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Morphological Comparisons of Taiwan Native Wild Tea Plant (Camellia sinensis (L.) O. Kuntze forma formosensis Kitamura) and Two Closely Related Taxa Using Numerical Methods

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Camellia sinensis (L.) O. Kuntze forma formosensis Kitamura is generally referred to as the tea plant growing naturally in mid-elevation mountains of Taiwan. Several taxonomic treatments have been published for this plant in the past, but some contradictory results have been obtained. To assess the taxonomic position of the wild tea plant and explore its relationship with two other closely related taxa, C. sinensis var. sinensis and C. sinensis var. assamica, 16 vegetative and 11 floral characters were examined on 165 OTUs. The data were analyzed using cluster analysis and nonlinear principal components analysis. All cluster phenograms consistently separated the native wild tea plant from two other related taxa. Conversely, pronounced admixture between C. sinensis var. sinensis and C. sinensis var. assamica was present. The nonlinear principal components analysis indicated that the surface features of buds and ovaries are two diagnostic characters. Based on the present study, it is proposed that the Taiwan native wild tea plant might deserve recognition as a distinct species.
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Taiwania, 52(1): 70-83, 2007
Morphological Comparisons of Taiwan Native Wild Tea Plant
(Camellia sinensis (L.) O. Kuntze forma formosensis Kitamura)
and Two Closely Related Taxa Using Numerical Methods
Mong-Huai Su(1), Chih-Hua Tsou(2) and Chang-Fu Hsieh(1,3)
(Manuscript received 30 August, 2006; accepted 22 December, 2006)
ABSTRACT: Camellia sinensis (L.) O. Kuntze forma
f
ormosensis Kitamura is generally referred to as
the tea plant growing naturally in mid-elevation mountains of Taiwan. Several taxonomic treatments
have been published for this plant in the past, but some contradictory results have been obtained. To
assess the taxonomic position of the wild tea plant and explore its relationship with two other closely
related taxa, C. sinensis var. sinensis and C. sinensis var. assamica, 16 vegetative and 11 floral
characters were examined on 165 OTUs. The data were analyzed using cluster analysis and nonlinear
principal components analysis. All cluster phenograms consistently separated the native wild tea plant
from two other related taxa. Conversely, pronounced admixture between C. sinensis var. sinensis and C.
sinensis var. assamica was present. The nonlinear principal components analysis indicated that the
surface features of buds and ovaries are two diagnostic characters. Based on the present study, it is
proposed that the Taiwan native wild tea plant might deserve recognition as a distinct species.
KEY WORDS: Camellia sinensis forma formosensis, Native wild tea plant, Taiwan, Numerical
taxonomy.
INTRODUCTION
The native wild tea plant, Camellia sinensis (L.)
O. Kuntze forma formosensis Kitamura, is generally
referred to the tea plant growing naturally in the
broad-leaved forests at mid-elevations of the Central
Mountain Range (CMR) of Taiwan (Wu et al., 1970;
Lai et al., 2001). According to collection data from
herbaria, it is widely distributed at elevations from
900 to 1800 m in the central, southern and eastern
regions of Taiwan, including Nantou, Chiayi,
Kaohsiung, Pingtung and Taitung County (Fig. 1).
The native wild tea plant in Taiwan was first
described in 1717 during the Ching Dynasty. In 1724
a political officer, Su-Jien Huang, mentioned it again
in his report to Taiwan. Therefore the native wild tea
plant has been known to exist in Taiwan for nearly
300 years.
Tea is one of the most popular beverages in the
world. Taxonomically, economic tea plants belong to
the genus Camellia of family Theaceae. There are
two main varieties of tea plants: C. sinensis var.
__________________________________________
1. Institute of Ecology and Evolutionary Biology, National Taiwan
University, 1, Sec. 4, Roosevelt Rd., Taipei 106, Taiwan.
2. Institute of Plant and Microbial Biology, Academia Sinica, 128,
Sec. 2, Academia Rd., Taipei 115, Taiwan.
3. Corresponding author. Email: tnl@ntu.edu.tw
sinensis and C. sinensis var. assamica (Masters)
Kitamura (Kitamura, 1950; Sealy, 1958; Ming,
2000). The former is usually processed for green teas
(un/hemi-fermented teas), while the latter is for black
teas (fully fermented teas). In Taiwan, tea plants are
also important crops and have been cultivated for
over 200 years. Nevertheless, the cultivated tea plants
(C. sinensis var. sinensis and C. sinensis var.
assamica) were imported, and have nothing to do
with that native wild tea plant (Hasimoto, 1967; Lai
et al., 2001). During the Japanese Colonized Period,
the government sought to popularize and encourage
the cultivation of the native wild tea plants, but for
some reasons it had never been done (Shih, 1995). As
a result of the 50-year-long effort, a new cultivar of
black tea, “TTES No. 18” or "Red Jade", was finally
released by the Tea Research and Extension Station
and extended to the farmers in 1999. The “TTES No.
