acute, 2/11-1/4 as long as tube, the lower oblong-lanceolate, 
acuminate, 3-3.25 mm long; corolla 21-27 mm long, the slender tube exserted more than 
half its length, the neck 3-4 mm long; upper lip ca. 3 mm long, cut to the middle into 2 
porrect lobes; middle lobe of lower Up 5-6 mm long, 8-9 mm wide, with deeply cordate 
base, the lateral lobes 0.4-0.5 mm long, 2.5-3 mm wide; nutlets ellipsoid, attenuate at 
both ends, 2-2.5 mm long, 1 mm wide, brown, nearly smooth. Fl. (June) first half of 
July; fr. from August. 

Pebbles in lower mountain belt. — Caucasus: E. Transc. (northeastern part). Endemic. 
Described from vicinity of Alty-Agach village, Mutyanka River, Khizinsk district in E. Azer- 
baidzhan (so far known only from this locaUty). Type in Baku. 

Cycle 1. Fissae Pojark. — Flowers in remote cymes, these simple or once or twice 
(thrice) loosely dichotomously branching, with peduncles 0.4-1 (the upper) to 4.5 cm 
long; aggregate inflorescence a very sparse panicle, rarely a raceme; corolla 11-20 cm 
long, the tube Wi-l (IVi) times as long as calyx; nutlets with very numerous minute 
395 (strong magnifying glass!) flat pellucid tubercles. Plants of middle and lower mountain 
belt covering the Caucasus and Southwest Asia (from Iran to eastern part of Asia Minor). 

60. N.fissaC.A.M. Verzeichn.Pfl. Cauc. kasp. Meer. (1831) 93; Benth. Lab. gen. et 
sp. 737 (non471). — N. rariflora C. A.M. in sched. non C. Koch. — N. laxa Benth. 1. c. 
483 (excl. syn.). - N. teucriifolia auct.: Ldb. Fl. Ross. Ill (1846-1851) 378; Benth. 
in DC Prodr. XII, 389, p. p.; Lipsk. Fl. Kavk. 420, p. p.; N. Pop. in Tr. Bot. sada Yur'evsk. 
univ. XIV, p.p.; Grossg. Fl. Kavk. Ill, 254, p.p.; idem,Opred. rast. Kavk. 331, p.p., non 
Willd.(1809). 

Perennial; root woody, coarsely fibrous; stems 50-90 cm long, with short simple hairs, 
densely leafy in lower part, with middle internodes 5-8 (1 1) cm long, strongly branching, 
mostly from base; axillary branche's long, with few pairs of cymes; cauline leaves ovate- 
triangular or oblong-ovate to lanceolate, with truncate or shallowly cordate base, obtuse 
or acute, each margin with 6-7 large obtuse or acute teeth, 1 .5-4 cm long, 1 .2-3.5 cm 
wide (the uppermost at base of upper axillary branches much reduced, entire, sublanceo- 
late), the upper side densely covered with very short and very fme appressed hairs, finally 
glabrescent, the lower side at first with usually dense indument of equally short but 
coarser spreading hairs, these sometimes persistent on the veins; lower petioles half as 
long again as the blade, the upper 1/5-1/3 the length of blade; floral leaves, except some- 
times the lowermost, small, narrow, entire, bracteiform; aggregate inflorescence (including 
inflorescences on axillary branches) paniculate, very lax, to 2/3-3/4 of whole plant length; 
upper cymes 3-7-flowered, sometimes unbranched, the others loosely dichotomous with 
axes of first order sometimes developing into elongate bostryx, 1-4 cm long; peduncles of 
cymes slender, divergent to horizontal, the upper 1-1.5 cm long, the others to 5 cm; bracts 
1 .5-2 mm long, narrow, acuminate, mostly violet; pedicels 0.8-1 mm, that of central flow- 
er to 1 .5 mm long; calyx (6.5) 7-8 mm long, incised nearly to the middle, often nearly to 
1/3, densely covered with sessile yellowish glands and spreading (2) 3-jointed hairs, the 
upper teeth (1/6) 2/11-2/9 as long as tube (1-1.3 mm long), ovate or ovate-lanceolate, 
abruptly attenuate to a short point, the lower 1.3-1.5 mm long, lanceolate, short-acuminate; 



265 



corolla blue, 17.5-19 mm long, the tube exserted from calyx to half its length or slightly 
more, expanding into neck 3-4 mm long and as wide; upper Up ca. 2.5 mm long, cleft to 

396 2/3-3/5 into obtuse lobes; middle lobe of lower lip 3-3.8 mm long, 5.5-6.5 mm wide, the 
lateral lobes 0.7-1 mm long, 1-1 .2 mm wide; nutlets ellipsoid, 3-angled, ca. 2 mm long, 

1 mm wide, very finely punctate-tuberculate. 

Exposed dry, gravelly and stony places in lower and central mountain belt. - Caucasus: 
S. Transc. (southwestern part of Azerbaidzhan, Dzhebrail and Gadrud areas), Tal. Gen. 
distr.: Iran. (Astrabad province, Elburz Range). Described from near Sivirs in Talysh, 
Type in Leningrad. 

Notes. 1) This species was reported by Karelin (1839) for "Northern Turkmenia" and 
there is a herbarium specimen with similar inscription; but to this day nobody else has 
found the plant in Turkmenia and it must be assumed that it does not occur there. 

2) Species most closely related to N. fissa C.A.M. are N. microphylla Stapf (C. Iran) 
and N. szovitsiana Pojark. (N. longiflora Boiss. p.p. non Vent.) from N. Iran 
(Kurdistan). 

61. N. trautvetteri Boiss. et Buhse in Nouv. Mem. Soc. Nat. Mosc. XII (1860) 175; 
Boiss. Fl. or. IV, 647, p. p. (quoad typum). - N. leptoclada Trautv. in Tr. Bot. sada, II 
(1873) 480; Boiss. 1. c, 648; N. Pop. in Tr. Bot. sada Yur'evsk. univ. XIV, 234; Grossg. 
Fl. Kavk. Ill, 294; idem,Opred. rast. Kavk. 331. - Exs.: PI. or. exs. No. 216. 

Perennial; root tough, woody, not forming a rhizome; stems few, 45-100 cm long, 2- 
5 mm thick, usually branching, mostly from base, beset (like all other parts of the plant) 
with small yellowish capitate sessile glands, leafy in lower part and densely covered with 
coarse subsetaceous white spreading hairs; axillary branches long, leafless, with few pairs 
of cymes; cauline leaves 1.5-4.5 cm long, 1.2-4 cm wide, broad-ovate to suborbicular- 
ovate, rounded or obtuse at apex, deeply cordate at base, each margin wiih 6-9 large crenae, 
the upper side with rather dense but inconspicuous indument of rather long, very fine, 
more or less implexed hairs, the lower side at first covered all over with long white fuga- 
cious hairs, the veins with mostly persistent subsetaceous hairs; petioles white-hispid, the 
lower about half as long again as blade, the upper short; floral leaves small, sessile, oblong- 
ovate to narrowly elliptic, entire or the lower with few teeth; aggregate inflorescence a 
large, broad, loose panicle; cymes remote, all except the upper dichotomously branching, 
the axes of first order to 2 cm long, sometimes repeatedly dichotomous or bostryciform; 
bracts ovate-elliptic or eUiptic, 2-3 mm long; pedicels 0.7-1 mm long; calyx 6-7.5 mm 
long, violet, densely covered with whitish or yellowish sessile glands and sparse spreading 

397 (2) 3-jointed simple hairs, frontally incised to the middle; teeth with scarious margin, the 
upper 0.8-1 mm long, ovate or broadly ovate, 2/15-1/6 times as long as tube, mucronate, 
the lower 1-1.5 mm long, lanceolate or narrowly lanceolate, acuminate; corolla 15.5- 

17 mm long, the slender curved tube exserted to the middle, abruptly attenuate [?] into 
neck 2.5-3 mm long and wide; upper lip 2.3-3 mm long, cleft to 1/2-2/3; middle lobe of 
lower Up 3-4 mm long, 4-6 mm wide, with large triangular teeth at margin, the lateral 
lobes 0.7-1 mm long, 2-3 mm wide; nutlets brown, 1.8-2 mm long, 1 mm wide, ellipsoid, 
minutely tuberculate, the tubercles rather pale and very flat. Fl. May-June; fr. second 
half of June to July. 

Dry exposed slopes in middle mountain belt. - Caucasus: S. Transc. (Nakhichevan 



266 



ASSR and SE Armenia). Gen. distr.: N. Iran (Karadag). Described from near AUiar in 
N. Iran. Type in Leningrad. 

Note. A comparison of the type specimens of N. trautvetteri Boiss. et Buhse and 
N. leptoclada leaves no doubt that they are identical. Yet, with the exception of Popov 
(1914), nobody has compared them as related species. This is due to the fact that N. 
trautvetteri has so far been Hsted by all investigators of the flora of the Caucasus as 
N. leptoclada Trautv. (described from Bilyav village in Nakhichevan ASSR) v^hereas the 
name N. trautvetteri has been associated with a species differing iharkedly from that to 
which it had been originally applied — N. velutina Pojark. (see Note to this species). 

62. N. velutina Pojark. in Bot. mat. gerb. Bot. inst. AN SSSR, XV (1953) 3 12. - N. 
trautvetteri Boiss. Fl. or. IV (1879) 647, p. p. (quoad pi. e Koschadara), non Boiss. et 
Buhse (1860); auct. fl. Cauc: N. Pop. in Tr. Bot. sada Yur'evsk. univ. XIV, 234; Grossg. 
Fl. Kavk. Ill, 294, p. p.; idem, Opred. rast. Kavk. 331, p. p., non Boiss. et Buhse (1860). 

Perennial; root woody, vertical, 2 cm across in upper part; stems 1-4, 30-120 cm long, 
2-8 mm thick at base, densely velutinous, the very short simple (eglandular) hairs forming 
a grayish powdery bloom, with internodes in middle part 5-10 cm long, branching from 
base but sometimes simple; branches usually long, leafy, terminating in racemiform or 
paniculate inflorescence; leaves broadly ovate, with truncate or cuneate or rarely cordate 
base, obtuse at apex, with 5-9 large crenae per margin, the upper side bright green, at first 
rather densely hairy, the hairs appressed, very fine, discernible only under high magnifi- 
cation, later sometimes almost completely disappearing, the lower side gray, velutinous 
398 all over; cauline leaves 1 .7-5 cm long, 1 .3-4.3 cm wide; petioles IVi-l times as long as 
blade, the upper 1/4-2/5 as long as blade; floral leaves much smaller, sessile, the lower- 
most ca. 1 cm, ovate or oblong, dentate or entire, the others bracteiform, 3-4 mm long, 
1-1 .5 mm wide, oblong-eUiptic; flowers in 3-7-flowered or many-flowered cymes, these 
except the uppermost dichotomously branching, often repeatedly to form a bostryx; ag- 
gregate inflorescence (including inflorescences of axillary branches) a very lax panicle to 
50-80 cm long; peduncles of upper cymes 2.5-10 mm long, others 1.5-4 cm long; bracts 
oblong or ovate-eUiptic, 1.5-2 mm long, ca. 1 mm wide, with ciliolate margin, velutinous- 
hairy like other parts of inflorescence; pedicels 0.7-1 .5 mm long; calyx 5.5-7 mm long, 
violet, the tube densely beset with sessile glands and on the nerves also with simple spread- 
ing white 2-3-jointed hairs; the frontal incision 1/3-2/5 (1/2) its length; teeth with broad 
membranous rim, the upper broadly ovate, 1/6-1/5 (2/9) as long as tube, short-acuminate 
or abruptly attenuate to a short point, the lower narrower, oblong-ovate, slightly longer; 
corolla blue, 11.5-15 mm long, with tube exserted to 1/3-2/5 its length, abruptly expand- 
ing above into neck 2-3 mm long and 2.5-3.5 mm wide; upper hp 2.3 mm long, nearly 
flat, cut to 2/3 into 2 obtuse lobes; middle lobe of lower Up 3.3-4 mm long, 5-6 mm 
wide, the lateral lobes broad, 0.6-0.8 mm long, ca. 2 mm wide; nutlets broadly eUipsoid, 
2 mm long, 1 .5 mm wide, obtusely 3-angled, brown, with minute flat tubercles. Fl. June- 
August; fr. from July. (Plate XXI, Figure 3.) 

Lower and middle mountain belt, at altitudes of 900-1700 m, exposed fine earth and 
gravelly-stony slopes. - Caucasus: S. Transc. (Nakhichevan ASSR and E. Armenia — 
Daralagez). Endemic. Described from Nakhichevan ASSR, Koshadara River. Type in 
Leningrad. 



267 



Note. The first specimens of this species, collected by Shovits along the Koshdara 
River, were quite erroneously included by Boissier (Fl. or. IV, 647) in N. trautvetteri 
Boiss. et Buhse, described earlier from Alliar village in Karadag (Iranian Azerbaidzhan), 
and ever since this species has been treated by aU botanists as N. trautvetteri. In actual 
fact the genuine N. trautvetteri differs fromN. velutina Pojark. in a number of charac- 
ters: 1) a completely different indument of leaves, stems and petioles, consisting of long 

399 spreading glandular hairs, interspersed with small sessile or subsessile capitate glands; 

2) longer and much narrower, lanceolate calyx-teeth, and 3) much narrower bracts cover- 
ed with glandular hairs. Specimens of the genuine N. trautvetteri Boiss. et Buhse from 
Transcaucasia were reported by students of the Caucasian flora under the name N. lepto- 
clada Trautv. which is merely a synonym of N. trautvetteri Boiss. et Buhse (see Note 
to N. trautvetteri). 

63. N. daghestanica Pojark. in Bot. mat. gerb. Bot. inst. ANSSSR, XV (1953) 315. - 
N. teucriifolia auct. fl. cauc: Lipsk. Fl. Kavk. (1899) 422; N. Pop. in Tr. Bot. sada 
Yur'evsk. univ. XIV, 234; Grossg. Fl. Kavk. Ill, 294; idem, Opred. rast. Kavk. 333, p. p. 
(quoad pi. daghest.) non Willd. (1809). 

Perennial; stems 40-70 cm long, arcuately ascending from nearly horizontal base, 2-3 cm 
across, simple or branching, in lower part leafy, violet, very shortly and densely hairy, 
green, with sparser pubescence above; internodes 7-1 1 cm long; branches fertile, mostly 
short, with 1 or 2 pairs of small leaves in lower part; cauline leaves 1.5-2.5 cm long, 1.2- 
2.2 cm wide, ovate-triangular, with broadly cordate, rarely truncate or cuneate base, ob- 
tuse, at margin with 5-7 large crenae, the upper side with minute (magnifying glass!) fine 
appressed hairs, the lower side more densely covered with thicker short 1-3-jointed hairs, 
often purple, with prominent main veins; petioles with short hairs sometimes interspersed 
with long hairs, the lower half as long again as blade, the upper (1/3) 1/2 the length of 
blade; floral leaves 3.5-5 mm long, the lower resembling cauline leaves, dentate, the others 
bracteiform, lanceolate, usually violet-tinged; flowers in 3-5-flowered cymes, these in 5-9 
pairs forming a lax raceme, usually compact, rarely the lower loosely dichotomous, the 
upper about 1-1 .5 cm apart, sessile or on peduncles 3-7 mm long, others 3-5 cm apart and 
on peduncles 2.3-5 cm long; bracts narrowly lanceolate, acuminate, violet, 2.5-3.5 mm 
long; pedicels 1 .5-2 mm, in bifurcation 2.5-5.5 mm long; calyx 6.5-8 mm long, the tube 
densely covered with round sessile yellow glands, the nerves and teeth with spreading 
white villous jointed hairs, incised at front to the middle or slightly deeper; teeth membra- 
nous at margin, the upper triangular- or lanceolate-ovate, 2/9-2/7 as long as tube (1.3- 
2 mm long, 0.6-0.7 mm wide), acute or short-acuminate, the lower narrowly lanceolate, 

400 1.5-2.2 mm long, long-acuminate; corolla blue, 16.5-18 mm long, pubescent outside, the 
slender curved tube exserted to the middle, abruptly expanding into neck (3) 3. 5-4. 5mm 
long and as wide; upper lip 2-3 mm long, cut to 2/3 into obtuse lobes; lower lip with 
middle lobe 3-4 mm long and 5-6 mm wide, the lateral lobes 0.7-0.8 mm long, 2-2.5 mm 
wide; upper stamens as long as upper lip of corolla or slightly shorter; nutlets brown, 
oblong-ellipsoid, 2 mm long, 1 mm wide, with small (high magnification!) round flattish 
tubercles. Fl. (?) July -August; fr. from first half of July. 

Lower mountain beU; exposed slopes, gravelly taluses and bluffs. - Caucasus: E. 
Transc. (Kuba region). Dag. Endemic. Described from S. Dagestan, Akhta village. Type 
in Leningrad. 



268 



Note. Of all the Caucasian species of the section Schizonepeta combined under the 
name of N. teucriifolia Willd., N. daghestanica and the closely related N. iberica 
most closely approach the diagnosis of N. teucriifolia. However, without examination 
of the original specimens, it is at present impossible to ascertain what in fact represents 
N. teucriifolia that was described from Turkish Armenia: too many different species 
come under this name. There are no grounds for assuming that N. teucriifolia occurs 
in the USSR. Of the species distributed in Turkish Armenia, two conform most closely 
to the description of N. teucriifolia, namely N. calvertii Boiss. (from near Erzerum) 
and N. rariflora C. Koch (from Ol'tin district). Thus Boissier, who had seen the origi- 
nals, may have been right in placing N. rariflora C. Koch among synonyms of N. teu- 
criifolia Willd. N. calvertii Boiss. and N. galatica Bornm. (from the northeastern part 
of Asia Minor) are most closely akin to N. daghestanica and N. iberica. 

64. N. iberica Pojark. in Bot. mat. gerb. Bot. inst. AN SSSR, XV (1953) 318. - N. 
teucriifolia auct.: Grossh. et Schischk. in Sched. pi. or. exs. No. 91; Grossg. Fl. Kavk. 
Ill, 294, p.p.; idem, Opred. rast. Kavk. 331, p.p.; Sosnovsk. Fl. Gruz. VII, 294. - Ic: 
Fl. Gruz. VII, fig. 319. - Exs.: PI. or. exs. No. 91. 

Perennial; root vertical, woody, coarsely fibrous, more or less twisted; stems few, 30- 
60 cm long, 1 .8-3 mm thick at base, erect, scabrous, with short dense hairs, violet in lower 
part, simple or with few fertile axillary branches; cauline leaves triangular, rarely ovate- 
triangular, obtuse or acute at apex, mostly deeply cordate, rarely truncate or cuneate at 
base, with 7-10 large ovate or oblong-ovate obtuse or acute teeth at each margin, scabrous 
on both sides with short hairs, spreading, these more copious and coarser beneath, finer, 
401 subappressed above; lower leaves to 8mm long, 3 mm wide, oblong, long-acuminate, with 
few teeth, the upper subulate-sublanceolate, violet; inflorescence of 5-10 cymes, mostly 
a fairly loose raceme; upper cymes 3-5-flowered, approximate, on peduncles 3-10 mm 
long, the lower 2-4.5 cm apart, on peduncles 1.5-3.5 cm long, often dichotomously 
branching, the axes of first order sometimes transformed into bostryces 1 .5-4 cm long, 
all axial parts of inflorescence scabrous, with short whitish hairs; bracts lanceolate, acu- 
minate, long-mucronate, ciliate, half as long again as pedicels, these 1 .2-1 .7 mm long, in 
bifurcation 2.5 mm long; calyx (7.5) 9-10.5 mm long, more or less suffused with violet, 
incised to the middle, covered with spreading hairs and small sessile glands; teeth with 
broad scarious sparsely ciliate margin, the upper triangular or ovate-triangular, (1/6) 2/1 1- 
2/9 as long as tube, the lower lanceolate, long-acuminate, mucronate; corolla 17.5-20mm 
long, blue, pubescent, the tube exserted to 2/5-1/3 its length, expanding into neck 3- 
4 mm long and as wide; upper lip 3 mm long, cut to 2/3 into obovate lobes; middle lobe 
of lower Up 3.5-4 mm long, 6 mm wide, the lateral lobes 0.6-0.8 mm long, 2.5-3 mm wide; 
upper stamens distinctly (1 .2 mm) longer than upper lip of corolla; nutlets ellipsoid, 
1.5 mm long, 1 mm wide, brown, with numerous small flat tubercles. Fl. (June?) July; 
fr. from second half of July. 

