Table Of Content"' CCG. m R S O N & D. G. LONG
ANIC GARDEN EDLNBURGH
OF BHUTAN
FLORA OF
BHUTAN
INCLUDING A RECORD OF PIANTS
FROM SIIUUM AND DARJEEI,ING
VOLUME 2 PART 3
A. J. C. GRIERSON & D. G. LONG
EDITED BY L. S. SPRINGATE
Illustrations bv M. Bates, Chen Yo-Jiun, C. Oliver,
L. Olley, G. Rodrigurs
& M. Tebbs
ROYAL BOTANIC GARDEN EDINBURGH
ROYAL GOVERNMENT OF BHUTAN
200 1
This volu~nei s dedicated to the memory of
Andrew J. C. Grierson (19 29-1 990) und Pauline Grierson (nke Shillubeer,
1927-1 996)
Published by the Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh
EH3 5LR, UK and the Royal Government of Bhutan
0R oyal Botanic Garden Edinburgh and Royal Govern~nento f Bhutan 2001
ISBN 1 87229 1 93 7
Typeset, printed and bound by
The Charlesworth Group, Huddersfield. UK. 01484 517077
CONTENTS
List of figures ........................................................................................................ iv
Introduction ...................................................................................................... 1035
Acknowledgements ........................................................................................... 1035
Abbreviations .................................................................................................... 1036
ANGIOSPERMAE continued
171. Solanaceae (R.R. Mill ) ............................................................................1 037
172. Buddlejaceae (S.J. Rae) ...........................................................................1 078
173. Scrophulariaceae (R.R. Mill ) ..................................................................1 082
174. Bignoniaceae (E. Aitken) .........................................................................1 236
175. Acanthaceae (J.R.I. Wood) ....................................................................1. 243
176. Pedaliaceae (E. Aitken) ...........................................................................1 293
177. Gesneriaceae (O.M. Hilliard ) .................................................................1. 296
178 . Orobanchaceae (S. J . Rae) ......................................................................1..3 30
179. Lentibulariaceae ( H.J. Noltie) ................................................................. 1334
1 80. Phrymaceae ( L.S. Springate) ................................................................... 1340
18 1 . Plantaginaceae ( L.S. Springate) ............................................................... 1341
182. Caprifoliaceae ( R.A. Clement). Loniwra (P.W. Meyer) ......................... 1343
182a. Carlemanniaceae (D.G. Long & J . Wright) ............................................ 1362
183. Valerianaceae (R.A. Clement) ...............................................................1. 363
1 84. Dipsacaceae ( R.A. Clement ) ..................................................................1. 368
185. Morinaceae (R.A. Clement) .................................................................... 1372
186. Campanulaceae ( R.A. Clement) .............................................................. 1375
187. Compositae ( A.J.C. Grierson & L.S. Springate) .....................................1 397
Bibliography ...................................................................................................... 1633
Index of botanical names .................................................................................. 1642
Index of common names ................................................................................... 1672
Index of families in published parts of the Floru ............................................. 1674
FIGURES
90 . Solanaceae ................................................................................................. 1053
9 1 . Solanaceae ................................................................................................. 1069
92 . Solanaceae and Buddlejaceae .................................................................... 1075
93 . Scrophulariaceae ........................................................................................ 1089
94 . Scrophulariaceae ........................................................................................ 1115
95 . Scrophulariaceae ........................................................................................ 1127
96 . Scrophulariaceae ........................................................................................ 1139
97 . Scrophulariaceae ........................................................................................ 1153
98 . Scrophulariaceae ........................................................................................ 1181
99 . Scrophulariaceae ......................................................................................1.2. 07
100. Scrophulariaceae ........................................................................................ 1223
10 1 . Scrophulariaceae and Bignoniaceae .......................................................... 1237
102. Acanthaceae ............................................................................................1..2.4 9
103. Acanthaceae ............................................................................................... 1267
104. Acanthaceae ............................................................................................... 1269
105. Acanthaceae ............................................................................................... 1277
106. Pedaliaceae and Gesneriaceae ................................................................... 1295
107. Gesneriaceae .............................................................................................. 1307
108. Gesneriaceae ..........................................................................................1..3..2 5
109. Gesneriaceae and Orobanchaceae ...........................................................1..3 29
1 10. Lentibulariacae, Phrymaceae and Plantaginaceae ..................................... 1335
1 1 1 . Caprifoliaceae ............................................................................................ 1351
1 12. Caprifoliaceae ............................................................................................ 1361
1 13. Carlemanniaceae and Valerianaceae .......................................................... 1365
1 14 . Dipsacaceae, Morinaceae and Campanulaceae .........................................1 371
11 5. Campanulaceae .......................................................................................... 1381
11 6. Campanulaceae .......................................................................................... 1391
1 17. Compositae ................................................................................................ 1423
1 18 . Compositae: Mutisieae .............................................................................. 1425
1 19. Compositae: Cirsieae ................................................................................. 1429
120. Compositae: Lactuceae .............................................................................. 1455
12 1 . Compositae: Lactuceae .............................................................................. 1465
122. Compositae: Lactuceae .............................................................................. 1471
123. Compositae: Vernonieae ............................................................................ 1491
124. Compositae: Inuleae .................................................................................. 1493
125. Compositae: Plucheae and Gnaphalieae ................................................. 1509
126. Compositae: Gnaphalieae .......................................................................... 1513
127. Compositae: Calenduleae and Astereae .................................................... 1527
128. Compositae: Astereae ................................................................................ 1543
129. Compositae: Anthemideae ......................................................................... 1 553
130. Compositae: Anthemideae ......................................................................... 1557
13 1 . Compositae: Anthemideae ......................................................................... 1561
132. Compositae: Anthemideae ......................................................................... 1563
133. Compositae: Senecioneae ..........................................................................1 591
134 . Compositae: Senecioneae and Helenieae ................................................... 1595
135 . Compositae: Heliantheae ........................................................................... 1609
136. Compositae: Heliantheae ........................................................................... 1613
13 7 . Compositae: Eupatorieae and Cavea ........................................................ 1629
INTRODUCTION
by David G. Long, Henry J. Noltie & Lawrence S. Springate
This part of the Flora oJ'Bhufcin completes the treatment of the Dicotyledons
and includes the important families Solanaceae, Scrophulariaceae, Acanthaceae,
Gesneriaceae, Caprifoliaceae and Compositae. The first drafts of many of the
accounts were completed between 1990 and 1994. although funding for
publication has only recently become available through the generous
support of Danida. Consequently the early drafts (particularly Compositae,
Scrophulariaceae and Solanaceae) have undergone thorough review recently.