18” is just an artificial hybrid between C. sinensis
var. assamica (maternal) and C. sinensis f.
formosensis (paternal). This tea smells like natural
cinnamon and fresh mint and is quite popular among
consumers.
Ancient people in the Mt. Alishan area of central
Taiwan called the native wild tea plant "Shuen-Cha"
which means teas from celestial beings. The folk
name is still used today by their descendants. In
March, 2007 Su et al.: Morphological comparisons of Taiwan wild tea plant and related taxa 71
Fig. 1. Distribution map of C. sinensis f. formosensis based on
authors' samples and materials from TAI, TAIF and PPI herbaria. 1:
Nantou County. 2: Yunlin County. 3: Chiayi County. 4: Kaohsiung
County. 5: Pingtung County. 6: Taitung County.
several reports on tea improvements, the native wild
tea plant was named as 'San-Cha' (means teas from
mountains) to be distinguished from the imported tea
plants (Shih, 1995; Li and Chang, 2003).
Botanical nomenclatures of the native wild tea
plant are somewhat complicated. Here we try to
clarify it in brief. For its affinity to C. sinensis var.
assamica, Masamune and Suzuki proposed Thea
assamica Masters var. formosensis Masamune et
Suzuki for the first time (Masamune, 1936).
However, Masamune didn't publish the name validly,
because there was no description provided. The
correct name first published for this plant should be
Thea formosensis Masamune et Suzuki (Suzuki,
1937). Later, this taxon was transferred as a forma
under C. sinensis (Kitamura, 1950) and named as C.
sinensis f. formosensis. In the following four decades,
however, this name has been consistently neglected
in all studies concerning the taxonomy of Camellia
(Keng, 1950; Liu and Lu, 1967; Li, 1976; Ying,
1995) until the publication of Flora of Taiwan,
second edition, in which Hsieh et al. (1996) followed
Kitamura's treatment. Ming (2000) also mentioned
the same name, but he treated it as a synonym of C.
sinensis. Although Suzuki (1937) had pointed that
native wild tea plant was "Thea assamica affinis, sed
foliis glabris" and Kitamura (1950) described that the
forma was unique by 'Foliis majoribus angustioribus
atroviridibus crassioribus', taxonomic uncertainties
of this wild tea plant still remain. There had been
several attempts to study C. sinensis f. formosensis
based on numerical methods (Wu et al., 1970, 1972;
Shih et al., 1972; Hu, 2004), however, all these
studies aimed at exploring the variation among
populations of C. sinensis f. formosensis, rather than
solving the fundamental taxonomic problems.
In the present study, the morphological variation
among C. sinensis f. formosensis and its two related
taxa, C. sinensis var. sinensis and C. sinensis var.
assamica is summarized by applying multivariate
numerical approaches. The aim is to detect
infraspecific boundaries, to identify reliable
distinguishing characters. A sound knowledge of
taxonomy is a prerequisite for the success of any
germplasm conservation program of the wild tea
plant of Taiwan.
MATERIALS AND METHODS
Materials
In total, 165 specimens were selected for
morphological study (Table 1). Sources of the
specimens used in this investigation were deposited in
the Herbarium of National Taiwan University (TAI),
Herbarium of Taiwan Forest Research Institute
(TAIF), Prof. Tzen-Yuh Chiang's Laboratory at the
National Cheng Kung University (NCKU), and
supplemented by the authors' own collection.
Scientific names given on the specimen labels and
annotated labels were tentatively used. Among 165
specimens, 72 were identified as C. sinensis var.
sinensis, 41 as C. sinensis var. assamica and 52 as C.
sinensis f. formosensis. Materials of C. sinensis f.
formosensis were collected from all natural habitats
we have known so far. Both C. sinensis var. sinensis
and C. sinensis var. assamica are not native to Taiwan.
In order to explore the variation of the two closely
related taxa, several collections from China were
made. Meanwhile, specimens from the Tea Research
and Extension Station, which represented tea plants
from India, Sri Lanka, Thailand and China, were
included.