Dry slopes in central mountain belt. — Caucasus: E. Transc. (Trialet Range, vicinity of 
Tbilisi). Endemic. Described from vicinity of Tbilisi. Type in Leningrad. 

Series 2. Brevifoliae Pojark. — Flowers in compact semiverticels; upper pairs of 
semiverticels sessile, crowded in a globular or ovoid head, the lower 1-3 pairs remote, with 



269 



short peduncles; corolla 15-20 mm long, the tube about half as long again as calyx; nut- 
lets smooth. Plants usually branching from base, with more or less arched stems and 
branches. Habitats in alpine and subalpine belts. 
Two Caucasian species belong to this series. 

65. N. brevifolia C. A.M. Verzeichn. Pfl. Cauc. kasp. Meer. (1831) 93; Benth. Lab. 
gen. et sp. 483; C. Koch in Linnaea, XXI, 675; Boiss. Fl. or. IV, 147; N. Pop. in Tr. Bot. 
sada Yur'evsk. univ. XIV, 234; Grossg. Fl. Kavk. X, 293; idem,Opred. rast. Kavk. 331. - 
N. lamiifolia auct.: Benth. in DC. Prodr. XII (1848) 390, p. p. (quoad pi. Meyer.); Ldb. 
Fl. Ross. Ill, 379 (excl. syn. Benth. et Tourn.), non Willd. (1809). - N. alexeenkoi 
402 Woron. apud Grossh. Fl. Kavk. Ill (1932) 293, nom nud. in synon. — Exs.: Fl. Cauc. exs. 
No. 217. 

Perennial; root brown, woody but not tough, fibrous, twisted, passing above into a 
branching rhizome, this and stem base buried in rubble, covered with small dark brown 
squamiform leaves; stems many, 15-45 cm long, ascending, often curved or more or less 
flexuous, densely covered, especially in lower part, with very short spreading simple hairs 
and minute sessile whitish glands forming a powdery bloom, simple or (usually only in 
lower part) with long fertile axillary branches; lower internodes 1.5-3 cm, the upper 5- 
7 cm long; cauline leaves 9-17 mm long, 7-17 mm wide, broadly ovate, with broadly cor- 
date or truncate base, obtuse or rounded at apex, with 4-5 (6) large obtuse marginal teeth, 
densely covered on both sides with very short hairs, these appressed, 3-5-jointed above, 
shorter, spreading, 1-3-jointed beneath, intermixed with sparse small sessile capitate whit- 
ish glands and yellow punctate glands; lower petioles IVi-l times as long as blade, the up- 
per 2/5-1/2 as long; semiverticels few-flowered, the upper sessile, approximate at summit 
to form a small, usually loose, 8-14-flowered head, the other 1-2 (3) pairs remote, pedun- 
cled; upper peduncles 2-5 mm, the lower 12-25 mm long; lower floral leaves resembling 
upper cauline leaves, the upper smaller, oblong, dentate or entire, to fully bracteiform and 
then lanceolate, violet; bracts 2/5-1/2 as long as calyx, tapering to a long point; calyx 
7.5-9 mm long, tubular, slightly dilated above, densely covered outside, like the bracts, 
with small sessile whitish glands, the nerves with simple jointed spreading basally thickened 
hairs, the wide frontal incision reaching the middle and sometimes beyond; teeth cuspi- 
date (but not spinescent), the upper 3 ovate or ovate-triangular, mostly abruptly attenuate, 
1/5-1/4 as long as tube, the lower 2 narrower to triangular-lanceolate; corolla 15-20 mm 
long, bright blue, pubescent outside; tube slender, curved, exserted to 1/3-2/5, abruptly 
405 expanding into neck 3-3.5 mm long and 3.5-4 mm wide; upper Up bent forward, flat, 
2.3-2.5 mm long, cut to 2/3 into obtuse obovate lobes; middle lobe of lower lip 4.5-5mm 
long, 7-8 mm wide, the lateral lobes 1 mm long, 2.5-3 mm wide; nutlets brown, smooth, 
oblong-obovoid, 3-angled, 2.5-2.7 mm long, 1 mm wide. Fl. June-August; fr. from second 
half of July. 

Alpine and subalpine belts, on gravelly and stony shding taluses and on slopes. — Cauca- 
sus: Main Range, southeastern part. Endemic. Described from Tfandag Mountain. Type 
in Leningrad. 

Note. The specimen pubUshed in "Flora Caucasica" (exs. No. 217) from Dibrar Moun- 
tain (southern tip of Main Range) is not typical: it differs from all other available speci- 
mens in the much denser indument of all parts; the hairs on stems and calyx are broad, 



270 




PLATE XXIII. 1 - Nepeta grossheimii Pojark., general aspect and flower; 2 - N. saccharata 
Bge., summit of plant, flower, calyx in front view and nutlet from ventral side; 3 - N. amoena 
Stapf, calyx in front view. 



271 



flat, and comparatively long. Moreover, the leaves of this specimen are shaped differently, 
often broader than long and triangular in outline. More material from this area would be 
desirable. 

66. N. lamiifolia Willd. Enum. pi. Berol. (1809) 602; Benth. Lab. gen. et sp. 484 et in 
DC. Prodr. XII, 300, p. p. quoad pi. armen.; Boiss. Fl. or. IV, 645. 

Perennial; very similar to the preceding species, distinguished mainly by its large, dense, 
many-flowered heads and triangular leaves; stems 20-60 cm long, to 3 mm across; middle 
internodes to 12 cm long; leaves to 3.5 cm long and wide, mainly triangular-ovate, with 
truncate or broadly cordate base, tapering toward apex, obtuse to subacute, the margin 
with large acute or rarely obtuse teeth; inflorescence consisting of a compact (20) 30-50- 
flowered head 2-2.5 cm long and wide (without corollas) and 2-4 pairs of remote semiver- 
ticels; upper peduncles 3-6 (8) mm long, the lower 18-30 mm; axillary branches almost 
leafless, shorter than in N. brevifolia, terminating in a rather many-flowered head and 
1 or 2 remote verticillasters. Otherwise, Uke the preceding species. Fl. July-August; fr. 
from end of July. 

Gravelly taluses in subalpine and alpine belts. - Caucasus: S. Transc. (Akhmagan Range 
and mountains in vicinity of Lake Sevan). Gen. distr.: Arm.-Kurd. Described from Ar- 
menia. Type in Berlin. 

Note. N. lamiifolia was described by Willdenow from specimens cultivated in the 
Berlin botanical garden, the reported provenance being Armenia. Leading taxonomists, 
who had access to both Willdenow's originals and to typical specimens of N. brevifolia 
406 C.A.M., differ in their evaluation of the relationship between the species concerned. 
Whereas Bentham (in DC. Prodr. XII, 390) and Ledebour (Fl. Ross. Ill, 379) found it 
feasible to unite N. lamiifolia and N. brevifolia C. A.M. into one species (Ledebour 
noted only a few distinctions in vesture), Boissier regarded them as separate species and 
even stressed the absence of any kinship between them. In the detailed description of 
N. lamiifolia in "Flora orientalis," apparently based on Willdenow's originals (Boissier 
does note cite any other specimens), there are no significant differences as compared with 
our specimens collected in Armenia that are unquestionably close to N. brevifolia but 
differ from it in the above-noted features. On this basis the Armenian plant should be 
considered as a separate species. 

Section 8. Orthonepeta Benth. Lab. gen. et sp. (1834) 467; Briq. in Pflanzenfam. 
IV, 3a, 238. — Flowers bisexual and female (with normal pistil and 4 staminodes); calyx 
straight, narrowly cylindrical in flower, tubular-ovoid, ribbed in fruit; throat straight, 
rarely slightly obHque; teeth erect, lanceolate-subulate, aristate; corolla with curved tube 
abruptly expanding into a broad neck, included to the base of the neck or slightly higher, 
in female flowers nearly up to the limb; middle lobe of lower lip obUquely descending, 
concave, bearded and without sweUing at base, the margin cut into large teeth; nutlets 
tuberculate, sometimes only at apex, rarely almost smooth; bracts narrow, shorter than 
calyx; flowers in more or less remote cymes clustered in narrow racemes at ends of stem 
and axillary branches; central flower of cyme pedicellate. Perennial, high plants, mostly 
with long virgate axillary branches. 

About 15 species mainly in the Mediterranean countries and Southwest Asia. 



272 



Series 1. Pannonicae Pojark. - Corolla violet or white; nutlets hairy at apex. 
Besides N. pannonica L., this series contains N. nuda L. (SE Europe), N. latifolia 
L. (Spain) and N. violacea L. (S. Europe and, possibly, Asia Minor). 

67. N. pannonica L. Sp. pi. (1753) 570; Jacq. Fl. austr. II, 18; Bess. Enum. pi. Volhyn. 
23; Bge. in Ldb. Fl. Alt. II, 403. - N. paniculata Crantz, Stirp. Austr. (1769) 270. - 
Cataria nuda Moench, Meth. (1794) 388. - N. ucrainica auct.: M. B. Fl. taur.-cauc. 
11(1808)41; Bge. inLdb. Fl. Alt.II,403,nonL. -N. nudaauct.: M. B. 1. c.42; C.A. 
M. Verzeichn. Pfl. Cauc. kasp. Meer. 92; C. Koch in Linnaea, XXI, 676; Boiss. Fl. or. IV, 
663, p. min. p.; Shmal'g. Fl. II, 322; Kryl. Fl. Zap. Sib. IX, 2309; Grossg. Fl. Kavk. 292, 
p. p.; idem, Opred. rast. Kavk. 330, p. p. non L. — N. violacea auct.: Bge. in Ldb. Fl. 

407 Alt. II (1830)403; C. Koch in Linnaea, XXI, 677, non L. - N. nuda a. grandiflora 
Benth. Lab. gen. et sp. (1834) 486, p. p.; id. in DC. Prodr. XII, 387, p. p.; Ldb. Fl. Ross. 
Ill, 377, p. maxim, p. - N. nuda /3. pauciflora Benth. 1. c. (1834) 486; Ldb. Fl. Ross. 
Ill, 377, p. p. -N. nuda jS, ucrainica C. Koch, 1. c. 296. - N. barbata Rgl. et Winkl. 
in Tr. Bot. sada, VI (1879) 296; 0. and B. Fedch. Perech. rast. Turk. V, 143. - N. tur- 
kestanica Gandog. in Bull. Soc. Bot. Franc. 60 (1913) 26. — N. nuda ssp. pannonica 
var. paniculata Gams in Hegi, 111. Fl. V, 4, 2372. - Ic: Jacq. 1. c. tab. 129; Rchb. Ic. 
Fl. Germ. tab. 1243, fig. Ill; Fl. Yugovost. VI, fig. 605; Viznachn. rosl. URSR, fig. 244.- 
Exs.: GRF,No.733; Fl. pol. exs. No.380; Fl. exs. Austr.-Hung. No. 2948. 

Perennial; root oblique, woody, nodose, branching; stems many, 50-120 cm long, 4- 
7 mm across, sturdy, 4-angled, canahculate, glabrous or subglabrous in lower part, more or 
less densely covered above with very short simple spreading basally thickened hairs, with 
internodes 4-12 cm long, sometimes unbranched or with slender sterile branches in lower 
part, always branching in upper part; branches often long, virgate, bearing numerous re- 
mote cymes; cauHne leaves sessile (the upper) or nearly so (only lowermost with distinct 
petioles to 10-1 2 mm long), to 10 cm long, 3.5 cm wide, thin to fairly thick, bright or 
pale green, slightly glaucescent beneath, with prominent veins and with numerous punc- 
tate glands, covered on both sides with very short hairs, rather densely at first, later gla- 
brescent above, with pubescent veins and scattered hairs between them on the lower side, 
oblong-ovate or oblong-elliptic to lanceolate, with shallowly cordate or truncate base, ob- 
tuse, acute or acuminate, the margin crenate or dentate; only the lowermost floral leaves 
sometimes resembUng the upper cauline, often all bracteiform, linear-subulate, 3-10 mm 
long; cymes numerous, loose, the upper (sometimes most) 2-5-flowered, on peduncles 
2-5 mm long, subremote, the lower (rarely most) many-flowered (10-25), once or twice 
dichotomously branching, on peduncles ca. 10-1 2 mm long; bracts narrowly linear, 2- 
2.5 mm long; pedicels 0.3-0.5 mm long; calyx 4-5 mm long, tubular in flower, ovoid in 
fruit, straight, green or more or less suffused with violet, densely covered with short 
spreading hairs (like bracts and axial parts of inflorescence) and with round sessile resinous 
glands; teeth 3/5-1/2 as long as tube, narrowly and inconspicuously scarious-margined; 
corolla hairy outside, 8-9 mm long in bisexual flowers; tube 3-3.5 mm long, abruptly 

408 expanding into neck 2.3-2.5 mm long, 3-3.8 mm wide, included to base of neck or slightly 
higher; upper lip 2.5-2.8 mm long, cut into 2 obtuse ovate lobes; lower lip with middle 
lobe 2.5-3 mm long, 4.5-5 mm wide, the triangular lateral lobes 1-1.3 mm long, 2-2.5 mm 
wide; corolla varying in color from pale violet or pink to white; in female flowers corolla 



273 



5-6.5 mm long, included nearly to the limb; the lower lip relatively less developed; stami- 
nodes with rounded sterile scarious rudiments of anthers, these included in neck of tube; 
style twice as long as upper Up; nutlets eUipsoid, 3-angled, 1.5-2. 2 mm long, 1-1.2 mm 
wide, brown, smooth or mainly in upper part and on the back with sparse flat tubercles, 
at apex always with acute papilliform protuberances and sparse hairs. Fl. June to first 
half of September; fr. July-September. 

Chernozem and steppe strips on meadows, steppes, thickets, forest margins, sometimes 
open woods, exposed slopes in mountain regions in lower and central belts on meadow 
and steppe slopes. — European part: V.-Kama, U. Dnp., M. Dnp., V.-Don, Transv., U. Dns., 
Bes., Bl., Crim., L. Don, L.V.; Caucasus: Cisc, Dag., E., W. and S. (northern part)Transc., 
Tal.; West Siberia: Ob (southern part), U. Tob., Irt., Alt.; East Siberia: Yenis. (south- 
western part); Centr. Asia: Dzu.-Tarb., T. Sh., Pam.-Al. (Alai and Zeravshan ranges, very 
rarely in southern regions: found in Karategin and near Yakkabag). Gen. distr.: Centr. 
Eur., Bal.-As. Min. (northern part of Balkan Peninsula), Dzu.-Kash. (Kuldja area), Mong. 
Described from Hungary. Type in London. 

Notes. 1) Up to the present time this species ("Viznanchnik rosl. URSR"; Hegi) is 
often presented under the name N. nuda L. whichis not fully classified and is used in dif- 
ferent meanings. Moreover, there is no doubt as to the appropriateness of the epithet 
N. pannonica L. and hence its substitution by the epithet N. nuda L., published at the 
same time, is at any rate superfluous. Linnaeus reported Spain as the homeland of N. nu- 
da L., which has now been proven to be inaccurate. It is not clear if N. nuda Jacq. (Fl. 
Austr. I, 1773, tab. 24) corresponds to Linnaeus' N. nuda. The problem of identity or 
specific distinctiveness of N. nuda L. and N. pannonica L. has been the subject of a 
number of specialized studies. Summing up the views of Bauer, Janchen and Kerner, Freyn 
concluded, in agreement with these authors, that N. nuda L., a species distinct from N, 
pannonica L., is distributed in more southern parts of Europe (Albania, south of the 
Balkan Peninsula). Hayek considers N. nuda L. to be a white-flowered form of N. pan- 
nonica L.; it is possible that N. nuda Jacq. corresponds to this interpretation. The form 
described as N. nuda var. albiflora Boiss. from Asia Minor differs from the white-flow- 
409 ered form of the original N. pannonica in the more compact many-flowered cymes and 
broader calyx-teeth. The white-flowered form occurring in Western Transcaucasia (Abkha- 
zia) resembles the Southwest Asian form and requires further investigation as do the other 
Transcaucasian forms (especially those from Talysh) among which specific, geographical 
races may be separated upon detailed field study. 

2) With respect to N. turkestanica Gandog. (type from Talka River near Kuldja), we 
were unable to find any consistent differences from N. pannonica L. 

3) Investigation of the type specimen of the enigmatic species N. barbata Rgl. et 
Winkl., grown in the Pomologicheskii garden in Peterburg from seeds collected in Dzhun- 
garia, revealed that it does not essentially differ from N. pannonica and represents a 
form of this species deviating slightly in leaf shape, which is apparently due to growing in 
the shade. 

4) A hybrid N. pannonica X N. strictifolia was discovered in the Nakhichevan 
ASSR (Karagut massif)- The plant was a female specimen that closely resembled N. pan- 
nonica in the structure of the corolla as well as in the character of branching and shape of 
the inflorescence, but differed conspicuously from this species in the shape of the calyx 



274 



(narrowly tubular, curved in front), with teeth intermediate in shape between those of the 
two above-named species; shape and dentation of cauline and floral leaves and bracts, as 
well as indument, are very similar to those of N. strictifolia; corolla whitish, with violet- 
blue limb. 

Economic importance. The aerial part of the plant yields oil, the output ranging from 
0.09 (Kudryashev, 1934) to 0.15-7.28% (Prokhorov and Lebedev, 1932). The oil is pale 
yellow, mobile, and has an unpleasant odor; it is not soluble in alcohol and hence its im- 
portance in the perfume industry is negligible. A nectariferous plant. 

Cycle 1. Sulphureae Pojark. - Corolla yellow or cream-colored; nutlets without 
hairs at apex. 

In addition to the 3 species occurring in the Soviet Union, this cycle contains N. me- 
dia Stapf (NE. Iran, Alwand mountains) and N. marrubioides Boiss. et Heldr. (Asia 
Minor). 

68. N. sulphurea C. Koch in Linnaea, XXI (1848) 677. - N. pontica C. Koch, 1. c. - 
N. nuda /3. albiflora Boiss. Fl. or. IV (1879) 663, p. p. - N. nawaschini Bordz. in 
Protok. zased. Kievsk. obshch. estestvoisp. (1907) XXXIV; idem, in Zap. Kiu,..k. obshch. 
estestvoisp. XXV, 1, 109; Grossg. Fl. Kavk. Ill, 292 (excl. pi. Lenkor.); idem, Opred. 
rast.Kavk. 330. 