Amongst these families the Compositae (also known as Asteraceae) are pre-
eminent as the largest family of Dicotyledons, encompassing 126 genera and
372 species, outnumbered in the angiosperms only by two Monocotyledon
families: Gramineae and Orchidaceae. Compositae was the special interest of
the late Andrew Grierson, whose vision and enthusiasm had a major influence
in the inauguration and running of the Floru of Bltutun from its beginning until
his untimely death in 1990. This account of Compositae stands as a tribute to
his enormous contribution. The Compositae is particularly well represented at
high altitudes in the East Himalaya; it shows many strong links with the Tibetan
flora, and for that reason does not show much endemism within the political
boundaries of the Floru. Amongst other families, Scrophulariaceae is noteworthy
as containing the largest single genus in the Flor-u of Bhuturz: Pediclr1tr.i.~.w ith
76 species represented. Many of these are local East Himalayan endemics.
Several of the family accounts were contributed by the late Rose A. Clement
(nee King) and these constitute her remaining posthumous publications.
ACKNOWLEDGEMENTS
The editors of the F1oi.u qf Bkutori wish to acknowledge our deep gratitude
to Danida, the Overseas Aid Division of the Royal Danish Ministry of Foreign
AfEziirs, for the Flora of Bhutan Completion Project. The present volume is
published under their auspices: this support has covered editing and printing
costs and the preparation of illustrations. We are also pleased to acknowledge
the continued interest and help of the Royal Government of Bhutan, in particu-
lar the Minister of Agriculture, Lyonpo Dr Kinzang Wangdi and Yeshey Dorji
of the National Biodiversity Programme.
We also wish to thank the authors of accounts contained in this part. most of
whom have had to put up with a very lengthy delay in publication of their work;
this delay was due to constraints of funding for publication and we are grateful
for their forbearance: Eona M. Aitken, Olive M. Hilliard, Paul W. Meyer. Robert
R. Mill, Sally J. Rae. John R.I. Wood and Jenny Wright. Thanks are also due
1035
to the artists for the drawings they have contributed: Mary Bates, Chen Yo-Jiun,
Christina Oliver, Louise Olley. Glenn Rodrigues and Margaret Tebbs.
The Directors and Curators of the Herbaria at the Royal Botanic Gardens,
Kew and the Natural History Museum, London, have continued to provide
facilities for study visits and many very large loans of specimens. Some interes-
ting nineteenth-century collections were supplied by Manchester Museum. The
staff of these organisations are also thanked for taxonomic advice and assistance,
as are the following specialists for advice on particular groups: Charles Jeffrey
(Compositae), Jan Kirschner (Tcrrcrscrcum), Valery Malecot (Vibu~.rzurna) nd Peter
Taylor (Utriculor.icr).
We also wish to thank Eona Aitken, Norma Gregory and Kim Howell for
technical and editorial help with this part.
As this part concludes both the treatment of Dicotyledons, and is also the final
outstanding part of the Floru to be written at the Royal Botanic Garden
Edinburgh, it is appropriate now to acknowledge those colleagues and friends
in Britain, Bhutan and elsewhere who have contributed to the Flora ojBlzutan
project over the years in many different ways, as collectors, contributors of
revisions, referees, taxonomic and nomenclatural advisers, horticulturists and
general enthusiasts for Himalayan plants and Bhutan. We are grateful to them
for their continued support: Peter Baxter, Prof. Stephen Blackmore, Simon
Bowes Lyon, B.L. Burtt, James Cullen, Lyonpo C. Dorji, Yeshey Dorji, Norma
Gregory, Chris Grey-Wilson, D.M. Henderson, Prof. David Ingram, Roy
Lancaster, Ron J.D. McBeath, Prof. David Mann, Robert R. Mill, Chris Parker,
Tony Peers, Rebecca Pradhan, Sally J. Rae, Keith Rushforth, Tony Schilling,
Ian W.J. Sinclair, Philippe de Spoelberch, W.T. Stearn, Sonam Tshering, Tandin
Wangdi, Mark F. Watson and John R.I. Wood.
We wish also to correct an earlier omission of acknowledgement to Prof. David
Mabberley, for refereeing Andrew Grierson's account of Meliaceae in Volume
2 Part 1 ( 1991 ), which contained several previously unpublished observations
and conclusions contributed by Prof. Mabberley in his review of the manuscript.
ABBREVIATIONS
Abbreviations for languages and dialects of common names of plants used in
this volume are:
Dz: Dzongkha language ( W Bhutan and official language)
Eng: English
Med: Bhutanese medicinal name
Nep: Nepali
Sha: Shachop dialect (E Bhutan)
For other abbreviations, e.g. botanical authorities, users are referred to
Volume 1 Part 1, p. 34.