Character measurement
In this study, each collection was designated as an
operational taxonomic unit (OTU). Specimen
duplicates were treated as one OTU. Characters were
72 TAIWANIA Vol. 52, No. 1
Table 1. List of voucher specimens included in the present study. Codes are comprised of an abbreviated scientific name, followed by a dash,
and then the collection site with a serial number. Codes with an asterisk are the specimens collected from the germplasm banks of the Tea
Research and Extension Station. TAI: Herbarium of National Taiwan University; TAIF: Herbarium of Taiwan Forest Research Institute; NCKU:
Prof. Tzen-Yuh Chiang’s Laboratory at National Cheng Kung University.
Name Collector with collection
number Site Date Herbarium Code
var. assamica M. H. Su 669 Taiwan, Taipei Co. (Burma type) 2005/10/6 Authors A-BU1*
K. S. Wang 5007 China, Yunan Prov. Elv. 1781m 2004/12/1 NCKU A-CN1
K. S. Wang 5010 China, Yunan Prov. Elv. 970m 2004/12/1 NCKU A-CN2
K. S. Wang 5016 China, Yunan Prov. 2004/12/1 NCKU A-CN3
K. S. Wang 5017 China, Yunan Prov. 2004/12/1 NCKU A-CN4
K. S. Wang 5019 China, Yunan Prov. 2004/12/1 NCKU A-CN5
K. S. Wang 5021 China, Yunan Prov. 2004/12/2 NCKU A-CN6
K. S. Wang 5024 China, Yunan Prov. 2004/12/2 NCKU A-CN7
K. S. Wang 5026 China, Yunan Prov. 2004/12/2 NCKU A-CN8
K. S. Wang 5033 China, Yunan Prov. 2004/12/2 NCKU A-CN9
K. S. Wang 5034 China, Yunan Prov. Elv. 1200m. 2004/12/3 NCKU A-CN10
K. S. Wang 5035 China, Yunan Prov. Elv. 1200m. 2004/12/3 NCKU A-CN11
K. S. Wang 5036 China, Yunan Prov. Elv. 1200m. 2004/12/3 NCKU A-CN12
K. S. Wang 5041 China, Yunan Prov. Elv. 1900m. 2004/12/3 NCKU A-CN13
K. S. Wang 5042 China, Yunan Prov. Elv. 1900m. 2004/12/3 NCKU A-CN14
K. S. Wang 5052 China, Yunan Prov. Elv. 1950m. 2004/12/4 NCKU A-CN15
K. S. Wang 5053 China, Yunan Prov. 2004/12/4 NCKU A-CN16
K. S. Wang 5054 China, Yunan Prov. Elv. 1470m. 2004/12/4 NCKU A-CN17
K. S. Wang 5058 China, Yunan Prov. Elv. 1300m. 2004/12/4 NCKU A-CN18
K. S. Wang 5066 China, Yunan Prov. Elv. 1020m. 2004/12/4 NCKU A-CN19
K. S. Wang 5072 China, Yunan Prov. Elv. 1400m. 2004/12/5 NCKU A-CN20
K. S. Wang 5076 China, Yunan Prov. Elv. 1380m. 2004/12/5 NCKU A-CN21
K. S. Wang 5079 China, Yunan Prov. Elv. 1000m. 2004/12/5 NCKU A-CN22
K. S. Wang 5084 China, Yunan Prov. Elv. 900m. 2004/12/5 NCKU A-CN23
K. S. Wang 5091 China, Yunan Prov. Elv. 900m. 2004/12/5 NCKU A-CN24