Perennial; root woody, thickening above; stems numerous, to Im long, erect, sturdy, 
410 to 5 mm across, scabrous, with very short antrorse hairs (these rather dense in upper part 
and mostly very sparse below), simple or branching; branches firm, usually short (only 
the lowermost to 1 5-20 cm long), with numerous subremote pairs of cymes; middle inter- 
nodes 3.5-7 mm long, usually shorter than leaves; cauline leaves short-petioled, the upper 
sessile (only the lowermost with petioles 8-15 mm long), oblong-ovate or oblong to sub- 
lanceolate, to 7-8.5 mm long, 2.5-3.5 mm wide, from cordate to orbicular or nearly cune- 
ate at base, mostly obtuse at apex, crenate or dentate, light green, paler beneath, scaberu- 
lous on both sides with very short hairs, the upper side with sparse appressed hairs, the 
lower side more densely hairy and beset with numerous punctate glands and with promi- 
nent coarse veins; lower floral leaves sometimes resembling the upper cauline, dentate, 
about equaUng the cymes, the others bracteiform, narrowly lanceolate or linear-lanceolate, 
entire, slightly exceeding peduncles; cymes gathered in long narrow racemes at ends of 
stem and of axillary branches, forming in aggregate a rather contracted narrow panicle; 
pairs of cymes, except sometimes the upper, distinctly remote; all cymes (even the upper) 
many-flowered, many of them once or twice dichotomously branching, the axes thick, 
large, densely covered with short spreading hairs; peduncles of upper cymes 2-4 mm, the 
lower 8-10 mm long; bracts pale green, with scarious colorless margin, shorter than calyx- 
tube; pedicels ca. 0.3 mm long, thick; calyx 5.5-7 mm long, pale green, densely covered 
with very short spreading hairs, the throat slightly obUque; teeth subequal, lanceolate- or 
linear-oblong, with green nerve and broad white margins, as long as or slightly shorter than 
tube; corolla white, sometimes with few lilac speckles on the yellow lower lip, hairy out- 
side, the hairs very short, longer only on the lips, in bisexual flowers 8-10 mm long; tube 
with neck included in calyx nearly to middle; upper Up 2.5-3 mm long, cut to the middle 
into ovate apically attenuate lobes; middle lobe of lower lip with a beard of long hairs at 



275 



base, 2.5-3 cm long, 4-5.5 mm wide, the lateral lobes triangular, 1.2-1.5 mm long, 2- 
2.8 mm wide; upper stamens as long as or slightly shorter than upper lip of corolla; style 
slightly exserted; corolla of pistillate flowers smaller, 7-8 mm long, included up to the 
limb, the middle lobe of lower Up relatively less developed; staminodes included in the 
dilated part of tube, with very small scarious rudiments of anthers; style long-exserted, 
curved, with large lobes 1mm long; nutlets 2-2.5 mm long, 1-1.2 mm wide, 3-angled, 

411 ellipsoid, brown, becoming almost black, finely and obtusely tuberculate on both sides, with 
acute, tightly crowded tubercles near apex, but not hairy. Fl. second half of June to July; 
fr. second half of July to August. 

Subalpine and central mountain belts, meadow slopes and forest margins. - Caucasus: 
S. Transc. and Tal. Gen. distr.: Bal.-As. Min. (northeastern part of Asia Minor) and Arm.- 
Kurd. (western part). Described from Gemshin at upper reaches of Fortuna River on 
Pontus Range. Type was in Berlin. 

69. N. sintenisiiBornm. in Fedde, Repert. sp. nov. X (1912) 421; 0. andB. Fedch. 
Perech. rast. Turk. V, 145. 

P lal; root woody, dense, oblique, 2.5-3 mm across, passing at apex into branch- 
ing rhizome; stems erect, sturdy, 4.5-6.5 cm long (and probably longer), 3-6 mm across, 
densely leafy (leaves being longer than internodes), above middle with (2) 4-6 axillary 
fertile branches 6-20 cm long, sometimes with few short slender sterile branches in lower 
part, grayish tomentose or almost arachnoid, the hairs fine, implexed, jointed, viscid, dense 
on upper part of stem especially beneath inflorescence and persistent, rather early disap- 
pearing in lower part; cauline leaves sessile (only the lower with petioles 5-8 mm long), 
5-8 (9)cm long, 2-2.5 (3. 5)cm wide, lanceolate, rarely ovate-oblong, cordate or subhastate 
(the upper sometimes cuneate) at base, gradually attenuate above, obtuse or acute at 
apex, evenly crenate-dentate, covered on both sides with gray implexed fine jointed 
viscid hairs and rather numerous round yellow resinous sessile glands; leaves of sterile 
stems and of axillary sterile shoots usually wider, the lower petioled; verticillasters com- 
pact but few-flowered (6-12 flowers), the upper closely approximate, short-peduncled, 
the lower 1-4 more or less remote and on longer peduncles (3-8 mm); lower floral leaves 
resembling the cauline, decreasing in size toward summit of inflorescence, becoming nar- 
rower (to Unear-lanceolate) and progressively less toothed, the upper bracteiform, narrow- 
ly Unear, bracts narrowly linear, 0.2-0.4 mm wide, half as long as calyx or rarely longer; 
pedicels 0.4-1.5 (2) mm long; calyx 7-8 mm long, 1.5-2 mm wide, cylindrical, becoming 
subobovoid in fruit; teeth nearly subulate, equal in length, as long as calyx-tube or sUghtly 
longer; calyx and bracts densely white-villous, with fine jointed hairs and densely covered 
with short whitish capitate glandular hairs and yellow sessile glands; corolla of bisexual 
flowers 10- 12 mm long (female ca. 8 mm long) pale yellow, with more intensely colored 

412 lower lip, hairy outside; tube not exserted; upper lip 2-2.5 mm long, cut to 1/3 into 2 
obtuse lobes; middle lobe of lower hp 2.25-3 mm long, 4-5 mm wide, coarsely crenate- 
incised, bearded at base, the lateral lobes nearly suborbicular, 1-1.5 mm long, 2-2.5 mm 
wide; upper stamens equaling upper lip, the lower half as long; anther-cells at first 
markedly convergent, later divergent at an angle of nearly 180°; style as long as upper 
lip; nutlets blackish-brown, bearded, broadly ellipsoid, 2-2.3 mm long, 1 .25 mm wide. 
Fl. second half of May to July; fr. from end of June. (Plate XXIV, Figure 1.) 



276 



Steppe plant communities in central mountain and subalpine belts. - Centr. Asia: Mtn. 
Turkm. (Kopet Dagh Range, western part). Gen. distr.: Iran. (N. Iran: Sammam moun- 
tains in Gilyan). Described from Sundzodag Mountain in W. Kopet Dagh. Duplicate of 
type in Leningrad. 

70. N. komarovii E. Busch in Bot. zhurn. SSSR, XXIV (1939) 423; Grossg. Opred. 
rast. Kavk. 331 ; Sosnovsk. Fl. Gruz. VII, 301. - Ic: E. Bush, op. cit. tabl. 

Perennial; root oblique, tough, woody; stems to 1 10 cm long, 3.5-5 mm across, sharp- 
ly 4-angled; slightly flexuous, in lower part usually glabrous, in upper part, especially be- 
low inflorescence, densely covered with very small (magnifying glass!) simple spreading 
hairs, in lower part with sterile, sometimes very short branches, above with 1 or 2 pairs of 
long fertile branches, the internodes usually slightly exceeding leaves; lower leaves brown, 
squamiform, other cauline leaves with stout petioles about 1/10 as long as blade, thin 
(dry — chartaceous), pale green, sparsely covered on both sides with very short simple 
hairs and with numerous punctate glands beneath, lanceolate, deeply cordate at base, 
obtuse at apex, evenly crenate or crenate-dentate along the whole margin, 5.5-8 cm long, 
2.7-3 cm wide; lower floral leaves resembling the cauline but at most half as long, decreas- 
ing in size and progressively less toothed toward summit of inflorescence, mostly bractei- 
form, narrowly linear, 5-10 mm long, 0.7-1.5 mm wide; upper cymes dense, 5-8-flowered, 
subsessile, forming compact verticillasters, the lower with up to 17 flowers, on peduncles 
5-12 (20) mm long, inflorescence on stem consisting of 10 pairs of cymes, 4-7 cm apart in 
lower part, 1-2 cm apart above; axillary branches with 5-10 pairs of few-flowered cymes ; 
bracts narrowly linear, 3.5-5 mm long, much shorter than calyx, acuminate; pedicels 1- 
1.5 mm long; calyx 8-10 mm long, pale green, the nerves densely covered with short 
413 spreading simple hairs; throat slightly oblique; teeth with ciliolate scarious margin, as 
long as tube, the upper 4-5 mm long, the lower slightly shorter; corolla cream-colored, 
with pale yellow lower Hp, 12.5-16 mm long, pubescent outside, included to base of neck; 
tube expanding into neck 3-4 mm long and as wide; upper lip 3.5-4.2 mm long, cut to 
middle into obtuse ovate lobes; lower lip with middle lobe 3-4 mm long and 5-6 mm 
wide, the obUquely triangular-semiorbicular lateral lobes 1.2-1.6 mm long and 2.7-3 mm 
wide; nutlets brown, eUipsoid, slightly enlarged below, obtuse at both ends, 3-angled, 
finely and acutely tuberculate. Fl. and fr. end of July. 

Subalpine belt, on rocks. - Caucasus: southern slope of Main Range. Endemic. De- 
scribed from S. Osetia - vicinity of Styr Gnukh village in basin of Malaya Liakhva 
River. Thus far only known from two locations near the above-mentioned village. Type 
in Leningrad. 



Section 9. Oxynepeta Benth. Lab. gen. et sp. (1834) 467 (excl. sp. 58-59); Boiss. 
Fl. or. IV, 639; Briq. in Pflanzenfam. IV, 3a, 238. - Oxynepeta Bge. Lab. Pers.(1873) 
58 (pro gen.). - Plants dioecious (or partly polygamodioecious?); flowers of two types: 
pistillate, the long style with large outwardly curved lobes and staminodes included in 
dilated part of corolla-tube; staminate, the normal stamens under upper lip and style 
shorter than stamens, lobes of stigma rudimentary, not divergent; individuals with such 
flowers usually sterile, but sometimes certain flowers (bisexual?) set fruits; calyx erect, 



277 



more or less cylindrical in flower; throat suberect; teeth lanceolate-subulate, aristate, 
covered inside and at margin of sinuses with coarse antrorse hairs; corolla completely 
included in calyx, with straight tube; upper lip 2/7-1/3 as long as the lower, with orbi- 
cular lobes; lateral lobes of lower lip erect, obHquely semiorbicular, the middle lobe up- 
right, tapering to a short broad claw, concave, with large-toothed margin; nutlets finely 
punctate and covered with small flat tubercles; flowers in cymes, forming small pyramidal 
panicles at ends of stem and of pyramidally disposed axillary branches; central flower of 
cyme sessile. Perennial plants. 

The section includes slightly more than ten species, most of them distributed in South- 
west Asia, including the Caucasus. 

Note. Species of this taxonomic group are erroneously characterized by Bunge and 
all later authors as gynodioecious, the central flower of each cyme supposedly being bi- 
414 sexual, the peripheral flowers imperfectly male. Examination of the enormous amount 
of herbarium material does not confirm this: the presence of dioecism is unquestionable 
(at least in all our species). 



Series 1. Parviflorae Pojark. — Flowers azure or blue. 

In addition to the species occurring in the USSR, this series also contains N. heliotro- 
pifolia Lam.s.l. (NE Asia Minor), N. curvidens Boiss. (S. and C. Asia Minor, in the 
north to Cappadocia inclusive), N. callichroa Hausskn. et Briq. (Iran) and N. hispa- 
nica Boiss. (Spain). 

71. N. parviflora M. B. PL taur.-cauc. II (1808) 41 ; Benth. Lab. gen. et sp. 486, in DC. 
Prodr. XII, 393, p. p.; Ldb. Fl. Ross. Ill, 380, p. p. (excl. pi. talysch.); Fl. Yugo-vost. VI, 
144; Grossg.Opred. rast. Kavk. 330, p. p. — N. ucrainica auct.: Boiss. Fl. or. IV (1879) 
668, p. p. (quoad pi. taur. et cauc. excl. pi. talysch.); Shmal'g. Fl. II, 324; Fedch. and Fl. 
Fl. Evrop. Ross. 808; Grossg. Fl. Kavk. Ill, 292, p. p. - N. euxina Velen. Fl. Bulg. Suppl. 
I (1892) 232; Hayek, Prodr. fl. Balcan. 263. - Exc: Callier, Iter taur. a. 1900, No. 703. 

Perennial; root woody, tough, more or less twisted, passing into a short rhizome; stems 
18-70 cm long, erect or somewhat ascending, densely villous the whole length or only below 
inflorescence with long white many-jointed simple (not glandular or viscid) hairs, in upper 
part usually with short crisp pubescence, strongly branching, the branches long, gradually 
shorter toward summit, fertile, usually branching in turn; leaves bright green or glaucous, 
firm, the main veins impressed above, prominent beneath, the upper side with sparse ap- 
pressed hairs, the lower side mostly rather densely covered with long spreading hairs, rare- 
ly the hairs sparse or shorter; cauline leaves to 5-6.5 cm long, to 2-3 cm wide, ovate or 
lanceolate-ovate, the upper often lanceolate, gradually narrower toward summit, cordate 
or the upper rounded or cuneate at base, usually obtuse at apex, the margin serrate-dentate 
or crenate-dentate; petioles villous or pubescent, the upper short, the lower long, some- 
times longer than blade; floral leaves mostly exceeding peduncles, the lower resembling 
upper cauline leaves, the upper bracteiform; bracts sessile, narrowly lanceolate, long- 
acuminate, entire, much longer than axes of cymes; cymes 3-5 pairs, on peduncles 3- 
7 mm (the upper) to 3-4 cm long, mostly twice or thrice dichotomously branching but 
compact; axes of cymes 3-7 mm long in flower (8-9 mm in fruit); calyx 6-7.5 (8) mm 



278 



long, usually green (like the bracts), rarely bluish, densely covered with crisp short or 

415 (especially in lower part long, white hairs; tube in flower cylindric-obconical, in fruit 3- 
4.5 mm long and as wide, subglobose or rounded-ovoid; teeth often shorter than tube, 
rarely as long, very rarely slightly longer, lanceolate-subulate; corolla 7.5-8.5 (9) mm long, 
blue-azure, as long as calyx; middle lobe of lower hp reniform, ca. 1.5 mm long, 2.5 mm 
wide; in staminate flowers stamens slightly shorter than upper hp, style usually scarcely 
reaching the base of stamens, rarely almost as long as lower stamens; in pistillate flowers 
style equaUng or slightly exceeding the upper lip of corolla, staminodes with distinct fila- 
ments and round scarious rudiments of anthers; nutlets 2-2.5 mm long, 1 .4-1 .6 mm wide, 
dark brown, eUipsoid, truncate at both ends, indistinctly and remotely finely tuberculate, 
usually covered at the top with thick hairs. Fl. second half of May to July; fr. from June. 

Steppe plants, also occurring on wastelands, chalky slopes, seldom among crops. — 
European part: M. Dnp., V.-Don (southwestern part), Transv. (western part, rarely), U. 
Dns., Bes., Bl., Crim. (Simferopol', Karasu-Bazar, Sudak, rarely), L. Don, L.V. (western 
part, rarely). Gen. distr.: Centr. Eur. (reported by Kerner for Central Hungary), Bal.-As. 
Min., Rumania (Dobrudja) and (according to Hayek) Bulgaria. Described from material 
from the Crimea and N. Caucasus. Type (lectotype) in Leningrad. 

72. N. ucrainica L. Sp. pi. (1753) 570; Bge. in Ldb. Fl. Ak. II, 403; Benth. Lab. gen. 
et sp. 487; in DC. Prodr. XII, 393 (excl. pi. taur.-cauc); Ldb. Fl. Ross. Ill, 381 (excl. pi. 
taur.-cauc); Fl. Yugo-vost. VI, 144; 0. and B. Fedch. Perech. rast. Turk. V, 148, p. p.; 
Kryl. Fl. Zap. Sib. IX, 23 10. - N. sibirica Bge. in Ldb. Fl. Ah. II (1830) 402 (cum auct. 
M.B.) non L. - Teucrium sibiricum L. 1. c. 564; Pall. Reise Russ. R. II, 269, III, 314; 
Georgi, Beschr. Russ. R. Ill, 5, 1077. - Ic: Rchb. Ic. Fl. Germ. tab. 1243, fig. II. 

Perennial; root woody, tough, branching; rhizome thickened, multicipital, sometimes 
branching; stems 17-50 cm long, erect or slightly ascending, subglabrous or covered with 
short simple upcurved hairs, sometimes with admixture of long white hairs on the ribs, 
pyramidally branching; leaves shorter than internodes, pale green, more or less pellucid, 
the upper side glabrous or with sparse short appressed hairs, the lower side hairy only on the 
somewhat prominent veins to (rarely) densely covered with short or sometimes longer, 
more or less spreading simple hairs; cauline leaves 2-4.5 cm long, 1-2.5 cm wide, ovate to 
lanceolate, the upper to narrowly lanceolate, with cordate or cuneate base, shortly or gra- 
dually tapering above, the upper attenuate, with acute or rarely obtuse tip, dentate or ser- 

416 rate, the uppermost sometimes entire; lower petioles as long as blade, the upper short; 
bracts, except sometimes the lower, longer than peduncles at flowering, as long or shorter 
in fruit, lanceolate, mostly long-acuminate, sessile, all entire except sometimes the lower; 
cymes at summit of stem 5-7 (8) pairs, fewer on axillary branches, on peduncles 0.5- 

1.5 cm (the upper) to 4-5 cm long, mostly simple, 3-flowered, the lower partly twice 
(thrice) loosely dichotomously branching, with axes of first order 0.5-0.8 mm long in flower, 
1-2 cm in fruit; bracts mostly shorter than calyx, rarely as long; pedicels 3.5 mm long, 
twice as long in fruit; calyx 9-12 (13) mm long (violet?) blue, densely covered, especially 
in lower part, with usually long spreading simple white hairs, in upper part (at times 
nearly all over) with sparser short crisp hairs sometimes intermixed with glandular hairs, 
narrowly cylindrical in flower, becoming oblong-obconical, expanding in fruit (2.5-3 mm 
wide), cylindrical or ovoid-cyhndrical, 2^2-3 times as long as wide; teeth usually shorter 
than tube or just as long, rarely slightly longer; corolla 9-12 mm long, blue, as long as 



279 



calyx; upper lip cleft nearly to base; lateral lobes [presumably of lower lip] slightly 
shorter than upper Up, the middle lobe reniform, 2-3 mm long, 3-5 mm wide; in stami- 
nate flowers upper stamens shorter than upper Up, style as in preceding species; in pistil- 
late flowers style as long as upper lip of corolla, with large downcurved lobes 1-1. 2 mm 
long; staminodes concealed in upper part of tube, with very short filaments and very 
smaU round rudiments of anthers; nutlets 2.5-2.7 mm long, 1.8-1. 9 mm thick, ellipsoid, 
rounded-truncate at both ends, blackish-brown in maturity, rather densely tuberculate, 
the tubercles small, very flat, sometimes more convex near apex, acute. Fl. May-June; 
fr. second half of June to July. 

Plains and mountain steppes and meadows, exposed gravelly and stony slopes. — Euro- 
pean part: V.-Kama (far north), Transv., L. Don (eastern part), L. v.; West Siberia: U. 
Tob., Irt.; Centr. Asia: Ar.-Casp. (far northern part), Dzu.-Tarb., T. Sh., Pam.-Al. (Alai 
and Transalai ranges). Gen. distr.: Centr. Eur. (N. Rumania, Hungary, very rarely), Bal.- 
As. Min. (reported for Dobrudja and Bulgaria by Hayek). Described from the "Ukraine" 
(in actual fact, from the southeastern European part of the USSR, apparently). Type in 
London. 

