Morphometric analysis of the Taxus wallichiana
complex (Taxaceae) based on herbarium material
MICHAEL MÖLLER
1
*†, LIAN-MING GAO
2
†, ROBERT R. MILL
1
, DE-ZHU LI
2
,
MICHELLE L. HOLLINGSWORTH
1
and MARY GIBBY
1
1
Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh EH3 5LR, Scotland, UK
2
Key Laboratory of Biodiversity and Biogeography, Kunming Institute of Botany, Chinese Academy of
Sciences, 132 Lanhei Road, Kunming, Yunnan, 650204, China
Received July 2006; accepted for publication May 2007
We used morphological and geographical data of 128 herbarium specimens of the Taxus wallichiana complex in
eastern Asia to investigate their utilization in discriminating and identifying taxa included in the complex. One bud
scale and 26 leaf characters were used to separate T. fuana, T. wallichiana var. wallichiana, T. wallichiana var.
mairei, T. wallichiana var. chinensis and T. sumatrana by K-means clustering and dendrograms using Ward’s
distance. Out of the 27 characters examined 21 were found to be well correlated with geographical patterns.
T. fuana was morphologically the most distant taxon, while T. sumatrana clustered among the T. wallichiana
varieties. After correcting misidentifications of the specimens in T. wallichiana, its varieties occupied discrete
geographical ranges, except for some limited sympatry of varieties mairei and chinensis east of the Tanaka–
Kaiyong line and the Sichuan Basin, China. Our analysis demonstrates the importance of consistency in character
selection and definition in the identification of morphologically difficult taxa and the power of combining
morphometric and geographical data in clarifying their spatial distribution. © 2007 The Linnean Society of
London, Botanical Journal of the Linnean Society, 2007, 155, 307–335.
ADDITIONAL KEYWORDS: China – geographical distribution – PCA – taxonomy – Taxus fuana – Taxus
sumatrana – variety delimitation.
INTRODUCTION
The taxonomy of the genus Taxus L. is complex and
inconsistent over large areas of its distribution range.
The main problem in this genus arises from a lack of
discrete (‘hard’) vegetative characters. This is aggra-
vated by the fact that herbarium specimens are often
sterile and so a whole suite of potentially suitable
generative morphological features cannot be used. A
large number of vegetative characters are hard to find
and constitute mainly characters of the leaf. The
majority of these are continuous and often difficult to
define unequivocally. Terminological problems and
interpretation of character states can further hamper
taxonomic communication. Proper identification is
paramount to comply with CITES regulations
because all Taxus species occurring in China have
been listed as first-class protected plants by the
central government of China since 1999 and were
listed in Appendix II of CITES since 1995 (http://
www.cites.org/eng/app/appendices.shtml#10).
Taxus includes four species in North America
(T. brevifolia Nutt., T. canadensis Willd., T. floridana
Nutt., T. globosa Schlecht.), one in Europe and
western Asia (T. baccata L.), and several in east and
south-east Asia. The taxonomy of these latter taxa is
still in flux. De Laubenfels (1988) included all the
Chinese and Indo-Chinese yews, except for T. cuspi-
data Siebold & Zucc., in T. sumatrana (Miq.) de
Laub., but other authors have not taken such a broad
view. Li & Fu (1997) established Taxus fuana Nan Li
& R. R. Mill for specimens from Xizang (Tibet), and
from central Nepal westwards to Pakistan and
Afghanistan. Fu, Li & Mill (1999), while retaining
T. cuspidata and T. fuana, recognized three varieties
*Corresponding author. E-mail: m.moeller@rbge.org.uk
†These two authors contributed equally to the study.
Botanical Journal of the Linnean Society, 2007, 155, 307–335. With 9 figures
© 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 155, 307–335 307
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