M. H. Su 670 Taiwan, Taipei Co. (Assam type) 2005/10/6 Authors A-IN1*
M. H. Su 667 Taiwan, Taipei Co. (Manipur type) 2005/10/6 Authors A-IN2*
M. H. Su 685 Taiwan, Nantou Co. (Sri Lanka type) 2005/11/15 Authors A-SR1*
M. H. Su 610 Taiwan, Nantou Co. Elv. 1000m. 2005/2/1 Authors A-NT1
M. H. Su 609 Taiwan, Nantou Co. Elv. 1000m. 2005/2/1 Authors A-NT2
M. H. Su 608 Taiwan, Nantou Co. Elv. 1000m. 2005/2/1 Authors A-NT3
M. T. Kao s. n. Taiwan, Nantou Co. 1955/2/12 TAI A-NT4
S. Hibino & S. Suzuki s. n. Taiwan, Nantou Co. 1926/7/17 TAI A-NT5
H. Keng, T. S. Liu & M. T.
Kao s. n. Taiwan, Nantou Co. 1955/7/20 TAI A-NT6
S. Y. Lu 18521 Taiwan, Nantou Co. Elv. 400m. 1986/3/2 TAIF A-NT7
S. Y. Lu 18291 Taiwan, Nantou Co. Elv. 700m. 1986/2/7 TAIF A-NT8
T. S. Liu & H. Keng 2853 Taiwan, Taitung Co. 1955/8/10 TAI A-TT1
Y.Y.-K.K. s. n. Taiwan, Taoyuan Co. 1937/3/2 TAI A-TY1
Y.Y.-K.K. s. n. Taiwan, Taoyuan Co. 1937/3/2 TAI A-TY2
Y.Y.-K.K. s. n. Taiwan, Taoyuan Co. 1937/3/2 TAI A-TY3
M. H. Su 684 Taiwan, Taipei Co. (Thailand type) 2005/11/15 Authors A-TH1*
var. sinensis C. H. Tsou 2117 China, Fujian Prov. Elv. 500-1200m. 2004/5/12 Authors S-CN1
C. H. Tsou et al. 1957 China, Guangdong Prov. 2004/8/6 Authors S-CN2
M. S. An 3357 China, Guizhou Prov. Elv. 920m. 2003/5/26 TAIF S-CN3
K. F. Wang 1-0560 China, Guizhou Prov. Elv. 650m. 2003/7/15 TAIF S-CN4
J. H. Hu 282 China, Hunan Prov. Elv. 500m. 2001/8/19 TAIF S-CN5
J. H. Hu 242 China, Hunan Prov. Elv. 880m. 2002/6/2 TAIF S-CN6
T. M. Taing 00748 China, Jiangxi Prov. 180m. 2000/10/22 TAIF S-CN7
Z. Chen 09769 China, Sichuan Prov. Elv. 810m 1996/10/5 TAIF S-CN8
T. Makino s. n. Japan, Tokyo City. 1910/10 TAIF S-JP1
M. H. Su 640 Taiwan, Chiayi Co. Elv. 1100m. 2005/9/12 Authors S-CY1
M. H. Su 644 Taiwan, Chiayi Co. Elv. 800m. 2005/9/13 Authors S-CY2
M. H. Su 95 Taiwan, Hsinchu Co. 2003/5/8 Authors S-HC1
Y. Shimada s. n. Taiwan, Hsinchu Co. 1913/11/15 TAIF S-HC2
A. T. Hsieh s. n. Taiwan, Miaoli Co. 1929/12/14 TAIF S-ML1
S. W. Chung 7520 Taiwan, Nantou Co. Elv. 700-800m. 2004/6/10 TAIF S-NT1
S. P. Chien s. n. Taiwan, Taipei City 1984/6/18 TAI S-TP1
C. S. Kuoh 2952 Taiwan, Taipei City 1971/11/23 TAI S-TP2
C. M. Kuo 5457 Taiwan, Taipei City 1974/7/6 TAI S-TP3
March, 2007 Su et al.: Morphological comparisons of Taiwan wild tea plant and related taxa 73
Table 1. Continued.