Notes. 1) N.ucrainica L. has been identified with N. parviflora M.B. by many of 
the earUer (as well as contemporary) authors. Bentham (1834), on the strength of exami- 
nation of the Linnaean specimens, estabUshed the faUacy of such an interpretation of N. 
ucrainica L. He pointed out that this species was identical with Teucrium sibiricum 
417 L. and that N. parviflora M. B. was a separate species which he interpreted in the same 
way as Bieberstein. N. ucrainica L. (in Bentham's interpretation) and N. parviflora 
M.B. are so markedly different from each other that it is difficult to imagine that Ben- 
tham, who distinguished them so weU, could have been mistaken as to the identity of 
Linnaeus' specimens. We take it that in this case we can fuUy rely on the authority of 
Bentham. In the European part of the USSR, the westernmost habitat of N. ucrainica 
is fixed by a herbarium specimen from the village of Mikhailovka on the Medveditsa River 
(tributary of the Don); the species is quite common on the right bank of the Lower Volga. 
It is noteworthy that the only reference cited by Linnaeus in his description of N. ucrai- 
nica was Herbert's list (1743) of plants from the Don and Volga. N. ucrainica is un- 
known outside the Ukraine. However, it has long been claimed for Transylvania and it is 
reported at present for Dobrudja and Bulgaria. It is possible that it occurs in Moldavia and 
Podolia. May not the N. campestris Andrz. (Izv.Kievsk.univ. 7, 1862, 126, nonBenth.), 
described from PodoUa, be N. ucrainica L.? 

2) In the scope adopted here, N. ucrainica is not wholly monotypic and represents a 
series of minor forms. In order to obtain an evaluation of their taxonomic significance, 
there is need of more material and field observations. A catmint of the Alai and Transalai 
ranges, recaUing N. kopetdaghensis in some characters, deserves special attention. 

73. N. schischkinii Pojark. in Addenda XIX, 358. - N. heliotropiifolia Ldb. Fl. Ross. 
Ill, 1 (1847-1849) 381, non Lam. - N. parviflora auct.: Benth. in DC. Prodr. XII(1848) 
393; Grossg. Opred. rast. Kavk. 330, p. p. quoad pi. talysch. - N. ucrainica auct.: C.A. 
M. Verzeichn. Pfl. Cauc. kasp. Meer. (1831) 92; Boiss. Fl. or. IV, 668, p. p.; Grossg. Fl. 
Kavk. Ill, 292, p. p., quoad pi. talysch. non M. B. 

Perennial; root tough, woody, branching; stems 30-60 cm long, 3-5 mm across, as- 
cending, acutely 4-angled, scabrous, with dense, very short, simple, spreading, more or less 



280 



reclinate, proximally thickened hairs, finally glabrescent at base, simple or with leafy 
sterile branches in lower part, with 3-5 pairs of long fertile branches above, these 2.5- 
4.5 (5.2) cm long, 0.8-2.4 (3.2) cm wide, sometimes branching in turn; leaves yellowish- 
green, rather firm, with veins impressed above and prominent beneath, covered on both 
sides with spreading curved or nearly crisp hairs, these longer than on stem, usually sparse 
on the upper side, much more copious beneath, especially on the veins; lower and middle 
cauline leaves approximately as long as internodes, rarely longer or shorter, with petioles 

418 2/3 (the lower) to 1/3 as long as blade, broad-ovate to ovate, truncate or cuneate or rarely 
cordate at base, obtuse at apex, the margin coarsely crenate or obtusely dentate; upper 
leaves much shorter than internodes, with petioles 2.5-6 mm long, oblong-ovate, rarely 
lanceolate, with cuneate base, gradually attenuate to obtuse or acute apex, toothed from 
base or only in upper part, usually entire near apex; floral leaves at flowering greatly ex- 
ceeding, later as long as or slightly shorter than peduncles, oblong or lanceolate, long- 
acuminate, entire, decreasing in size toward summit of inflorescence, becoming narrower, 
bracteiform; cymes 4-7 (9) pairs on stem, with peduncles 0.5-1 (the upper) to 4 cm long, 
the upper simple, 3-flowered, the lower twice or thrice loosely dichotomously branching, 
with axes of first order 5-8 mm long (in flower), later 12-20 mm long; lateral flowers of 
cymes on pedicels 1 .5-4 mm, finally to 8-10 mm long; calyx in flower narrow, tubular- 
cylindrical, (8) 9-1 1 mm long, in fruit with ovoid-cylindrical tube, more or less suffused 
with violet, densely covered with thick crisp hairs; teeth linear-subulate, very long-aristate, 
usually slightly longer than tube; corolla 6.5-9 mm long, shorter than calyx (calyx-teeth 
exceeding it by 1.5-2 mm), only in male flowers sometimes as long as calyx, blue, appress- 
ed-hirsute outside; upper Up 1-2 mm long, cut to the middle; lateral lobes of lower Up 
0.7-1 .2 mm long, 1 .5-2 mm wide, the middle lobe flabelliform, 2-3 mm long, 4.5-5 mm 
wide; in male flowers upper stamens as long as upper hp of cotolla, style reaching base of 
filaments, the lobes not divergent; in pistillate flowers style exceeding corolla by 1.5- 

2 mm, staminodes with very small round scarious rudiments of anthers and often with 
very short filaments; nutlets broadly ellipsoid, 2-2.2 mm long, 0.7-0.8 mm wide, truncate 
at both ends, dark brown, finely tuberculate on both sides, the tubercles obtuse, near apex 
more numerous and sometimes more convex or even acute. Fl. June to first half of July; 
fr. second half of July to September. 

Steppe and meadow slopes in central mountain belt. — Caucasus: E. Transc. (near 
Shemakha and southwestern part of Azerbaidzhan), S. Transc, Tal. Gen. distr.: may 
possibly occur in neighboring parts of Iran. Described from ravine Pir-Seid on Bozdag 
Mountain in southwestern Azerbaidzhan. Type in Leningrad. 

74. N. kopetdaghensis Pojark. in Addenda XIX, 359. — N. ucrainica a grandiflora 

419 f subsimplex et |3. parviflora f pubescens O. Ktze. in Tr. Bot. sada, X (1887) 228. — 
N. ucrainica auct. fl. turkest.: 0. and B. Fedch. Perech. rast. Turk. V (1913) 148, p. p. 
quoad pi. turcom. — Exs.: Sintenis, It. transcasp.-pers. a. 1900-1901, No. 185. 

Perennial, densely leafy, with woody, tough, often twisted, branching root and short 
rhizome; stems 25-65 cm long, to 5 (6) mm thick, sharply 4-angled, more or less densely 
covered with short crisp hairs interspersed, mainly along ribs, with very fine long many- 
jointed, arachnoid-implexed pellucid lustrous viscid hairs (these often very copious in 
lower part), unbranched below or developing short sterile branches, higher (rarely nearly 



281 



from base) with 5-8 pairs of leafy fertile branches, these sometimes in turn branching in 
lower part; leaves yellowish-green, thin, the veins (except the main) usually thin beneath, 
distinct but not prominent, impressed above; the lower side mostly densely covered v^th 
fine spreading pellucid viscid hairs (these shorter than on stem), the upper side usually 
with sparse appressed hairs (the upper leaves usually densely pubescent on both sides), the 
margins finely ciliate; cauline leaves to 4.5-5.5 cm long, 2.2-2.5 (3) cm wide, the lower 
oblong-ovate, with cordate or truncate base, obtuse, crenate, the others lanceolate, gradual- 
ly attenuate from cuneate base to an obtuse tip, the margin with subremote antrorse or 
spreading teeth; petioles puberulent, the lower ca. 2/3 as long as blade, often arachnoid- 
villous, the upper short; uppermost leaves sessile; lower floral leaves lanceolate, dentate, 
the upper entire, at flowering longer than peduncles, later as long or slightly shorter; 
cymes 3-4 pairs on stem, with peduncles 5-12 mm long (the upper) and to 4.5-5 cm (the 
lower), often simple, 3-flowered, some of the lower with 8(10) flowers, twice (thrice) 
loosely dichotomously branching, the axes of first order 5-10 mm long, later nearly twice 
as long; lateral flowers of cymes on pedicels 2-5 mm long, in fruit to 10-13 mm long; 
calyx in flower cylindric-obconical, 9-1 1 mm long, in fruit ovoid, violet; tube tomentose, 
with long fine spreading white hairs; teeth usually slightly longer than tube, lanceolate at 
base, long-acuminate, sparsely covered with very long hairs; corolla 8.5-1 1 mm long, 
usually equaUng or slightly shorter than calyx (only in staminate flowers sometimes ex- 
ceeding calyx-teeth), dark blue, puberulous outside; upper lip 1.5-2mm long, deeply cleft; 
420 lateral lobes of lower Up 1 mm long, 1.5-1.8 mm wide, the middle lobe flabelliform; in 
staminate flowers upper stamens 1/2 to 2/3 as long as upper lip of corolla, style hardly 
reaching the base of upper stamens, with short rudimentary connivent lobes; in pistillate 
flowers style not exserted from corolla, staminodes with very short filaments and very 
small round scarious rudiments of anthers; nutlets broad, ellipsoid-obovoid, 2.8 mm long, 
1 .8 mm wide, brown, obscurely tuberculate on both sides. Fl. May-June; fr. from first 
half of June. 

Steppe and meadow coenoses from foothills to subalpine belt. - Centr. Asia: Mtn. 
Turkm. (C. Kopet Dagh). Endemic. Described from Kopet Dagh, mountains above Nef- 
ton village near Ashkhabad. Type in Leningrad. 



Series 2. Erivanenses Pojark. - Flowers yellow or reddish. 

In addition to the Soviet species, this series should contain N. involucrata Bge. (N. 
Iran) and N. cryptantha Boiss. et Hausskn. (N. Syria and Mesopotamia). 

75. N. erivanensis Grossh. in Beih. Bot. Centralbl. XLIV (1928) 234; Fl. Kavk. Ill, 
291; Opred. rast. Kavk. 292. 

Perennial; stems many, 30-70 cm long, densely covered with short simple and capitate 
glandular hairs, sometimes interspersed with solitary long jointed hairs; upper part of 
stem, axillary branches and pefioles with glandular hairs only; branches often arising 
nearly from base of stem, rarely from its middle, nearly all fertile; leaves yellowish, light 
green, dull, thin, with thin but prominent veins, 2-3.5 cm long, 1-2.2 cm wide, all ovate 
or the upper oblong-ovate, with cordate or (mainly the upper) rounded-cuneate base, ob- 
tuse at apex, crenate or crenate-dentate, the uppermost sometimes entire; lower petioles 



282 



2/3 as long as blade, the upper short; lower floral leaves resembling cauHne leaves, the 
others lanceolate or oblong, often entire, acuminate; cymes on stems 5-6 pairs, the lower 
with peduncles to 4-5 cm long, 3-4 cm apart, the upper with peduncles 0.6-1 cm long, 
approximate ; many cymes twice (the lower thrice) dichotomously branching, only the 
upper simple, 3-flowered, all dense, short, the axes being 5-6 mm long, the pedicels 1- 
4 mm long, scarcely elongating after flowering; bracts oblong or lanceolate-linear, aristate- 
acuminate, as long as calyx or shorter; calyx 5-7.5 mm long, pale green, densely covered 
along the ribbed nerves with short thick antrorse hairs intermixed to varying extent with 
421 short-stipitate capitate pellucid glands, cylindrical in flower, with broadly ovoid tube; 
teeth mostly exceeding, sometimes up to half as long again, rarely as long as calyx, lanceo- 
late-subulate, the margin with short simple and glandular cilia; corolla yellow, puberulent 
outside, 6-8 mm long; upper lip 1 .2-1.5 mm long, deeply cut into ovate lobes; lateral 
lobes of lower lip nearly as long as lobes of upper hp, the middle lobe with a very short 
claw, the margin of the reniform limb with few large teeth; in staminate flowers upper 
stamens nearly as long as upper Up of corolla, style merely reaching the base of filaments, 
the rudimentary lobes connivent; in pistillate flowers style slightly exceeding the upper 
lip of corolla, with recurved lobes 0.8-1 mm long, staminodes with rather large round 
scarious rudiments of anthers and short filaments; nutlets broadly ellipsoid, truncate at 
ends, 2-2.5 mm long, 1.3-1.5 mm wide, sparsely tuberculate, the tubercles large but flat, 
at apex numerous, small, more convex. Fl. May-June; fr. from end of May. (Plate XXIV, 
Figure 2.) 

Taluses, stony and rocky slopes in lower mountain belt. — Caucasus: S. Transc. (south- 
ern part of Armenia and Nakhichevan ASSR). Endemic. Described from vicinity of 
Erevan. Type in TbiUsi? 



Section 10. Micronepeta Benth. in DC. Prodr. XII (1848) 394; Boiss. Fl. or. IV, 
639, p. p. (pro subsect.); Briq. in Pflanzenfam. IV, 3a, 238, p. p. (pro subsect.). — Calyx 
cylindrical (in fruit accrescent, usually ovoid or campanulate), straight with straight throat; 
teeth erect, covered, especially inside, at base and at margin of sinuses, with long stiff an- 
trorse hairs, these forming a fringe at margin of throat; corolla small (5-9 mm long), in- 
cluded mostly to limb, rarely to base of dilated part of tube; middle lobe of lower hp 
erect, with swelling at base; nutlets smooth, shining. Flowers in verticillasters or cymes, 
the upper mostly crowded in a head, rarely all remote; floral leaves usually large, longer 
than inflorescence. Annual plants. 

Species of this section occur mainly in Iran, also in Central Asia and Afghanistan. 



Series 1. Bracteatae Pojark. - Verticillasters forming a terminal capitate inflores- 
cence, sometimes 1 (very rarely 2) verticillaster remote; floral leaves of semiverticels large, 
ovate to lanceolate-linear, nerved, not spiny, tapering to short petiole, usually forming 
together with terminal floral leaves a kind of involucre around the head (and axillary 
422 semiverticels); calyx narrowly cylindrical in flower, slightly broader in fruit, the lanceo- 
late-subulate teeth separated by broad sinuses; upper lip of corolla deeply cut into oblong- 
ovate lobes; middle lobe of lower Up tapering to a short broad claw, entire, with concave 
lobes. Plants of subalpine belt and Central Asian juniper belt. 



283 



(423) 




PLATE XXrV. 1 - Nepeta sintenisii Bornm., summit of plant, bisexual flower, expanded corolla of 
pistillate flower, fruiting calyx, nutlet; 2 - N. erivanensis Grossh., summit of plant, staminate flower, 
expanded corolla of staminate flower, expanded corolla of pistillate flower, nutlet. 



284 



In addition to two species distributed through Central Asia, Iran and Afghanistan, 
this series also contains N. globiflora Bge. which is endemic to Iran. 

76. N.bracteataBenth. inDC.Prodr. XII(1848)395; Boiss. Fl. or. IV, 666; Lipsk. 
in Tr. Bot. sada, XVIIl, 69; O. and B. Fedch. Perech. rast. Turk. V, 147. - Zataria ? 
humilis Benth. 1. c. 183. - Exs.: Kotschy, PI. Pers bor. No. 179. 

Annual, 5-25 cm high, usually divaricately branching from base, with decumbent lower 
branches; only weak, small plants unbranched; stem and branches stiff, violet, covered 
with very short white curved proximally thickened hairs forming a glaucescent bloom; 
branches long, often equaling the stem, terminating in inflorescence, often bearing short 
sterile axillary branchlets; cauline leaves with petiole about the length of blade, ovate or 
rhombic-ovate (the upper often sublanceolate), 9-16 mm long, 5.5-12 mm wide, truncate 
rounded or cuneate at base, obtuse or acute, entire or often with 3-5 large acute remote 
teeth on each margin, rather densely covered on both sides with white 2-3-jointed basally 
thickened hairs, the main veins rather thick, violet beneath; terminal leaves (subtending 
the head) short-petioled, resembling the upper cauline or those subtending verticillasters; 
inflorescence a terminal head formed by several closely crowded, few- flowered verticil- 
lasters; one lower pair of remote capitate short-peduncled semiverticels sometimes present; 
leaves subtending semiverticels usually ovate, rarely elliptic, short-petioled, entire, dense, 
strongly nerved, to 10 mm long, together with terminal leaves forming a kind of involucre; 
bracts linear-lanceolate, ciliate, usually longer than calyx (rarely slightly shorter), Hke 
leaves subtending cymes green or violet; pedicels 1 .5-3 mm long; calyx cylindrical, in 
425 flower 4.5-6.5 mm long, in fruit 7-9.5 mm long, with 1 5 prominent nerves, green or violet, 
covered outside, especially along nerves, with long spreading white hairs; teeth linear- 
subulate, as long as tube or IVi (2) times as long, separated by rounded sinuses, these like 
teeth covered within along the margin with upright bristly hairs; corolla subglabrous out- 
side, 6-7.5 mm long, included to the limb; tube slightly curved, 3-4 mm long, abruptly 
expanding into elongate infundibular neck 2-2.8 mm long and 1 .5-2 mm wide; upper Hp 
1 .2-1 .5 mm long; lower hp with tetragonal-orbicular subpatent lateral lobes 0.5-0.7 mm 
long and 0.7-1 mm wide, the middle lobe tapering at base to a short broad claw, its limb 
1-1 .5 mm long, 1 .5-2 mm wide, rounded below, inflated from base along median line, 
shallowly emarginate, with entire upturned margins; upper stamens reaching only the 
middle of upper Hp of corolla; lower stamens and style not longer than lateral lobes of 
lower lip; nutlets ellipsoid, 1 .2-1 .5 mm long, 0.6-0.7 mm wide, pale brown, with obtuse 
edge on ventral side, slightly depressed at dorsal side. Fl. from May-June; fr. from June. 
(Plate XXV, Figure 2.) 

Stony slopes and taluses in Central Asian juniper belt and sheep's fescue steppe. - 
Centr. Asia: Pam.-Al. (Kugitang, Turkestan, Zeravshan, Darvaz range and W. Pamir). 
Gen. distr.: Iran. (Iran, Afghanistan and Baluchistan). Described from vicinity of Isfahan 
in Iran. Type in London. Duplicates of type in Leningrad. 

77. N. daenensis Boiss. Diagn.ser. 1, VII (1846) 53; ej. Fl. or. IV, 667; Benth. in DC. 
Prodr. XII, 395; Bge. Lab. Pers. 58; Lipsk. in Tr. Bot. sada, XVIII, 105; 0. and B. Fedch. 
Perech. rast. Turk. V, 147. -Exs.: Kotschy, PI. Pers. ausir. No. 91 5^. 