Name Collector with collection
number Site Date Herbarium Code
var. sinensis H. Simizu 418 Taiwan, Taipei City 1934/12/15 TAI S-TP4
Y. Yamamoto s. n. Taiwan, Taipei City 1930/11/29-30 TAI S-TP5
C. M. Kuo 9279 Taiwan, Taipei City 1978/1/2 TAI S-TP6
S. Suzuki s. n. Taiwan, Taipei City 1931/8/2 TAI S-TP7
G. Masamune 1534 Taiwan, Taipei City 1931/11/24 TAI S-TP8
S. Sasaki s. n. Taiwan, Taipei City 1922/10/15 TAI S-TP9
S. Suzuki s. n. Taiwan, Taipei City 1932/12/18 TAI S-TP10
S. Suzuki s. n. Taiwan, Taipei City 1929/11/30 TAI S-TP11
S. Sasaki s. n. Taiwan, Taipei City 1916/5 TAI S-TP12
S. Sasaki s. n. Taiwan, Taipei City 1925/12/9 TAI S-TP13
K. C. Yang 1162 Taiwan, Taipei City 1982/11/28 TAI S-TP14
C. C. Chou 41 Taiwan, Taipei City. 1984/6/18 TAI S-TP15
Y. Yamamoto s. n. Taiwan, Taipei City. 1938/2/27 TAI S-TP16
N. Y. Gu s. n. Taiwan, Taipei City. 1936/10/28 TAI S-TP17
B. L. Shie s. n. Taiwan, Taipei City. Elv. 850m. 1985/7/16 TAIF S-TP18
H. L. Chiang 423 Taiwan, Taipei City. 1997/6/15 TAIF S-TP19
S. Sasaki s. n. Taiwan, Taipei City. 1923/2 TAIF S-TP20
S. Sasaki s. n. Taiwan, Taipei City. 1927/10 TAIF S-TP21
K. C. Yang et al. 5265 Taiwan, Taipei City. 1996/12/31 TAIF S-TP22
S. Y. Lu 3335 Taiwan, Taipei City. Elv. 600m. 1975/11/19 TAIF S-TP23
M. H. Su 197 Taiwan, Taipei Co. Elv. 500m. 2003/5/13 Authors S-TP24
C. C. Hsu 5211 Taiwan, Taipei Co. 1968/12/27 TAI S-TP25
S. F. Huang K188 Taiwan, Taipei Co. 1987/7/31 TAI S-TP26
W. S. Tang s. n. Taiwan, Taipei Co. 1984/12/1 TAI S-TP27
S.Suzuki s. n. Taiwan, Taipei Co. 1924/11/2 TAI S-TP28
C. M. Kuo 6689A Taiwan, Taipei Co. 1978/8/29 TAI S-TP29
T. C. Huang 9756 Taiwan, Taipei Co. 1982/8/1 TAI S-TP30
S. Suzuki s. n. Taiwan, Taipei Co. 1923/10/21 TAI S-TP31
T. C. Huang, T. I. Yang & K.
C. Yang 1737 Taiwan, Taipei Co. 1985/10/18 TAI S-TP32
C. M. Kuo 5549 Taiwan, Taipei Co. 1974/7/28 TAI S-TP33
T. C. Huang & K. C. Yang
1924 Taiwan, Taipei Co. 1985/11/22 TAI S-TP34
J. H. Lii 236 Taiwan, Taipei Co. Elv. 200-400m. 2000/8/31 TAIF S-TP35
S. Y. Lu 12906 Taiwan, Taipei Co. 1983/9/15 TAIF S-TP36
H. M. H. Chang Taiwan, Taipei Co. Elv. 400m. 1999/10/12 TAIF S-TP37
H. L. Chiang 218 Taiwan, Taipei Co. Elv. 500m. 1996/10/6 TAIF S-TP38
W. F. Ho 314 Taiwan, Taipei Co. 1996/6/13 TAIF S-TP39
Y. H. Chang 4810 Taiwan, Taipei Co. Elv. 200-500m. 2001/10/11 TAIF S-TP40
S. C. Wu et al. s. n. Taiwan, Taipei Co. Elv. 350-500m. 1996/8/22 TAIF S-TP41
W. F. Ho Taiwan, Taipei Co. 1997/6/6 TAIF S-TP42
C. M. Chen s. n. Taiwan, Taipei Co. 2002/9/23 TAIF S-TP43
Y. H. Chang 4865 Taiwan, Taipei Co. 2001/10/18 TAIF S-TP44
W. F. Ho 126 Taiwan, Taipei Co. 1996/5/16 TAIF S-TP45
M. F. Loa & K. C. Yang 80 Taiwan, Taipei Co. Elv. 180-200m. 1996/10/5 TAIF S-TP46
Y.Y.-K.K. s. n. Taiwan, Taoyuan Co. 1937/3/2 TAI S-TY1
Y.Y.-K.K. s. n. Taiwan, Taoyuan Co. 1937/3/2 TAI S-TY2
Y.Y.-K.K. s. n. Taiwan, Taoyuan Co. 1937/3/2 TAI S-TY3
Y.Y.-K.K. s. n. Taiwan, Taoyuan Co. 1937/3/2 TAI S-TY4
Y. Shimada s. n. Taiwan, Taoyuan Co. 1913/11/20 TAIF S-TY5
Y. Shimada s. n. Taiwan, Taoyuan Co. 1913/11/20 TAIF S-TY6
Y. Shimada s. n. Taiwan, Taoyuan Co. 1913/11/20 TAIF S-TY7
Y. Shimada s. n. Taiwan, Taoyuan Co. 1913/11/20 TAIF S-TY8
Y. Shimada s. n. Taiwan, Taoyuan Co. 1913/11/20 TAIF S-TY9
Y. Shimada s. n. Taiwan, Taoyuan Co. 1913/11/15 TAIF S-TY10
Unknown (TAIF no. 31474) Unknown Unknown TAIF S-NN1
f. formosensis M. H. Su 642 Taiwan, Chiayi Co. Elv. 1300m. 2005/9/12 Authors F-CY1
C. I. Hu s. n. Taiwan, Chiayi Co. 2004/6 Authors F-CY2*
H. Yamada s. n. Taiwan, Chiayi Co. - TAIF F-CY3
Tang et al. 606 Taiwan, Kaohsiung Co. Elv. 1800m. 2005/1/2 Authors F-KH1
C. I. Hu s. n. Taiwan, Kaohsiung Co. 2004/6 Authors F-KH2*
C. C. Chuang & M. T. Kao
3369 Taiwan, Kaohsiung Co. Elv. 1350m. 1965/2/8 TAI F-KH3
74 TAIWANIA Vol. 52, No. 1
Table 1. Continued.
Name Collector with collection
number Site Date Herbarium Code
f. formosensis T. Kiang & M. T. Kao
KT439 Taiwan, Kaohsiung Co. 1971/5/12 TAI F-KH4
M. T. Kao 7448 Taiwan, Kaohsiung Co. 1968/12/10 TAI F-KH5
T. C. Huang 4952 Taiwan, Kaohsiung Co. 1968/12/10 TAI F-KH6
A. Tanimura s. n. Taiwan, Kaohsiung Co. 1935/1/12 TAI F-KH7
S. Sasaki s. n. Taiwan, Kaohsiung Co. 1936/3/8 TAI F-NN8
S. Y. Lu 18256 Taiwan, Kaohsiung Co. Elv. 1150m. 1986/1/30 TAIF F-KH9
S. Y. Lu 18664 Taiwan, Kaohsiung Co. Elv. 1500m. 1986/3/11 TAIF F-KH10
Y. H. Lai 83 Taiwan, Kaohsiung Co. Elv.
750-800m. 1996/12/2 TAIF F-KH11
C. P. Lin s. n. Taiwan, Kaohsiung Co. Elv. 650m. 2004/5/25 TAIF F-KH12
C. H. Tsou 2134 Taiwan, Nantou Co. Elv. 1200m. 2005/3/30 Authors F-NT1
C. H. Tsou 2132 Taiwan, Nantou Co. Elv. 1200m. 2005/3/30 Authors F-NT2
C. H. Tsou 2137 Taiwan, Nantou Co. Elv. 1200m. 2005/3/30 Authors F-NT3
C. H. Tsou 2139 Taiwan, Nantou Co. Elv. 1200m. 2005/3/30 Authors F-NT4
M. H. Su 687 Taiwan, Nantou Co. 2005/11/15 Authors F-NT5
M. H. Su 683 Taiwan, Nantou Co. 2005/11/15 Authors F-NT6
M. Hasimoto s. n. Taiwan, Nantou Co. 1966/1/13 TAI F-NT7
S. Sasaki s. n. Taiwan, Nantou Co. 1935/11/8 TAI F-NT8
S. Taniguchi s. n. Taiwan, Nantou Co. 1931/7/12 TAI F-NT9
S. Suzuki 3245 Taiwan, Nantou Co. 1935/11/8 TAI F-NT10
S. Sasaki s. n. Taiwan, Nantou Co. 1935/11/8 TAI F-NT11
M. Hasimoto s. n. Taiwan, Nantou Co. 1966/1/16 TAI F-NT12
S. Sasaki s. n. Taiwan, Nantou Co. 1935/10/8 TAI F-NT13
M. T. Kao 6668 Taiwan, Nantou Co. 1966/4/23 TAI F-NT14
B. J. Wang 15069 Taiwan, Nantou Co. 1988/12/25 TAIF F-NT15
S. Sasaki s. n. Taiwan, Nantou Co. 1922/11/30 TAIF F-NT16
M. H. Su 575 Taiwan, Pingtung Co. Elv. 1400m. 2004/4/14 Authors F-PT1
M. H. Su 269 Taiwan, Pingtung Co. 2003/9/20 Authors F-PT2
M. H. Su 270 Taiwan, Pingtung Co. Elv. 1300m. 2003/9/20 Authors F-PT3
M. H. Su 497 Taiwan, Pingtung Co. Elv. 1200m. 2004/1/24 Authors F-PT4
M. H. Su 498 Taiwan, Pingtung Co. Elv. 1200m. 2004/1/24 Authors F-PT5
M. H. Su 646 Taiwan, Pingtung Co. Elv. 1100m. 2005/9/27 Authors F-PT6
M. H. Su 645 Taiwan, Pingtung Co. Elv. 1100m. 2005/9/27 Authors F-PT7
M. H. Su 544 Taiwan, Pingtung Co. Elv. 1100m. 2004/3/10 Authors F-PT8
M. H. Su 545 Taiwan, Pingtung Co. Elv. 1100m. 2004/3/10 Authors F-PT9
E. Matuda s. n. Taiwan, Pingtung Co. 1919/7/11 TAI F-PT10
E. Matuda Taiwan, Pingtung Co. 1912/11/7 TAI F-PT11
K. C. Yang et al. 4583 Taiwan, Pingtung Co. Elv. 850m. 1995/12/2 TAIF F-PT12
K. C. Yang et al. 4527 Taiwan, Pingtung Co. Elv.
750-1100m. 1995/12/3 TAIF F-PT13
S. W. Chung 7090 Taiwan, Pingtung Co. Elv.
800-1000m. 2004/5/29 TAIF F-PT14
M. H. Su 659 Taiwan, Taitung Co. Elv. 1000m. 2005/9/28 Authors F-TT1
M. H. Su 661 Taiwan, Taitung Co. Elv. 1100m. 2005/9/28 Authors F-TT2
M. H. Su 660 Taiwan, Taitung Co. Elv. 1100m. 2005/9/28 Authors F-TT3
M. H. Su 655 Taiwan, Taitung Co. Elv. 1100m. 2005/9/28 Authors F-TT4