Annual; plants 3-18 (25) cm high; stem slender, erect, usually with axillary branches 



285 



at all nodes, green or reddish-violet, with very fine farinaceous bloom of minute hairs, 
these mainly simple, thickened toward base, interspersed with glandular hairs, on upper 
part of stem and in inflorescence almost exclusively capitate-glandular; nodes 2-4 (5), 
very far (often 6-1 1 cm) apart; axillary branches spreading or erect, arched at base, paral- 
lel to stem, sometimes branching in turn, terminating in inflorescence; leaves intense green 
or dark bluish-green, reddish-violet beneath, rarely anthocyanin-colored on both sides, 
linear or lanceolate, often curved and condupUcate, gradually tapering to narrowly cune- 
426 ate base, 6-8 mm long, 0.25-0.5 mm wide, obtuse or acute, entire or with few distant 

acute often subulate-linear teeth, with thin dense simple and glandular hairs, petioles near- 
ly as long as blade; verticillasters few-flowered, the upper subsessile, crowded in dense 
spicate-capitate terminal inflorescence, 0.5-1 .5 cm long and 0.6-1 .7 cm wide; floral leaves 
subtending inflorescence closely approximate, usually resembling cauHne leaves, petiolate, 
exceeding head width (often considerably so); leaves subtending verticillasters rather large, 
to 7 mm long, 3 mm wide, oblong-ovate to oblong or narrowly eUiptic, the lower green, 
the upper violet, firm, thick-nerved, entire, acute or acuminate; below terminal inflores- 
cence usually one pair of semiverticels (rarely 2), subapproximate or more often remote, 
short-peduncled; bracts narrowly lanceolate or narrowly eUiptic, acuminate, with densely 
white-ciliate margin, as long as calyx-tube or slightly longer; pedicels 0.5-1.5 (3) mm long; 
calyx 4-5 mm long, violet, glandular outside and with sparse spreading simple basally 
thickened hairs on the nerves; teeth linear-subulate, 1/2-2/3 times as long as tube, ciliate 
at margin, separated by rounded sinuses, with long upright bristles at margin and within; 
corolla 5.2-5.8 mm long, reddish (violet-pink?), sparsely hairy outside, the slender curved 
tube 3.2-3.6 mm long, included in calyx to base of abruptly expanding neck, this 1.2- 
1.3 mm long, 1.3-1.6mm wide; upper lip erect, 0.8-1.1 mm long; lower lip with horizon- 
tally disposed (at a right angle to upper hp) tetragonal subpatent lateral lobes 0.3-0.4 mm 
long and 0.7-1 mm wide, the large middle lobe subvertical, tapering to short claw, its Umb 
0.7-0.8 mm long, 1.7-2.1 mm wide, rounded at base, emarginate at apex, inflated at cen- 
ter from base to middle, concave at sides, with upturned margins; upper stamens slightly 
shorter than upper lip of corolla; style equaling upper stamens; nutlets brown, elUpsoid, 
1.2-1.3 mm long, 0.6-0.7 mm wide, with obtuse edge on ventral side, slightly concave on 
dorsal side. Fl. June-August; fr. from June. 

Stony slopes in subalpine belt. - Centr. Asia: Pam.-Al. (Shugnan and central part of 
S.Pamir). Gen.distr.: Iran. Described from Kukh-Daena mountains in S. Iran. Type 
in Geneva; duplicates of type in Leningrad. 



Series 2. Spathuliferae Pojark. — Flowers at summit of stem and of axillary branches 
i427 forming a capitate inflorescence, 1 or 2 distant verticillasters usually present below it; 

leaves subtending verticillasters not spinescent, spatulate or lanceolate-spatulate, tapering 
to long petiole, forming with floral leaves a kind of involucre around head and axillary 
semiverticels; calyx cylindrical-campanulate, in fruit ovoid-campanulate, the teeth lanceo- 
late-subulate, separated by rounded sinuses; upper lip of corolla deeply cut into oblong- 
ovate lobes; middle lobe of lower lip clawless, cordate, concave, entire. This series con- 
sists of one species native in the high-mountain region of Pamir and Hindu Kush. 



286 



78. N.spathuliferaBenth. inDCProdr.XII(1848)380; Boiss. Fl. or. IV, 668; Lipsk. 
in Tr. Bot. sada, XVIII, 106; 0. and B. Fedch. Perech. rast. Turk. V, 147. - N. renifor- 
mis Briq. in Bot. Tidskr. 28 (1908) 236. - N. fallax Briq. 1. c. 237. - Ic: in Bot. Tidskr. 
28 (1908) fig. 3. 

Annual; root slender, flexuous; stems 5-20 cm long, violet, sparsely covered with short 
glandular hairs, loosely covered in upper part (and when young also below) with long floc- 
culent arachnoid often fugacious white hairs, rarely simple, usually divaricately branching 
all the way up or only in lower part; branches arched-ascending, the lower decumbent at 
base, with inflorescence as developed as on stem; leaves semiorbicular or reniform, some 
ovate-rhombic, usually broader than long, with cordate or truncate or rarely cuneate base, 
rounded or obtuse apex, unevenly and remotely crenulate to entire, 5-12 mm long, 6- 
1 5 mm wide, usually more or less suffused with violet, both sides with loose tomentose- 
arachnoid hairs, these sometimes fugacious; petioles glandular hairy, about as long as 
blade; semiverticels sessile, (2) 3-(5)-flowered, 2 or 3 pairs at summit of stem (and 
branches) forming a 10-12-flowered capitate inflorescence 1.5-2 cm long and as wide; 
other 1 or 2 verticillasters remote; floral leaves scarcely differing from cauline leaves, 
long-petioled; leaves subtending verticillasters of the terminal head, especially the lower, 
more or less foliacious, spatulate or lanceolate-spatulate, forming together with terminal 
leaves a kind of involucre; semiverticels sometimes replaced by short (3) 5-flowered 
bostryces with numerous narrow bracts and then heads more compact, with more numer- 
ous flowers and with less distinct "involucre"; bracts petiolate, spatulate-ovate, to sessile, 
narrowly oblanceolate, acuminate, usually terminating in a rigid point, loosely covered 
428 outside with long implexed white hairs, often glandular, especially at base and along mar- 
gin, about the size of calyx; calyx bluish- or reddish-violet, 4-6.5 mm long, in fruit 7- 
8 mm, membranous, with indument as on bracts, (13) 15-nerved, the 5 nerves leading to 
teeth thick, firm, the others thin, sometimes indistinct; teeth equal, approximately half 
as long as tube, separated by broad rounded sinuses, these at margin and the teeth within 
densely covered with long hairs; corolla 7.5-9 mm long, dark blue or sometimes reddish- 
violet, the tube 4.5-5 mm long, shghtly exserted from calyx, expanding into an elongate 
neck (2-2.2 mm long, 1.5-1. 8 mm wide); upper lip erect, 1.3-1. 6 mm long, lateral lobes 
of middle [lower ?] Up short-tetragonal, slightly oblique, downcurved, 0.3-0.4 mm long, 
1-1. 2 mm wide, the middle lobe obUquely ascending, 1.2- 1.6 mm long, 2-2.5 (3) mm 
wide, with triangular swelling from base to middle, concave in front, shallowly emarginate 
at apex; upper stamens shorter than upper lip; nutlets oblong-eUipsoid, 2 mm long, 0.5- 
1 mm wide, pale brown, with indistinct obtuse edge on ventral side. Fl. June-August; 
fr. from second half of July. 

Stony and gravelly taluses in alpine belt and upper part of subalpine belt at altitudes 
of 2900-3800 m. - Centr. Asia: Pam.-Al. (Pamir, central, eastern and southwestern parts). 
Gen. distr.: Ind.-Him. (E. Afghanistan and probably NW Himalayas). Described from 
mountains near Kabul in eastern part of Afghanistan. Type in Kew. 

Note. This species was described under the three above-mentioned names. Duplicates 
of the types of N. reniformis Briq. and N. fallax Briq. are in the herbarium of the Bota- 
nical Institute of the Academy of Sciences of the USSR in Leningrad. Plants described 
under the name N. fallax develop short bostryces instead of semiverticels, a phenomenon 
very often observed in certain groups of Nepeta. Similar specimens of N. spathulifera 
have repeatedly been collected in Pamir. 



287 



Series 3. Pungent cs Pojark. — Inflorescence a small teniiinal head tomied by (1) 2-4 
pairs ot crowded cymes and several remote pairs ot" cymes; leaves subtending cymes (and 
bracts) rigid, with thick prominent nerves, spiny-tipped, usually longer than cymes; calyx 
narrowly cylindrical, in fruit ovoid, with linear- or lanceolate-subulate spinescent teeth 
separated by narrow incisions; upper lip of corolla shallowly cut into semiorbicular lobes; 
middle lobe of lower Up with round swelling at base, concave in front, entire, tapering to 
a claw. Plants of footluUs and lower (sometimes middle) mountain belt. 

429 Besides the three Central Asian species, N. chenopodiifolia Stapf from Iran also 
belongs to this series. 

79. N. pungens (Bge.) Benth. emend. Stapf; Benth. Lab. gen et sp. (1834) 487, quoad 
nomen; Stapf, Bot. Ergebn. Polak. Exped. 1 (1885) 47; Lipsk. in Tr. Bot. sada, XVIll, 
101, p. p.; Fedch.Perech. rast. Turk. V, 147, p.p.; Kryl. Fl. Zap. Sib. IX, 2312. - Zizi- 
phora pungens Bge. in Ldb. Fl. Alt. I (1829) 23. - Nepeta pusila Benth. 1. c. 438; in 
DC. Prodr. XII, 394; Ldb. Fl. Ross. IV, 382; Rgl. in Tr. Bot. sada, VI, 361. - Ic: Ldb. 
Ic. pi. tl. Ross. II, tab. 124. 

Annual; stem 5-25 cm long, erect, slender but sturdy, rather densely covered with very 
short downcurved basally thickened simple white hairs. 4-angled crimson-tinged along ribs 
and often all over, sometimes (in weak individuals) simple, mostly with 1-2 pairs of short 
branches, sometimes with a few long ramified branches from base; leaves bright green, cov- 
ered with very short hairs, these sparse, simple, appressed above, more copious (to very 
copious) beneath, interspersed with capitate glandular hairs; lower leaves with petioles as 
long as or longer than blade, ovate, 4-12 mm long, 3-6 (*^) mm wide, with truncate or 
rounded-cuneate base, obtuse or acute, with remotely toothed margin, thickish, with 
broad but not prominent nerves; middle leaves like the lower or narrower, to elliptic- 
lanceolate, acuminate, with shorter petioles; upper subtending leaves of 2-3 upper inflo- 
rescences short-petioled, narrowly lanceolate, somewhat rigid, with prominent thick 
nerves, rigidly subulate-pointed, the uppermost sometimes scarcely distinguishable from 
bracts; cymes 3-5-flowered, the upper 2-3 (4) pairs sessile or subsessile, forming an ovoid 
temiinal head (10) 12-1 7 mm long, 12-15 mm wide, containing 12-20 tlowers; 2-3 pairs 
of remote cymes below also sessile or subsessile, only the lower with developed peduncles 
1-2.5 cm long; semiverticels sometimes replaced by bostryces, these 2.5 cm long, densely 
beset with bracts; bracts slightly longer than calyx, narrow, elliptic-lanceolate to lanceo- 
late-linear, rigid, with thick hard nerves, curved, mostly conduplicate, long-spinescent; 
pedicels undeveloped; stems below inflorescence, peduncles, bracts and calyx outside 
roughened with very short thickened simple hairs and copious stipitate glands; cal.vx 5.5- 

430 7 mm long, (13) 15-nerved" teeth subequal, 2/5 to 1/2 as long as calyx, erect, long-acumi- 
nate, spiny-tipped, the margin (as in bracts) stiffly ciliolate, the teeth inside and margins 
of incisions covered with antrorse long bristly hairs; corolla 7-8.5 mm long, sparingly 
pubescent outside, reddish or white, with red speckles on lower Up; tube slender, 4.5- 
5.8 mm long, slightly curved, abruptly expanding into neck 1-1 .5 mm long, included to 
base of neck; upper lip 1.3-1. 7 mm long, concave, slightly cleft; lateral lobes of middle 
[lower?] lip obliquely ovate, subpatent, 0.7-1 mm long. 1.5-1.7 mm wide; upper stamens 
2/3 as long as upper hp. lower stamens barely exceeding its base, with anthers slightly 
smaller than in upper stamens; nutlets obovoid, 1.5 mm long. 1 mm wide, brownish-yellow, 
with indistinct edge on ventral side. Fl. May-June; fr. June-July. 



Rocks, bluffs, slopes of riverbanks in low mountains and foothills. — Centr. Asia: Ar.- 
Casp. (western part), Balkh., Dzu.-Tarb. (foothills). Gen. distr.: Mong. (northwestern 
part). Described from Arkaul mountains in E. Kazakhstan. Type in Leningrad. 

80. N. fedtschenkoi Pojark. sp. nov. in Addenda XIX, 360. — N. pungens auct. fl. 
turk. non Benth.: Lipsk. in Tr. Hot. sada, XVIII (1900) 101, p.p.; 0. and B. Fedch. 
Perech. rast. Turk. V, 147, p. p. 

Annual; root slender; stem 6-28 cm long, erect, with dense grayish indument, the 
hairs short, simple, white, recurved, more or less thickened toward base; stem simple only 
in the smallest individuals, usually branching, with 2-3 pairs of axillary fertile branches 
and often branching at base; branches long, often branching in turn; leaves pale, glauces- 
cent, the lower side paler, rather densely covered with short simple spreading hairs, the 
upper side subglabrous or with similar but appressed hairs; lower leaves (3) 6-22 mm long, 
(2.5) 5-13 mm wide, suborbicular to broadly ovate, with truncate or cuneate base, rounded 
or obtuse at apex, entire or incised-dentate with remote small acute teeth; the petioles 
slightly longer than blade; middle leaves similar to the lower or narrower, sublanceolate, 
acute, with larger teeth (4-6 per margin), like the lower leaves without prominent nerves, 
thickish, petioles as long as blade; uppermost leaves, i.e. those subtending the 2-3 upper 
pairs of cymes, sometimes resembling the middle leaves, but more often narrower, lanceo- 
late, with thick prominent midribs and lateral veins, entire, acuminate but often without 
431 a rigid point, rarely mucronate, short-petioled, usually exceeding inflorescence, including 
the terminal; cymes 3-8-flowered, 3-4 upper pairs clustered in capitate inflorescence, 
1 .5-2 cm long and as wide, 16-30-flowered, other pairs remote, the upper 1 or 2 sessile or 
on peduncles 2-5 mm long, the rest with peduncles 2-6 cm long; cymes sometimes re- 
placed by elongate cymose racemes (bostryces) densely covered with bracts; bracts green 
or more or less anthocyanin-tinged, narrowly lanceolate, rigid, nerved, curved, terminating 
in subulate mucro, the outer slightly longer than calyx, the inner usually as long or slightly 
shorter; flowers sessile; pedicels not quite distinct; calyx 6-8 mm long, in flower narrow- 
ly cylindrical, in fruit ovoid, dilated above base, 13-15-nerved, erect, covered, like bracts, 
with very short basally thickened simple hairs, the lower part of tube (and margin of 
bracts) longer; teeth lanceolate-subulate, erect, ciliate, beset within and at margin of 
sinuses with long antrorse bristly hairs, spiny-tipped, the upper 2-2.5 mm long, ca. 1/2- 
2/3 times as long as tube, the lower 2.3-2.7 mm long: corolla 7.5-9 mm long, bluish judg- 
ing by traces of color, hairy outside, included nearly to limb or to middle of neck; tube 
slender, long, 5-6.5 mm long, curved, expanding into neck 1 .4-1 .7 mm long; upper Hp 
erect, 1.3-1.7 mm long, shallowly incised above; lateral lobes of middle lip sub patent, 
obhquely ovate, 0.8-1 .2 mm long and wide; middle lobe with reniform limb ca. 1 mm 
long, 2 mm wide; upper stamens reaching the middle of upper lip of corolla, the lower 
disposed at level of margin of throat; anthers blue; nutlets eUipsoid-obovoid, 1.7 mm 
long, 1 mm wide, pale brown. Fl. May-June; fr. from first half of June. 

Stony and rocky slopes in mountain-steppe belt. — Centr. Asia: Pam.-Al. (mountains 
of southwestern end of Gissar Range, Zeravshan Range), T. Sh. (Syugaty mountains). 
Gen. distr.: Dzu.-Kash. (Kuldja, T. Sh.), Iran, eastern part of Afghanistan. Described 
from Zeravshan Range, near Madm village. Type in Leningrad. 

Note. In the structure of the inflorescence and in general habit, this species is very 
similar to N. pungens (Bge.) Benth. from which it is distinguished by the following 



289 



characters: absence of glaiuiular hairs on parts of the inflorescence; bhie (or lilac?) color 
of corolla; and tloral leaves, at least in the upper cymes, not spinescent, foliaceous or, in 
any case, with well developed blade. In tlie structure of floral leaves in the upper inflores- 
cences and the absence of glandular hairs. N. t>dtschenkoi approaches the Iranian spe- 
432 cies N. chenopodiifolia Stapf (N. pungens Benth. quoad typ.), whicii, however, is 
readily distinguisliable by its inflorescence: terminal head witli more numerous flowers 
(up to 40). floral leaves of a more herbaceous consistency, length of peduncles gradually 
increasing toward base but not reaching that observed in N. fedtschenkoi. 

81. N. microcephala Pojark. sp. nov. in Addenda XIX, 525. -N. pungens auct.: 
Bobrov in Tr. Bot. sada ANSSSR. XLIV (1931) 72. non Benth. 

Annual; stem 4-24 cm long, erect or slightly ascending, with 4-5 (7) remote nodes, 
rather densely covered with minute recurved basally thickened simple white hairs some- 
times interspersed with glandular hairs, green or violet-red, simple or with 1-2 (3-4) pairs 
of usually sliort fertile branches; leaves intensely or dark green, often violet-red beneath, 
the upper side with sparse appressed short thickish simple hairs, tlie lower side with rather 
copious spreading simple and glandular hairs, smooth (veins not prominent); lower leaves 
broadly ovate, 5.3-20 mm long, 7-14 mm wide, with rounded or truncate base, obtuse or 
rarely acute at apex, each margin with 3-6 remote small acute teeth, the petioles usually 
slightly longer than blade; other leaves (e.xcept those subtending terminal head) resembling 
the lower or narrower, narrowly elliptic to sublinear, tapering to narrow cuneate base, 
acute or acuminate at ape.x. the margin incised-serrate, the petioles 1/3-1/2 as long as 
blade; leaves subtending terminal head green or violet, narrowly lanceolate, long-acumi- 
nate, often aculeate, but more often narrower, bracteifomi, rigid, curved, spinescent, only 
slightly wider than inflorescence; cymes loosely 3-5-flowered, upper pairs sessile, crowded 
in a small, 5-10 (13)-tlowered head 7-12 mm long and as wide; other cymes distant, with 
slender peduncles (1) 2-5.5 cm long; bracts reddish-violet, as long as or slighfly longer 
than calyx, curved, narrowly lanceolate, rigid, with tliick midrib and 2 lateral nerves, 
spinescent, densely covered with minute simple hairs and sessile or short-stipitate small 
435 lustrous glands, the margin vsdth sparse longer basally thickened simple wliite hairs; flowers 
sessile; calyx in flower 5-7 mm long, 12-15-nerved, with indument as on bracts; teeth 
lanceolate-subulate, the upper half as long as tube, with bristly-ciliate margin, covered 
witliin and along margin of incisions wdth longer antrorse bristly hairs; corolla bluish (or 
bluish-lilac?), 6-7 mm long, scarcely exserted or even slightly shorter than calyx; tube 3- 
4 mm long, expanding into neck 1 mm long and 1.2-1.3 mm wide; upper lip 1.1-1.3 mm 
long, slightly cleft; lateral lobes of lower lip very oblique, ovate, ca. 0.8 mm long, 0.7 mm 
wide, sub patent, the middle lobe with reniform limb 0.6-0.8 mm long and 1.2-1.3 mm 
wide; upper stamens reaching the middle of upper lip of corolla, the lower not exceeding 
its base and nearly enclosed in neck of tube; anthers blue; nutlets obovoid. 1.2-1.3 mm 
long. 0.9-1 mm wide, brownish-yellow. Fl. (April?) May-June; fr. from second half of 
May. (Plate XXV, Figure 3.) 