M. T. Kao 6612 Taiwan, Taoyuan Co. 1966/1/11 TAI F-TY1
K. Mori 1901 Taiwan, Yunlin Co. 1906/11/5 TAIF F-YL1
Tanimura s. n. Unknown 1935/1/12 TAI F-NN1
chosen with respect to variation among taxa
mentioned in literature and based on personal
observations on specimens. Finally, a total of 35
characters were scored, including 17 vegetative and
18 floral characters (Table 2). For each specimen,
five mature, healthy-look leaves were scored and
averaged. The measurement on floral characters was
averaged from one to three flowers, depending on the
condition of the specimens. We also measured the
angles between the midrib and one major lateral vein
at two different positions (Fig. 2), because the usual
curved lateral veins cannot be expressed by a single
value (often measured at the base in most studies).
Upon further examination, it was found that one
vegetative and seven floral characters were constant
(not informative) and should be eliminated from the
following analyses. Finally, 27 (16 vegetative and 11
floral) characters were selected. Since not all
specimens were in the flowering stage, the data for
the numerical analyses were divided into three
March, 2007 Su et al.: Morphological comparisons of Taiwan wild tea plant and related taxa 75
Table 2. List of morphological characters examined for
multivariate analyses. The asterisk (*) denotes constant characters
that are excluded from the data analysis. The angles measured
between midrib and lateral vein are shown in Fig. 2.
Character Data type
Vegetative Characters
Bud pubescence Multi-state
Young branchlet pubescence Binary
Leaf length Quantitative
Leaf width Quantitative
Leaf thickness (dry specimen) Quantitative
Leaf shape Multi-state
Leaf apex shape Multi-state
Leaf base shape Multi-state
Leaf serration density (count / 4 cm) Quantitative
Leaf pubescence on adaxial surface* Binary
Leaf pubescence on abaxial surface Binary
Abaxial midrib pubescence Binary
Pairs of lateral veins Quantitative
Lateral vein angle at base (Fig. 2a) Quantitative
Lateral vein angle at middle (Fig. 2b) Quantitative
Petiole length Quantitative
Petiole pubescence Binary
Floral characters
Pedicel length Quantitative
Pedicel pubescence* Binary
Petal number per flower Quantitative
Petal length Quantitative
Petal width Quantitative
Petal pubescence* Multi-state
Sepal number per flower* Quantitative
Sepal length Quantitative
Sepal width Quantitative
Sepal pubescence Multi-state
Filament length Quantitative
Filament pubescence* Binary
Style length Quantitative
Style number per flower* Quantitative
Style pubescence* Binary
Stigma number* Quantitative
Ovary pubescence Binary
Flower number per cluster Quantitative
categories: (1) only vegetative characters (165 OTUs
× 16 characters); (2) only floral characters (62 OTUs ×
11 characters); and (3) all characters (62 OTUs × 27
characters).