Gravelly and stony slopes and bottoms of ravines at altitudes of 1200-1600 mm. — 
Centr. Asia: Mtn. Turkm. (Greater Balklian Mountains and central part of Kopet Dagh). 
Gen.distr.: Iran. (N.Afghanistan: Badkhyz). Described from Greater Balkhan Moun- 
tains, Dyuineg Mountain. Type in Leningrad. 



290 



(433j 




PLATE XXV. 1 - Nepeta santoana M. Pop., general aspect, flower, nutlet; 2 - N. bracteata 
Benth., general aspect, flower, nutlet; 3 - N. microcephala Pojark., general aspect, flower, nutlet; 
4 - N. satureioides Boiss., flower. 



291 



Note. This species is very readily distinguisiied from other species of the series Pun- 
gentes by the few-flowered, small, terminal head (smaller than the axillary semiverticels), 
elongated peduncles in all the axillary semiverticels and smaller flowers with scarcely ex- 
serted corolla. 



Series 4. Satureioideae Pojark. - Flowers in remote, sessile, few-flowered verticil- 
lasters; leaves subtending semiverticels bracteiform, slightly longer tlian calyx, rigid, lan- 
ceolate-linear, spiny-tipped; calyx-teeth lanceolate, aristate, separated by narrow incisions; 
upper lip of corolla concave, slightly emarginate, curved; middle lobe of lower lip tapering 
to broad claw, with drooping lobes. Monotypic series. 

82. N. satureioides Boiss. Diagn. ser. 1, V (1844) 24; ej. Fl. or. IV, 667; Benth. in 
DC. Prodr. XII, 395; Bge. Lab. Pers. 58; Lipsk. in Tr. Bot. sada, XVIII, 104; 0. and B. 
Fedch. Consp. V, 147. 

Annual; stem 4.5-55 cm long, slender but sturdy, erect, rather densely covered with 
fine spreading hairs interspersed, especially in upper part, with capitate glandular hairs, 
with 7-16 elongate internodes, branching from middle or from base; branches 1-4 pairs, 
436 long, with numerous verticillasters, branching; leaves grayish, covered above with sparse, 
sometimes copious, appressed, simple hairs, usually densely covered beneath with fine, 
more or less implexed simple hairs interspersed with glandular, the margins with glandular 
and simple cilia; radical leaves long-petioled, ovate to oblong-elliptic 7-13 mm long, 4- 
7.5 mm wide, with cuneate base, obtuse or acute at apex, the margin closely serrate; cau- 
line leaves 2.5-4.5 cm long, 4-7 mm wide, lanceolate, acuminate, gradually tapering to 
petiole, with few acute teeth in upper part; floral leaves lanceolate-linear or Unear, spiny- 
tipped, tlie lower with few teeth in upper part or entire, 2^/^-3 times as long as semiverti- 
cels, the upper entire, bracteiform, equaling or slightly exceeding semiverticels; bracts 
linear-lanceolate, often conduplicate, curved, rigid, spiny-tipped, green or in upper part 
violet-tinged, as long as or slightly longer than calyx; verticillasters (4) 5 to 10, remote, 
sessile, small, dense, few-flowered, the lower larger, 5-8 (lO)-flowered, the upper smaller, 
usually 3-, sometimes 2-flowered; calyx 6.2-7.2 mm long, narrowly tubular, straight, 
15-nerved, beset (like bracts) with sparse short glandular hairs intermixed with simple, the 
margin with a thick fringe of long thin white simple hairs; teeth erect, equal (3-3.3 mm 
long), attenuate to a rigid spiny point, covered within and at margin of incisions with long 
antrorse bristly hairs; corolla 8.5-9 mm long, pubescent outside, in dry state light straw- 
colored, with dark brown middle lobe of lower lip; tube 5-5.5 mm long, curved, slender, 
abrupfly expanding into neck, 1.6-2.2 mm long and 2-2.2 mm wide; upper Up 1.8-2 mm 
long, reclinate; lateral lobes of middle [lower?] lip obliquely ovate, 0.7 mm long, 1 mm 
wide, patent, the limb of middle lobe reniform, entire, strongly inflated at base; upper 
stamens inserted at base of upper Up, slightly exserted; lower stamens distinctly shorter, 
inserted at base of neck, with slightly smaller anthers; anthers bluish-black; style shorter 
than upper stamens; nutlets ellipsoid, ca. 1.7 mm long, 1 mm wide, pale brown, with an 
obtuse edge on ventral side. Fl. May -July; fr. from June. (Plate XXV, Figure 4.) 

Dry, mainly stony slopes, rocks, river pebbles in foothills and low mountains to 2400 m. 
- Centr. Asia: Ar.-Casp. (Mangyshlak Peninsula: mountains of Kara-Tau), Mtn. Turkm. 



292 



(Badkhyz mountains), Kyz. K. (mountain outliers), T. Sh. (western part - northern tip of 
Syr Darya Kara-Tau), Pam.-Al. (Zeravshan Range, eastern part of Turkestan Range, Alai 
437 Range, Darvaz, W. Pamir). Gen. distr.: Iran (Iran, Afghanistan). Described from C. Iran, 
near Isfahan. Type in Geneva. DupUcate of type in Leningrad. 



Genus 1255.* Glechoma** L. 

L. Sp. pi. ed. 1 (1753) 578. Chamaeclcma Moench, Meth. (1794) 393. - Generis Nepeta sect. 
Glechoma Benth. Lab. gen. et sp. (1832-1836) 484. 

Flowers short-peduncled in axils of middle and upper cauline leaves; calyx tubular- 
campanulate, straight, 15-nerved, 2-lipped, with longer upper teeth, declined after flower- 
ing; corolla bluish-lilac, the tube inflated at throat; upper lip cut into semiorbicular lobes; 
lower lip broad, 3-lobed, with rather long thick hairs at middle of lower lip; anther-cells 
at right angles to each other. Perennials, with creeping stems and long radicant shoots; 
leaves reniform or cordate, long-petioled, crenate. 

Note. This genus contains 12 species of which five, G. longituba (Nakai) Kuprian., 
G. grandis (A. Gray) Kuprian., G. hirsuta W. et K., G. brevituba Kuprian., and G. 
hederacea L., are distributed in the forests of Eurasia. G. hederacea has been intro- 
duced into N. America. 

G. decolorans (Hemsley) Turr., G. nivalis Jacqum., G. complanata (Dun.)Tur., 
G. rotundifolia Briq., G. pharica (Prain) Tur. and G. tibetica Jacqm. make up a dis- 
tinctive group of xeromorphous Himalayan-Tibetan species, most of which were quite 
judiciously referred by Levin (Tr. Bot. inst. ANSSSR, V, 1941) to a new genus, Pseudo- 
lophanthus Levin. 

1. Calyx-teeth 1/4-1/3 as long as tube; corolla 1 0- 1 8 mm long 

1. G. hederacea L. 

+ Calyx-teeth half as long as tube; corolla 18-25 mm long 2. 

2. Whole plant covered with white spreading hairs; flowers in axils of middle and up- 
per leaves in 4-6-flowered whorls 2. G. hirsuta W. et K. 

+ Whole plant covered with short coarse hairs; flowers in axils of middle leaves in 2- 
flowered whorls 3. G. longituba (Nakai) Kuprian. 



1. G. hederacea L. Sp. pi. ed. 1(1753)578; Maevsk. Fl. Sr. Ross. (1940) 612; Grossg. 
Fl. Kavk. Ill, 296; Kryl. Fl. Zap. Sib. IX, 2316; Kupriyan. in Bot. zhurn. SSSR, XXXIII, 
237, Plate l,Fig. 1.- Nepeta glechoma Benth. Lab. gen. et sp. (1832-1836) 485; Ldb. 
438 Fl. Ross. Ill, 379. - Chamaeclema hederacea Moench, Meth. (1794) 393. - Ic: 
Maevsk., op. cit.; Hegi, 111. Fl. V, 4, 2373. - Exs.: PI. Finl. exs. No. 889; Fl. pol. exs. 
No. 67; Fl. exs. Austro-Hung. No.3721 (1). 

Perennial, glabrous or very sparingly pubescent; stems 20-50 cm long, glabrous or with 
very short appressed downcurved hairs along ribs, procumbent or ascending, with numerous 

* Treatment by L.A. Kupriyanova. 
** From the Greek glechon, ancient name for corn mint. 



293 



radicant shoots; petioles of lower cauline leaves 5.6 cm long, those of middle leaves 2- 

3 cm long, nearly as long as blade; leaves with crenate margin, reniform or orbicular- 
reniform, the upper usually reniform-cordate; flowers in whorls of 3-4 in axils of middle 
and upper leaves; bracts shorter than pedicels, filiform, 1-1.5 mm long; corolla bluish- 
lilac, 10-18 mm long, usually l-lVz times as long as calyx; calyx narrow, tubular, the 
triangular teeth abruptly acuminate, 1/4-1/3 as long as tube; nutlets brownish, 2 mm 
long. May- July. 

Forests of the Soviet Union, shady riverbanks, coppices at forest margins, meadows, 
near dwellings. - European part: Kar.-Lap., Dv.-Pech., Bait., Lad.-Ilm., V.-Kama, U. Dnp., 
U.V., M. Dnp., V.-Don, Transv., U. Dns., Bes., Bl., L. Don (exceptionally!), L.V., Crim.; 
Caucasus: Cisc, Dag., W. and E. Transc; West Siberia: universal; East Siberia: Yenis. 
(rarely), Lena-Kol. (rarely), Ang.-Say., Dau.; Far East: Ze.-Bu. (collected one time in 
Bureya), Uda. (Okhotsk area, introduced); Centr. Asia: Dzu.-Tarb. Gen. distr.: Scand., 
Atl. and Centr. Eur., Med. (NW), N. Am. (introduced). Described from N. Europe. Type 
in London. 

2. G. hirsute W. et K. Icon, et Descr. pi. rar. Hungar. II (1805) 124, tab. 1 19; Fedch. 
and Fl. Fl. Evrop. Ross. Ill, 803; Kupriyan.inBot. zhurn. SSSR, XXXIII, 237, Plate 1, 
Fig. 2. - G. hederacea var. hirsuta Baumg. Enum. Stirp. Transs. II (1916) 165; Turr. 
Rep. Bot. soc. Exch. cl. Brit. Isles (1919-1920) 695. - Nepeta glechoma var. hirsuta 
Benth. Lab. gen. et sp. 485; Ldb. Fl. Ross. Ill, 380. - Ic: W. et K. 1. c; Kupriyan., op. 
cit. Exs.: Fl. pol. exs. No.669; Fl. exs. Austro-Hung. No.3722; Fl. exs. reip. bohem.- 
slov.No.53; Gerb. Fl. SSSR, No. 3771 a, c. 

Perennial; whole plant covered with long spreading white hairs; stems erect or slightly 
ascending, 30-80 cm long, with numerous creeping shoots; petioles of lower leaves 2-3 cm 
long, those of middle leaves up to 1 cm; leaves with crenate margin, the lower reniform- 
cordate, the middle cordate; flowers in 4-7-flowered whorls in axils of middle and upper 
leaves; bracts thin, filiform, shghtly shorter than pedicels; corolla bluish-lilac, 18-20 
(25) mm long, the broad tube 3H times as long as calyx; calyx hairy, narrow, tubular, 
439 8-10 mm long, the teetli subulate-acuminate, half as long as tube; nutlets brownish, 

4 mm long. April-May. 

Broadleaved forests. — European part: V.-Kama (vicinity of Kazan'), U. Dnp. (south), 
U.V. (south and southwest), V.-Don (rarely), M. Dnp., U. Dns., Bes., Bl. Gen. distr.: 
Centr. Eur., W. Med., Bal. (north). Described from Hungary. Type in Prague. 

Note. Forms that are intermediate between G. hederacea L. and G. hirsuta W. et 
K. occur in forests with elements of broadleaved species in the Leningrad and SW Vologod- 
skaya regions. In 1920, Grebner described plants from Lithuania similar to these as a sepa- 
rate species, G. hindenburgiana Grebn. (Fedde, Repert. 1920, 61). 

3. G. longituba (Nakai) Kuprian. in Bot. zhurn. SSSR, XXXIII (1948) 236, Plate 1, 
Fig. 4. - G. hederacea var. longituba Nakai in Tokyo Bot. Mag. XXXV (1921) 173, 
Kom. and Alis. Opred. rast. Dal'nevost. kraya, 11, 900. - Ic: Kupriyan., op. cit. 

Perennial, tinged with violet in lower and upper parts; stem 30-70 cm long, slender, 
ascending, covered, especially at the edges, with short coarse downturned hairs, with 
radicant shoots; petioles of lower leaves 1 .5-2.5 cm long, those of middle leaves ca. 1 cm 



294 



long; lower leaves reniform-cordate, the middle cordate, roughened above \\ith short 
coarse hairs, these densely covering the veins beneath where intermixed with short black 
glands; flowers usually paired in axils of middle leaves; bracts equaUng or exceeding 
pedicels, subulate-fihform, 2-3 mm long; corolla bluish-lilac, 20-25 mm long, with slender 
long tube; corolla usually 2^-3 times as long as calyx; calyx narrow, tubular, 8-10 mm 
long, the teeth subulate-acuminate, half as long as tube; nutlets brownish, 2 mm long. 
May-June. 

Broadleaved forests, coppices, riverbanks. — Far East: Uss. (south). Gen. distr.: 
Korea, China (Manchuria). Described from Korea. Type in Tokyo. 



Genus. 1256.* Dracocephalum** L. 

L. Sp. pi. (1753) 594. - Ruyschiana Boehr. ex Mill. Card. Diet. ed. VII (1759). - Moldavica Adans. 
Fam. II (1763) 190. - Zornia Moench, Meth. (1794) 410 ex parte. 

Calyx tubular or tubular-campanulate, 1 5-nerved, straight or curved, 2-Upped, the upper 
lip 3-toothed, its teeth 1/3-1/2 as long as the lanceolate teeth of lower lip, or else calyx 
obscurely 2-lipped, with subequal teeth, the middle tooth of upper Up broader than the 
others, the base of teeth nearly always with a thickened fold, this very noticeable when 
440 viewed from the side; calyx-teeth usually with very prominent transverse nerves (anasto- 
moses); corolla 2-lipped; tube narrow at base, widening at throat; upper Up erect or 
slightly curved, notched at apex or more or less deeply cleft; lower Up 3-lobed, v^th larger 
middle lobe; stamens 4, the upper pair slightly longer than the lower, the filaments as long 
as corolla or (rarely) exserted; anthers glabrous or hairy, 2-celled, the cells divergent; 
style with 2 equal subulate lobes; nutlets oblong, smooth. Perennial herbs, sometimes 
becoming woody below; flowers in verticillasters in the axils of upper leaves, usually 
forming a compact globular or a lax oblong inflorescence. 

Forty-five species in extratropical parts of Asia and in Centr. Europe; one species 
(D. parviflorum Nutt.) in N. America. 

1. Anthers pubescent 2. 

+ Anthers glabrous 4. 

2. Leaves deeply 3-5-partite, with linear or oblong obtuse lobes; calyx villous . . . 
34. D. austriacum L. 

+ Leaves entire; calyx short-hairy 3. 

3. Stem and calyx puberulent; bracts subovate, densely ciliate; corolla 20-25 (28)mm 
long 33. D. ruyschiana L. 

+ Stem subglabrous; bracts ciliate; calyx finely pubescent only in lower half, sub- 
glabrous above; corolla 35-40 mm long 32. D. argunense Fisch. 

4. Annuals 5. 

+ Perennials 7. 

5. Flowers small; corolla 7-9 mm long, slightly exceeding calyx 

18. D. thymiflorum L. 

* Treatment by B.K. Shishkin. 
** From the Greek dracon, dragon, and cephale, head, from the shape of the corolla. 

295 



+ Flowers larger; corolla 15-25 mm long, 2-2V^ times the length of calyx . ... 6. 

6. Plant 5-1 5 cm high, with disagreeable odor; leaves 0.5-1 .5 cm long, 0.3-0.8 cm 
wide; corolla 1 5- 18 mm long 22. D. foetid um Bge. 

+ Plant (15) 20-50 cm high, pleasantly scented; leaves 1.5-3 cm long, 0.7- 1.8 cm 

wide; corolla 18-25 mm long 23. D. moldavica L. 

7. Stamens more than twice as long as corolla (Centr. Asia) 

35. D. stamineum Kar. et Kir. 

+ Stamens not exceeding corolla or very slightly exserted 8. 

441 8. Calyx markedly 2-lipped, the upper lip cleft to 1/3-1/4, its 3 teeth subequal, ovate, 

the lower lip cleft to 1/4 or nearly to base 9. 

+ Calyx not markedly 2-lipped, all teeth subequal, the middle tooth of upper lip often 
much wider than the others 16. 

9. Bracts with 1-3 aristate teeth on each margin 10. 

+ Bracts foliaceous, with obtuse teeth or entire, sometimes with aristate teeth . . 14. 

10. Cauline leaves broadly ovate, subulate-dentate, all teeth subulate-pointed (W.T. Sh.) 
28. D. spinulosum M. Pop. 

+ Teeth of cauline leaves obtuse or acute but not subulate-pointed or very rarely some 
teetli subulate-pointed 11. 

1 1 . Stems usually decumbent or the central (if present) erect, 5- 1 9 cm long; leaf blade 
subreniform or broadly ovate, with cordate base, 1.2-2.5 cm long, 2-2.5 cm wide, 
evenly toothed 26. D. heterophyilum Benth. 

+ Stems erect or ascending, 20-70 cm long; leaf blade ovate or lanceolate-ovate, cune- 
ately tapering to base, 1-4 cm long, 0.3-1 .5 cm wide, unevenly toothed, pinnatifid, 
sometimes bipinnate or leaves entire 12. 

12. Leaves deeply pinnatisect (sometimes nearly to midrib), sometimes bipinnate. . . 
25. D. bipinnatum Rupr. 

+ Leaves almost entire or with shallow teeth or lobes not reaching midrib of leaf . 13. 

13. Bracts with a point 1-2 mm long; cauline leaves acuminate, entire or toothed, the 
teeth aristate 24. D. peregrinum L. 

+ Bracts with awn 3-5 mm long; cauline leaves obtuse, entire or toothed, the teeth 

not aristate 27. D. diversifolium Rupr. 

14. Leaves broadly ovate, with cordate base; bracts similar to leaves but smaller, not 
aristate; calyx densely villous, without yellow glands . . 29. D. komarovii Lipsky. 

+ Leaves ovate, with cuneately tapering or lanceolate base; bracts with a long subu- 

442 late point; calyx very short-hairy, with some remote, yellow, punctate glands . 15. 

15. Calyx-teeth with point 0.5 mm long (Mtn. Turkm.) 

31. D. subcapitatum (Ktze.) Lipsky. 

+ Calyx-teeth with subulate point 1.5-2 mm long (Caucasus) 

30. D. multicaule Montbr. et Auch. 

16. Leaves lanceolate or lanceolate-Unear, acute, entire or few-toothed 17. 

+ Leaves orbicular-ovate or oblong, obtuse at apex, the margins with rounded obtuse 

teeth or pinnatifid 18. 

17. Leaves and their lateral teeth (if present) terminating in a subulate point 0.5- 

1.5 mm long (E. Siberia) 15. D. fruticulosum Steph. 

+ Leaves acute, not subulate-pointed (Altai and Centr. Asia). . 16. D.integrifolium L. 



296 



18. Stems procumbent, 1.5-8 (12) cm long, with ascending branches, forming a dense 
tuft; leaves small, the blades 3-1 5 (20) mm long, orbicular-ovate or ovate, more or 
less deeply and evenly cut into rounded teeth 19. 

+ Stems solitary or many, usually erect, 1 2-70 cm long, not forming a dense tuft; 

leaves often larger 25. 

19. Bracts long-cuneate at base, entire or with 3 short teeth above (Caucasus) .... 
7. D. botryoides Stev. 

+ Bracts with 4-7 aristatc teeth 20. 

20. Middle tooth of upper lip of calyx 1 ^/i-2 times broader than long and 4-5 times as 
broad as lateral teeth; bracts equaling or shorter than calyx 21. 

+ Middle tooth of upper lip of calyx longer than broad and up to 3 times as broad as 
the lateral 29. 

21. Bracts and calyx without punctate glands, their teeth short-pointed at obtuse apex; 
corolla 10-1 2 mm long 6. D. paulsenii Briq. 

+ Bracts and often calyx copiously punctate-glandular, their teeth subulate-pointed; 
corolla 13-1 5 mm long (Alt., Ang.-Say.) 5. D. discolor Bge. 

22. Corolla 1 5-30 mm long, the upper lip much longer than the lower; style exsertcd 
5-7 mm 2. D. palmatum Steph. 

+ Corolla 12-18 mm long, upper lip about as long as the lower; style only slightly 

exsertcd 23. 

23. Bracts with 2-3 teeth, often entire; leaf blade 2 cm long, 1-2 cm wide (E. Siberia) 
1 . D. pinnatum L. 

+ All bracts with 3-7 teeth; leaf blade 0.5- 1. 2 cm long, 0.4-1 cm wide 24. 

24. Corolla 12-1 5 mm long; bracts often bluish, all teeth aristate, the terminal only 
slightly broader than the others 3. D. origanoides Steph. 

+ Corolla 20-22 mm long; bracts reddish-violet, the terminal tooth much broader 

than the others, commonly obtuse 4. D. bungeanum Schischk. et Serg. 

25. Bracts entire or very often with 1 tooth 26. 

+ Bracts toothed in upper half 29. 

26. Leaves dark green above, whitish-gray beneath with fine tomentum (Kuznetskii 
Ala-Tau) 14. D. krylovii Lipsky. 

+ Leaves concolor, green or grayish-green on both sides 27. 

27. Calyx 6-8 mm long; flowers often nodding; inflorescence elongate and lax after 
flowering, to 10-40 cm long, ca. 2 cm wide; corolla blue ... 17. D. nutans L. 

+ Calyx 13-17 mm long; inflorescence capitate, only slightly elongating after 

flowering 28. 

28. Leaf-blade orbicular or broadly ovate, with cordate or cuneately tapering base, 
evenly and obtusely toothed, about as long as wide; bracts shorter than calyx . . 
11. D. stellerianum Hiltebr. 

+ Leaf-blade oblong or ovate, toothed or sometimes nearly entire, cuneate at base; 

bracts longer than calyx 12. D. fragile Turcz. 

29. Leaves green above, white-tomentose beneath (Far East). . 13. D. multicolor Kom. 
+ Leaves more or less hairy beneath but not tomentose 30. 

30. Lower and middle cauhne leaves with petioles slightly shorter than blade (Centr. 
Asia) 31. 



297 



+ Lower and middle leaves with petioles as long as or longer than blade 33. 

31. Teeth of bracts terminating in awn 2-4 mm long; calyx 1 1-12 mm long, calyx-teeth 
aristate 19. D. nodulosum Rupr. 

444+ Teeth of bracts terminating in mucro 0.5-1 mm long; calyx 12-20 mm long . .32. 

32. Calyx 20mm long; corolla 35 mm long; leaves 1-3 cm long, 4-7 mm wide. . . . 
21. D. scrobiculatum Rgl. 

+ Calyx 12-15 mm long; corolla 25-30 mm long; leaves 1-2 cm long, 5-10 mm wide 
20. D. oblongifolium Rgl. 

33. Radical leaves oblong-ovate, longer than broad; flowers large; corolla 30-45 mm 
long, the upper lip long-lanate inside 34. 

+ Radical leaves orbicular-cordate or orbicular-reniform, not longer than broad; corol- 
la 25-30 mm long, the upper lip glabrous inside 10. D. imberbe Bge. 

34. Stems hairy, especially in lower part; corolla intensely blue 

8. D. grandiflorum L. 

+ Stems glabrous; corolla pink-lilac (Gissar Range). ... 9. D. formosum Gontsch. 



Subgenus 1. Eudracocephalum Briq. inPflanzenfam. 3a and 3b (1897) 239. -Anthers 
glabrous; stamens not exserted from corolla. 



Section 1. Buguldea Benth. in DC. Prodr. XII (1848) 398. - Calyx 2-hpped, upper 
lip 3-toothed, the middle tooth 2-3 times as wide as the others, lower hp with 2 lanceo- 
late teeth. 



Subsection 1. Keimodracontes Briq., I.e. - Sect. Keimodracon Benth. Lab. gen. 
et sp. (1832-1836) 491 et in DC. Prodr. XII (1848) 396. - Stems spreading, strongly 
branching, forming dense tuft; leaves incised-crenate or palmatipinnatisect; flowers in 
capitate or oblong inflorescence. 

1. D. pinnatum L. Sp. pi. (1753) 594; Hiltebr. Monogr. Dracoceph. 43, 72; Turcz. Fl. 
baic.-dahur. II, 377. - D. pinnatifidum Lam. Encycl. Meth. II (1786) 319. - D. pinna- 
tum j3. baicalense Bge. Suppl. Fl. Alt. (1836) 53. - D. pinnatum oc. vegetius Ldb. Fl. 
Ross. 111(1846-1848)388. -Ic: Hiltebr. 1. c. tab. 10; Pall. Fl. Ross. tab. 111. 

Perennial; rhizome creeping or ascending, 3-4 mm thick, branching above; stems few, 
erect or ascending, densely covered with short spreading hairs, simple or few-branched, 
5-15 cm long; leaves with petiole 5-1 2 mm long, the blade ovate-orbicular or oblong, 
1-2 cm long, 0.8-1 .5 cm wide, cordate or abruptly cuneate-attenuate at base, villous on 
both sides or white-tomentose beneath, pinnatisect, sometimes palmatisect, the oblong 
teeth rounded at the ends; flowers short-pediceled, in verticillasters at end of stem form- 
445 ing an inflorescence, this compact capitate in flower, later oblong; bracts longer than 
calyx, cuneate, villous-ciliate, pinnatifid in upper half, the lobes produced into a fine 
point, the middle lobe ovate, the lateral lobes lanceolate; calyx 12-13 mm long, villous; 
middle tooth of upper lip elHptic-oblong, IVi-l times as broad as the others, all teeth 



298 



subulate-pointed; corolla bluish-azure, ca. 18 mm long, soft-pubescent outside, upper lip 
slightly curved, as long as the lower. June-July. (Plate XXVI, Figure 2.) 

Stony and sandy slopes, coastal pebbles. — East Siberia: Ang.-Say. (western shore of 
Baikal), Dau. Gen. distr.: Mong. Described from Siberia. Type in London. 

Note. According to Bentham, the Caucasian plant D. botryoides Stev. is preserved 
in the Linnaean herbarium under the name of D. pinnatum. 

2. D. palmatum Steph. ex Willd. Sp. pi. Ill (1800) 151 ; Hiltebr. Monogr. Dracoceph. 
52, 76; Ldb. Fl. Ross. Ill, 382. - Ic: Hihebr. 1. c. tab. 14. 

Perennial; rhizome creeping or ascending, 0.5-0.8 cm thick; stems many, covered 
with spreading hairs, 5-12 cm long; petioles patent-hairy, slightly widening at base, 0.5- 
2 cm long; leaves ovate-orbicular or broadly ovate, with truncate or obscurely cordate 
base, deeply pinnatifid (but not to midrib), with oblong obtuse often involute lobes, 
densely hairy on both sides or almost canescent beneath, 4-10 mm long and nearly as 
wide; upper leaves often smaller, short-petioled or subsessile; flowers short-pediceled, in 
verticillasters at end of stem forming an oblong inflorescence; bracts cuneate, with lanceo- 
late spiny-tipped teeth; calyx often violet, ca. 10 mm long; middle tooth of upper lip 
rounded-obovate, mucronate, obscurely denticulate, 3 times as wide as lateral teeth, the 
lower lip with 2 narrowly lanceolate teeth; corolla 20-30 mm long, pale straw-colored, 
slightly villous outside, upper lip much longer than the lower; style often strongly exsert- 
ed; nutlets ovoid, faintly 3-angled, ca. 2 mm long, 1 mm wide. July. (Plate XXVII, 
Figure 2.) 

Sandy deposits and bluffs, gravelly slopes in stony and mottled tundras. — Arctic: 
Chuk.,An.; East Siberia: Lena-Kol.; Far East: Okh. Endemic. Described from Siberia. 
Type in Berlin. 

3. D.origanoidesSteph. exWilld.Sp.pl. 111(1800) 151; Kryl. Fl. Zap. Sib. IX, 2321. 
446 ~ D. pinnatum a. altaicum Bge. Suppl. Fl. Alt. 1836 53. — D. pinnatum a. minus 

Ldb. Fl. Ross. Ill (1846-1848) 384; Kryl., Fl. Alt. IV, 1034. - D. pinnatum L. var. 
pallidiflorum Kar. et Kir. in Bull. Soc. Nat. Mosc. XV (1842) 422. - D. villosum 
Krassn. in Bot. zap. II (1887-1888) 19. - D. pinnatum Lipsky in Tr. Bot. sada, XXVI 
(1909) 604, non L. — D. pinnatum var. songaricum Lipsky, 1. c. 605. 

Perennial; rhizome 3-5 mm thick, branching in upper part; stems numerous, procum- 
bent and often radicant; branches erect, 2-8 cm long, covered with long recUnate simple 
hairs; leaves with hairy petiole as long as or longer than blade, orbicular-ovate or ovate, 
at base often subcordate, 0.5-1 .2 cm long, 0.4-0.8 cm wide, rather shallowly cut into 
5-9 rounded-ovate lobes or large teeth, hairy on both sides (more densely beneath), with 
sessile punctate glands; flowers at ends of stems in dense ovoid or subglobular heads; 
bracts 10-14 (15) mm long, obcuneate, bluish, finely long-ciliate, cut at apex into 5-7 
(rarely 4) teeth, the lateral linear-lanceolate, the middle one ovate-lanceolate, all thin- 
pointed; calyx 8 mm long, often bluish, finely pubescent, obscurely 2-hpped, middle 
tooth of upper lip aristate, Wi times as wide as the others, these lanceolate, also aristate; 
corolla blue, 12-15 mm long, puberulent outside and at base of lower Up within; lower 
lip slightly longer than the upper. June-July. 

Gravelly and stony slopes, moraines near glaciers. - West Siberia: Alt.; East Siberia: 



299 



Ang.-Say.; Centr. Asia: Dzu.-Tarb., T. Sh. Gen. distr.: Mong., Ch. (Sinkiang). Describ- 
ed from Siberia. Type in Berlin. 

4. D. bungeanum Schischk. et Serg. in Kryl. Fl. Zap. Sib. IX (1937) 2322. - D. ori- 
ganoides Bge. Suppl. Fl. Alt. (1836) 54, non Steph.: Ldb. Fl. Ross. Ill, 384. 

Perennial; rhizome long, 3-5 mm thick, branching above; stems many, procumbent; 
branches erect or ascending, 2-8 cm long, patent -hairy, the hairs short in lower part, 
longer above, interspersed with short glandular hairs; leaves on hairy petioles nearly as 
long as blade, orbicular-ovate or ovate, sometimes cordate at base, shallowly cut into 5- 
9 ovate or round large teeth, glandular-hairy on both sides, more densely beneath, 5- 
12 mm long, 4-10 mm wide; flowers in compact globular or ovoid heads; bracts longer 

447 than calyx, obovate, cuneate at base, violet-tinged, long-hairy at margins, cut at apex into 
5-7 lobes, the terminal subovate, commonly obtuse, the lateral lobes narrower, lanceolate, 
thin-pointed; calyx tubular-campanulate, long-hairy, pale purple, obscurely 2-lipped, up- 
per Up 3-lobed, the middle lobe 1 ^i times as wide as the lateral, aristate, lower lip 2-partite, 
with lanceolate aristate lobes; corolla dark blue, 20-22 mm long, puberulent outside and 
at base of lower hp within, upper hp orbicular-ovate, shallowly cleft at apex, lower Up 
nearly as long as the upper, the middle lobe reniform, nearly twice as broad as long, very 
slightly emarginate, the lateral lobes broadly ovate, 2/7-1/3 as wide as the middle lobe. 
June-July. 

Alpine belt along pebby slopes and old moraines. — West Siberia: Alt. Gen. distr.: 
Mong. Described from Altai. Type in Leningrad. 

5. D.discolorBge.Suppl.Fl. Alt. (1836)51; Benth. in DC. Prodr. XII, 397; Ldb. Fl. 
Ross. Ill, 383; Kryl. Fl. Zap. Sib. VIII, 2320. - Ic: Ldb. Ic. pi. fl. Ross. II, tab. 128 
(sub. D. origanoide). 

Perennial; rhizome 3-5 mm thick, branching in upper part; stems numerous, procum- 
bent and often radicant, covered with remnants of dead leaves; branches erect, 2-8 cm 
long, short-hairy; leaves with pubescent usually long-ciliate petioles as long as blade, ovate, 
with subcordate or truncate base, 0.4-1 cm long, 0.2-0.8 cm wide, deeply (nearly to mid- 
rib) pinnatisect, with linear-oblong obtuse often revolute-margined lobes, green puberu- 
lent above, white-tomentose beneath; flowers at ends of branches in dense ovoid or 
oblong inflorescences; pedicels 1-3 mm long; bracts obcuneate, long-ciliate at margin, 
often punctate-glandular, cut at apex into 3-5 aristate lobes, the lateral lanceolate, the 
middle one ovate or oblong-ovate; calyx pale purple, pubescent, commonly with punctate 
lustrous glands, the tube cylindrical, the 2-Hpped limb nearly as long as tube; lateral lobes 
of upper lip ovate-lanceolate, aristate, the middle lobe 1^2-2 times as wide as long, 4-5 
times as wide as lateral lobes, emarginate at apex, with 1-3 aristate teeth, lobes of lower 
Hp lanceolate, awned-acuminate; corolla dark blue, short-hairy outside and at base of 
lower lip within, 12-15 mm long; upper lip cut to middle into 2 lanceolate lobes, incised 
at apex, crenate at margins, 4-5 times as broad as the ovate lateral lobes; style usually 
not exserted. May-June. 

448 Exposedstonyandgravelly slopes, dry steppes. -West Siberia: Alt.; East Siberia: Ang.- 
Say. Gen. distr.: Mong. Described from Altai. Type in Leningrad. 



300 



(449) 




PLATE XXVI. 1 - Dracocephalum scrobiculatum Rgl., part of plant, calyx, flower, bract; 2 
D. pinnatum L., general aspect, calyx, flower, bract; 3 - D. bipinnatum Rupr., part of plant, 
flower, bract, calyx. 



301 



6. D. paulsenii Briq. in Bot. Tidsskr. 28 (1908) 238. - D. discolor Lipsky in Tr. Bot. 
sada, XXVI (1906) 599, non Bge. - Ic: Briq. 1. c. fig. 4. 

Perennial; rhizome ca. 8 mm thick, branching at the end; stems numerous, densely 
covered with short spreading hairs, 5-15 cm long; leaves with petioles 2-3 mm long, ovate, 
deeply pectinately pinnatifid with revolute-margined incisions, green, appressed-puberulent 
above, white-tomentose beneath (?), 3-4 mm long and as wide; flowers short-pediceled; 
verticillasters in the axils of upper leaves fonning globular-ovoid or oblong inflorescences 
2-3 cm long and 1.5 cm wide; bracts elliptic, pinnatifid, with obtuse incisions, short- 
acuminate lobes, purple or purpHsh-blue, sparsely villous; calyx tubular-campanulate, 
more or less patent-hairy, 6-7 mm long, 2-Upped; middle tootli of upper Up broadly ovate, 
the lateral teeth narrowly ovate, the 2 teeth of lower Up ovate-lanceolate, aU short-acumi- 
nate; coroUa violet-blue, often with more deeply colored small spots, densely puberulent 
outside, 10-12 mm long. July-August. 

Stony slopes in alpine belt. — Centr. Asia: Pam.-Al., T. Sh. Gen. distr.: China (Sin- 
kiang). Described from Pamir. Type in Copenhagen. 

7. D. botryoides Stev. in Mem. Soc. nat. Mosc. Ill (1812) 266; Bge. Suppl. Fl. Alt. 41 ; 
Ldb. Fl. Ross. Ill, 383; Boiss. Fl. or. IV, 671; Grossg. Fl. Kavk. Ill, 297. - Nepeta pin- 
natifida Fisch. Catal. hort. Gorenk. (1812) 11, nom. nud. 

Perennial; rhizome ascending, 0.5-1 cm thick, branching above; stems usually numer- 
ous, spreading or ascending, simple or branching, 10-12cm long, densely covered with 
patent or slightly retrorse hairs; leaves with hairy petioles 0.5-1 cm long, ovate-orbicular, 
at base truncate or cuneate, sometimes subcordate, 10-15 mm long, 8-12 mm wide, deep- 
ly pinnatisect with oblong obtuse teeth, covered on both sides, more densely beneath, 
with grayish hairs; flowers short-pediceled; verticillasters forming capitate or oblong in- 
451 florescences, rarely lowermost whorl more or less distant; bracts cuneate, with mucro ca. 
1 mm long, entire or with 1-2 aristate teeth, villous, 7-8 mm long; calyx ca. 10 mm long, 
violet-tinged, middle tooth of upper lip broadly ovate, twice as wide as lateral teeth, aU 
teeth aristate; coroUa 15-18 mm long, pale violet, upper lip erect, slightly shorter than 
lower lip. Fl. July-August; fr. August-September. 

Stony slopes, pebbles near streams, rocks in alpine belt at altitudes of 2500-3600 m. - 
Caucasus: Dag., E. and S. Transc. (Zangezur). Endemic. Described from Caucasus. Type 
in Helsinki. 



Subsection 2. Calodracontes Briq. in Pflanzenfam. IV, 3a and 3b (1897) 239. - 
Section Calodracon Benth. Lab. gen. et sp. (1832-1836) 421. - Stems erect, sometimes 
few-leaved ; radical leaves long-petioled, crenate ; flowers in capitate or oblong inflorescences. 

8. D. grandiflorum L. Sp. pi. (1753) 595, ex parte; Benth. Lab. gen. et sp. (1832- 
1836) 421 et in DC. Prodr. XII, 399; Hiltebr. Monogr. Dracoceph. 48, 74; Ik.-Gal. and 
Il'in in Tr. Bot. inst. AN SSSR, ser. 1 , 3 (1936) 271 . - D. altaiense Laxm. in Nov. Comm. 
Ac.Petrop. XV (1770) 556; Ldb. Fl. Ross. Ill, 385; Lipsk. in Tr. Bot. sada, XXVI, 577; 
Kryl. Fl. Zap. Sib IX, 2323. - D. turkestanicum Gand. in BuU. Soc. Bot. Franc. LXV 
(1918) 66. - Ic: Laxm. 1. c. tab. 29, fig. 3; Bot. Mag. 25, tab. 1009; Pall. Fl. Ross. tab. 
112; Ik.-Gal. and Il'in., op. cit. Fig. 1. 



302 



Perennial; rhizome horizontal or ascending, 0.5-1 cm thick, branching above, producing 
few fertile stems and bundles of radical leaves; stems erect or ascending at base, simple, 
puberulent, more densely in upper part, 10-30 cm long; petioles IVi-l times as long as 
blade, dilated at base; leaf-blade oblong-elliptic or oblong, ovate, obtuse, cordate at base, 
crenate at margin, 2-8 cm long, 1-4 cm wide, more or less hairy beneath along the veins, 
subglabrous above; cauline leaves 3-4 pairs, short-petioled, ovate or orbicular-ovate, 
smaller; flowers short-pediceled in a dense subcapitate inflorescence, sometimes with a 
distant lower whorl; bracts obovate, cuneately tapering toward base and here entire, cut 
in upper half into teeth of which the middle one ovate-lanceolate, the others lanceolate, 
thinly acuminate, slightly exceeding or about as long as calyx, this covered with long fine 
hairs, nearly 2-Upped; middle lobe of upper lip ovate, mucronate, IVi-l times as long as 

452 the broadly lanceolate aristate lateral lobes; lobes of lower lip narrowly lanceolate; corol- 
la 35-45 mm long, intensely blue, puberulent outside, long-hairy within on lobes of upper 
lip and at base of middle lobe of lower lip; upper fip cut to 1/3 into semiorbicular lobes; 
lower lip IVz times as long as the upper, the middle lobe reniform, 2-3 times as broad as 
lateral lobes; nutlets ovoid, black, ca. 3 mm long. June-July. 

Alpine meadows, near glaciers, on rocks and stony slopes. — West Siberia: Alt.; East 
Siberia: Ang. -Say. , Dau., Lena-Kol. (Kalarskii district); Centr. Asia: Dzu.-Tarb., T. Sh., 
Pam.-Al. (Alai Range). Gen distr.: Mong., Ch. (Sinkiang). Described from Altai. Type 
in London. 

Note. The synonymy of the genus is very confusing. In establishing this species, Lin- 
naeus refers to the brief description of the plant supplied to him by J. GmeUn. The geo- 
graphical indications given by GmeUn in "Flora sibirica" (III, 234) are: "Intra valles 
rupium Bargusinensium et earum quae existunt ad Maiam et Indomam b. Stellerus obser- 
vavit." A species widely distributed in the Barguzin mountains is better known as D. al- 
taiense Laxm., while a different species, determined as D. stellerianum Hiltebr., occurs 
in Maia and Indom. We propose to retain the Linnaean name for the plant that grows 
extensively in Siberia and in the mountains of Central Asia. 

Economic importance. The plant is beautiful in flower and could be used as ornamental. 

9. D. formosum Gontsch. in Bot. mat. gerb. Bot. inst. AN SSSR, VII, 5 (1938) 101. 

Perennial; stems erect, glabrous, 4-angled, obscurely sulcate, 20-60 cm long, to 4 mm 
thick at base; radical leaves with petiole 9-13 (28) cm long, broadly triangular-ovate or 
oblong-ovate, 6-10 cm long, cordate at base, rounded at apex, crenate, glabrous, bright 
green; lower cauline leaves with petioles shorter than blade, ovate, cordate at base, 3-5 cm 
long, rounded at apex, crenate; upper leaves subsessile, subcordate at base; floral leaves 
sessile, broadly ovate, obtuse at apex, dentate, 2-2.5 cm long; inflorescence dense, ovaloid 
or subglobose, 7-10 cm long; outer bracts suborbicular, the upper oval, acute, with den- 
tate margin, the teeth with a purple awn 1-2.5 mm long; inner bracts oblanceolate, acute, 
with long aristate teeth in upper half; bracteoles lanceolate, acuminate, pubescent, 1.5- 
3 mm long; pedicels 2-3 mm long, hairy; calyx tubular, 20-22 mm long, puberulent and 

453 sparsely punctate-glandular, nearly 2-hpped; upper lip less deeply cleft and hence its lobes 
slightly shorter than the lanceolate lobes of lower lip; middle lobe of upper lip much 
broader and slightly longer than the others, ca. 8 mm long, the others ca. 7 mm long; 
corolla 3-4.3 cm long, pink-lilac, hairy outside, the tube 14-18 mm long, narrow in lower 



303 



part, inflated above, 8-10 mm long, 2-lobed at apex, with rounded lobes; lower lip slight- 
ly longer than the upper, 3-lobed, the middle lobe reniform, ca. 5 mm long, 8- 10 mm 
wide, emarginate, 2-3 times as wide as lateral lobes; filaments hairy. June-July. 

Subalpine belt, damp meadows. — Centr. Asia: Pam.-Al. Endemic. Described from 
Gissar Range. Type in Leningrad. 

10. D. imberbe Bge. Suppl. Fl. Alt. (1836) 50; Ldb. Fl. Ross. Ill, 385; Lipsk. in Tr. 
Bot. sada, XXVI, 579; Kryl. Fl. Zap. Sib. VIII, 2322. - D. altaiense Hiltebr. Monogr. 
Dracoceph. 50, 75, non Laxm. — D. imberbe var. laniflorum Herd, in Bull. Soc. Nat. 
Mosc. XLI, 1 (1868) 383. - D. laniflorum Rupr. in Mem. Acad. Petersb. VII ser. XIV, 
4 (1869) 65. - D. alberti Rgl. in Tr. Bot. sada, VI (1880) 362. - Ic: Hiltebr. 1. c. tab. 
13; Gartenfl.XXXI, tab. 1080(1882). -Exs.: Herb. Fl. As. Med. No. 207. 

Perennial; rhizome 4-10 mm thick, ascending or creeping; stems few, erect or ascend- 
ing, simple, with short fine hairs, 10-30 cm long; in addition to fertile stems there are 
bundles of radical leaves; leaves orbicular-cordate or orbicular-reniform, obtuse at apex, 
with large rounded teeth at margins, 1-4 cm long and nearly as wide, glabrous or very 
sparingly hairy above, densely pubescent or sometimes finely grayish-tomentose beneath; 
radical leaves with basally dilated petioles 2-4 times as long as blade; cauline leaves 3 pairs, 
the lower long-petioled, the middle with petioles as long as blade or shorter, the upper- 
most pair (at base of inflorescence) with very short petioles; flowers in verticillasters, sub- 
sessile, crowded at summit of stem in a dense ovoid inflorescence, one whorl in axils of 
lower pair of leaves; bracts shorter than calyx, bluish, obovate, cut into ovate-lanceolate 
or lanceolate aristate lobes, cuneate or entire at base; calyx puberulent outside, 1 5-1 8mm 
long, obscurely 2-lipped, teeth subequal, lanceolate, aristate, half as long as tube, the 
upper half as broad again as the others; corolla 25-30 (40) mm long, dark blue, pubescent 
outside, glabrous within or sometimes with sparse short hairs at base of lower lip; upper 
Up cut to 1/3 into 2 semiorbicular lobes; lower Up half as long again as the upper, the 
454 reniform middle lobe 3-3.5 times as wide as the lateral lobes; style exserted from upper 
lip, nutlets ovoid, obscurely trihedral, 4 mm long, 2 mm wide. June-July. 

Alpine belt, on rocks, taluses, stony and gravelly mountain tundras, old moraines near 
glaciers, sometimes alpine meadows. — West Siberia: Alt.; East Siberia: Ang.-Say.; Centr. 
Asia; Dzu.-Tarb., T.Sh., Pam.-Al. (Altai and Turkestan ranges). Gen.distr.: Ch. (Sin- 
kiang). Described from Altai. Type in Leningrad. 

11. D. stellerianum Hiltebr. Monogr. Dracoceph. (1805) 21, 36, 65. - D. chamaedry- 
folium Turcz. ex Benth. in DC. Prodr. XII (1848) 399. - D. grandiflorum ^. minus 
Benth. 1. c. 399. - D. grandiflorum a. latifolium et ^. angustifolium Ldb. Fl. Ross. 
111(1846-1851)386. 

Perennial; rhizome ascending, branching above; stems few, ascending at base, puberu- 
lent, more densely so in upper part, 6-20 cm long; leaves with patent-hairy petiole shorter 
than blade, broadly ovate or orbicular, subcordate or truncate or short-cuneate at base, 
with crenate to obtusely dentate ciUate margins, puberulent or subglabrous on both sides, 
1.2-3.5 cm long, 0.8-2 cm wide; floral leaves narrower, less dentate, sometimes entire; 
flowers short-pediceled, in verticillasters at end of stem forming an oblong inflorescence, 
sometimes with one remote lower verticillaster; bracts oblong or ovate, entire, long-ciliate, 



304 



subulate-pointed at apex; calyx ca. 13 mm long, 2-lipped, middle tooth of upper lip broad- 
ly ovate, 2-3 times as broad as the other lanceolate teeth; corolla ca. 30 mm long, densely 
pubescent outside, azure-blue. June-July. 

Stony slopes and taluses in alpine and subalpine belts. — East Siberia: Lena-KoL; Far 
East: Okh., Uss. (Dzhugdzhur Range). Endemic. Described from Siberia (between 
Yakutsk and Okhotsk). Type in Moscow (?). 

12. D. fragile Turcz. ex Benth. Lab. gen. et sp. (1832-1836) 495; Bge. Suppl. Fl. Alt. 
50; Ldb. Fl. Ross. Ill, 386; Turcz. Fl. baic.-dahur. II, 379. - Ic: Turcz. in Bull. Soc. 
Nat. Mosc. XXIV, 4, tab. 8. 

Perennial; rhizome long, 3-4 mm thick, ascending, branching above; stems few to 
many, ascending at base, later erect, covered with soft long spreading hairs, more densely 
so in upper half; leaves with petioles shorter than blade, ovate, 1-4.5 cm long, 0.5-2 cm 
wide, obtuse at apex, abruptly or gradually tapering at base, entire or crenate in upper 
half, subglabrous on upper side, prominently veined beneath, beset with white thickish 
455 hairs, the margin with long fine ciUa or subglabrous; flowers short-pediceled, in few- 
flowered verticillasters at summit of stem forming an oblong inflorescence, lower whorls 
distant; bracts entire, ovate or narrowly ovate, 12 mm long, 4 mm wide, v:"'^"s; calyx 
15-17 mm long, middle tooth of upper lip twice as wide as the lanceolate lateral teeth, 
all teeth subulate-pointed; corolla 25-30 mm long, whitish-yellowish, lower lip dark blue, 
with azure spots. July. 

Stony and gravelly riverbanks, often on bald peaks. — East Siberia: Yenis. (vicinity of 
Yeniseisk), Ang.-Say., Dau. Gen. distr.: Mong. (Lake Khubsugul). Described from Lake 
Khubsugul (Kosogol). Type in Leningrad. 

Note. In his monograph on the genus Dracocephalum, Hiltebrandt presents a 
drawing (No. 10) of D. grandiflorum. According to Bunge (Suppl. Fl. Alt., p. 50) the 
drawing may possibly refer to D. fragile Turcz. We have refrained from citing Hilte- 
brandt's key above because we doubt the accuracy of Bunge's supposition. 

13. D. multicolor Kom. in Izv. Bot. sada, XVI (1916) 177; Kom. and Alls. Opred. 
rast. Dal'nevost. kraya, II, 903. 

Perennial; rhizome ascending, to 1.5 cm thick, branching above; stems many, more or 
less suffused with violet, covered with short retrorse hairs, branching, 10-50 cm long; 
lower leaves soon withering; middle leaves ovate or ovate-oblong, subcordate or rounded 
or short-cuneate at base, obtuse at apex, pectinate-dentate with obtuse slightly antrorse 
teeth, the upper side glabrous, with depressed veins, the lower side white-tomentose, pro- 
minently veined; petioles 0.5-1 cm long, hairy on one side; blade 1.5-4 cm long, 0.8- 
2.5 cm wide; leaves of short axillary shoots smaller; flowers with pedicels 2-3 mm long, 
in few-flowered verticillasters at end of stem and branches, forming a rather dense oblong 
inflorescence; bracts obcuneate, gradually tapering to short petiole, sparingly pubescent, 
at margins with 4 lanceolate teeth terminating in long subulate slightly hairy point, the 
central tooth broadly ovate, abruptly subulate-pointed; calyx ca. 17 mm long, usually 
purple-violet, with scattered punctate glands, 2-Hpped; middle tooth of upper Up broadly 
ovate, mucronate, 2-3 times as broad as the other lanceolate acuminate teeth; corolla 
3 5-50 mm long, white -hairy outside, with violet spots on Umb; upper lip much longer than 
the lower. July-August. 



305 



456 Stony slopes and taluses, rocks, sometimes in oak forests. — Far East: Uss. Endemic. 
Described from Tetyukhe River. Type in Leningrad. 

14. D. krylovu Lipsky in Tr. Bot. sada, XXIV, 2 (1905) 121 ; Kryl. Fl. Zap. Sib. IX, 
2331. 

Perennial, stems few, woody at base, procumbent or ascending, 15-25 cm long, cover- 
ed with short retrorse hairs, branching from base with remote short branches, only the 
upper fertile; leaves glabrous above, dark green, short-hairy beneath, with numerous, very 
small glands; lower leaves with petiole half as long as blade, broadly ovate, 10-20 mm 
long, 6-12 mm wide, obtuse, large-toothed or incised, each margin with 3-5 obtuse or 
rounded antrorse teeth; leaves on branches oblong-elUptic or oblong, much narrower than 
the cauline; flowers short-pediceled, in 2-5-flowered verticillasters forming a loose Inflo- 
rescence at the ends of stem and upper branches, 3-4 cm long; bracts 1/5-1/3 as long as 
calyx, ovate or eUiptic, acuminate, entire or the lower with 1 tooth at each side, tapering 
to petiole at base; calyx ca. 12 mm long, puberulent, with numerous punctate glands; up- 
per lip cut to 2/3 into lobes, the middle lobe ovate, aristate, the lateral lanceolate, aristate, 
3-3^/ " _ -^ as broad as middle lobe and slightly shorter; teeth of lower lip lanceolate; 
corolla 30-35 mm long, azure, covered outside with fine and rather long hairs, hairy with- 
in only at base of lower lip; upper lip shallowly cut into semiorbicular lobes; lower Up 
half as long again, the reniform middle lobe obscurely crenate, 3^/4 times as broad as 
lateral lobes. July. 

Rocks. — West Siberia: Alt. (Mrassu River near Chelei). Endemic. Described from site 
indicated. Type in Leningrad. 



Subsection 3. Idiodracontes Briq. in Pflanzenfam. IV, 3a and 3b (1897) 239, ex 
parte. - Section Moldaviae Benth. Lab. gen. et sp. (1832-1836) 496, ex parte. — Leaves 
elongate, entire or few-toothed. 



Series 1. Fruticulosa Schischk. — Floral leaves indistinguishable in shape from 
cauhne leaves. 

15. D. fruticulosum Steph. in Willd. Sp. pi. Ill (1800) 152; Hiltebr. Monogr. Draco- 
ceph. 29, 65; Ldb. Fl. Ross. Ill, 386; Turcz. Fl. baic.-dahur. II, 380. - Ic: Hiltebr. 1. c. 
tab. 3. 
457 Perennial; rhizome thick, 5-10 mm in diameter, ascending, dark brown, branching 
above, woody; stems many, erect, woody in lower part, covered with short retrorse hairs, 
15-30 cm long; leaves ovate or lanceolate-ovate, 3-veined, entire or with 1-2 teeth on each 
margin, subulate-mucronate, covered with short hairs and numerous punctate glands, 6- 
12 mm long, 2-4 mm wide, abruptly terminating in a subulate point 0.5-1 mm long; 
abbreviated axillary shoots with smaller leaves; flowers short-pediceled, crowded at ends 
of stems in the axils of upper floral leaves and forming rather dense inflorescence 1-3 cm 
long, 1 .5-2 cm wide, sometimes with one distant verticillaster below; bracts leaflike, 
usually much shorter than calyx; calyx violet-tinged, ca. 10 mm long, minutely puberulent. 



306 



slightly curved; teeth lanceolate, acuminate, awn-tipped, the upper tooth slightly broader 
than the others; corolla half as long again as calyx, ca. 15 mm long, densely covered out- 
side with soft white hairs; anthers blackish-purple, slightly exserted from corolla; nutlets 
dark purple, 3.2 mm long, 1.2 mm wide. Fl. July; fr. August. 

Rocks and shrubby thickets. - East Siberia: Ang.-Say., Dau. Gen. distr.: Mong. 
Described from Siberia. Type in Berlin. 

16. D. integrifolium Bge. in Ldb. Fl. Alt. II (1830) 387; Ldb. Fl. Ross. Ill, 386; 
Lipsk. in Tr. Bot. sada, XXVI, 588; Kryl. Fl. Zap. Sib. IX, 2325. - Ic: Ldb. Ic. pi. fl. 
Ross. II, tab. 120. 

Perennial, more or less woody, the divaricately branching stems covered with grayish- 
brown peeUng bark; herbaceous branches erect or ascending, covered with very short 
appressed retrorse hairs, 15-60 cm long, leaves lanceolate or ovate-lanceolate, acute, sub- 
sessile or tapering to short petiole, 3-veined, entire or with 1-2 teeth on each margin, glabrous 
or short-ciUate at margins, 1.5-3 cm long, 3-8 mm wide; abbreviated axillary shoots vvith 
smaller leaves; flowers in verticillasters, in threes on short pedicels in axils of upper floral 
leaves, forming a rather dense inflorescence 2-5 cm long and ca. 2.5 cm wide; bracts slight- 
ly shorter than calyx, eUiptic, tapering at base, long-acuminate at apex, at sides with 1 pair 
of fiUform lobes 2-3 times longer than width of bracts; calyx short-hairy, often dingy 
violet, slightly curved, nearly 2-hpped; middle lobe of upper hp broadly obovate or sub- 
orbicular, aristate, 2-3 times as wide as the lanceolate lateral lobes, these about as long as 
lobes of lower lip; all calyx -teeth with very prominent transverse anastomoses; corolla 
bluish-lilac, puberulent outside and at base of lower Up within, 15-18 mm long; upper lip 
cut to 1/3 into semiorbicular lobes; lower Hp half as long again, the middle lobe reniform, 
emarginate, obtusely crenate at margins, nearly SYz times as wide as the rounded-ovate 
lateral lobes; style scarcely exserted from upper lip; nutlets dark brown, obscurely 3- 
angled, ovoid, 2.5 mm long, ca. 1 .5 mm wide. June-July. 

Stony, gravelly and grassy slopes, broadleaved forests and thickets at altitudes of 900- 
2000 m. - West Siberia: Alt. (SW); Centr. Asia: Dzu.-Tarb., T. Sh. Gen. distr.: Ch. 
(Sinkiang). Described from Altai. Type in Leningrad. 



Series 2. Nutantia Schischk. — Leaves more or less evenly toothed; floral leaves 
entire. 

17. D. nutansL.Sp.pl. (1753)596; Hiltebr. Monogr. Dracoceph. 54, 76; Ldb. Fl. 
Ross. Ill, 387; Turcz. Fl. baic.-dahur. II, 381 ; Lipsk. in Tr. Bot. sada, XXVI, 592; Kryl. 
Fl. Zap. Sib. IX, 2325. — D. nutans var. alpinum Kar. et Kir. in Bull. Soc. Nat. Mosc. 
XV (1842) 424. - D. alpinum Turcz. in Bull. Soc. Nat. Mosc. XXIV (1851) 383, non 
Sahsb. — D. microphyllum Turcz. 1. c. (1851) 384. — Zornia nutans Moench, Meth. 
(1794) 41 1. - Ic: Pall. Fl. Ross. tab. 115. 

Perennial; stems soUtary or few or many, simple or branching, puberulent, 20-70 cm 
long; radical and lower cauUne leaves with puberulent petiole longer than blade, ovate, 
cordate at base, obtuse at apex, obtusely subulate-toothed at margin, 1.5-4 cm long, 1- 
3 cm wide; middle cauUne leaves lar