Cluster analysis
Similarity matrices were generated using the
coefficient proposed by Gower (1971). Gower's
similarity coefficient (GSC) was designed to deal with
mixed type of characters, and was thus widely used
(Schultze-Motel and Meyer, 1981; Zaharof, 1988;
Cheng, 1990; Ward, 1993; Gugerli, 1997; St-Laurent
et al., 2000; Muvaffak et al., 2001; Binns et al., 2002;
Bayly et al., 2003; Mckenzie et al., 2004). This
similarity matrix was then used to perform a cluster
analysis using the Unweighted Pair Grouping Method
Based on Arithmetic Averages (UPGMA) (Sokal and
Michener, 1958) with the software MVSP v3.01
(Kovach Computing Service, 1999).
ab
ab
Fig. 2. Scheme of a leaf showing
the angles (a) formed between
the midrib and a lateral vein, and
(b) created by the midrib and the
interception of the tangent to the
middle portion of a lateral vein.
Nonlinear principal component analysis
To further explore the pattern of variation in
measured characters and to find those characters
which are decisive to distinguish taxa, a nonlinear
principal components analysis (NLPCA, de Leeuw,
1982) was undertaken. Similar to principal
components analysis, NLPCA can be used for
transforming attributes of a dataset into a new set of
uncorrelated attributes (principal components), while
still retaining as much of the variability of the dataset
as possible. It can handle variables of different types
(nominal, ordinal and numerical) simultaneously, and
deal with nonlinear relationships between variables.
NLPCA is performed by the program CATPCA
implemented in the software SPSS v13.0 (SPSS Inc.).
In addition, Cronbach's Alpha (Cronbach, 1951) was
calculated for each of the components extracted. If
Alpha value of a specific component is high, it would
be interpreted as indicating that the component has a
strong one-dimensional structure, or, the dimension
is reliable to account for the total variance. Generally,
an Alpha value of 0.70 or greater is considered to be
reliable (Bland and Altman, 1997).
RESULTS
Vegetative characters
The UPGMA phenogram based on vegetative
characters showed two discrete clusters (Fig. 3),
namely Group I-1 (GSC = 0.62) and Group I-2 (GSC
= 0.67). Group I-1 was composed entirely of C.
sinensis f. formosensis from central and southern
Taiwan. Within this cluster, there did not appear to be
any regional patterns. Group I-2, however, contained
all the samples of C. sinensis var. sinensis and C.
sinensis var. assamica, with C. sinensis f.
formosensis from eastern Taiwan. Despite of that,
samples of eastern C. sinensis f. formosensis formed
a consistent subgroup (Group I-2-1; GSC = 0.87)
within Group I-2. In contrast, all samples of C.
sinensis var. sinensis and C. sinensis var. assamica
overlapped extensively and together they formed a
large subgroup (Group I-2-2; GSC = 0.71).
76 TAIWANIA Vol. 52, No. 1
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Fig. 3. Phenogram of UPGMA cluster analysis based on vegetative
characters. The coefficient was defined as the Gower's similarity
coefficient.
The results of NLPCA on the vegetative
characters are presented in Table 3. The first three
components accounted for 65.3 % of the total
variance in the dataset. The first component alone
accounted for 43.2% of the total variance and was far
more important than other components. The first
component with a Cronbach's alpha value of 0.91 was
the only one considered to be reliable. The
component loadings correspond to the correlation
coefficients between characters and the derived
components. Characters with high loadings were bud
pubescence, young branchlet pubescence, abaxial
midrib pubescence and petiole pubescence (over 0.9),
followed by leaf length, leaf apex shape and pairs of
lateral veins (over 0.7). The plot by the first two
components (Fig. 4) shows a similar grouping with
the cluster analysis. However, the eastern samples of
C. sinensis f. formosensis were placed onto the
intermediate positions in NLPCA. It didn't group
these eastern samples with C. sinensis var. sinensis
and C. sinensis var. assamica absolutely.
Table 3. Loadings of the 16 vegetative characters on the first three
components from NLPCA. Eigenvalues, percentage of variance
explained and cumulated, and Cronbach's Alpha are given for each
component.
Component
Character 1 2 3
Bud pubescence -0.93 0.29 -0.05
Young branchlet pubescence 0.91 -0.32 0.03
Leaf length -0.77 -0.55 -0.09
Leaf width -0.55 -0.71 0.17
Leaf thickness 0.13 -0.10 0.25
Leaf shape -0.33 0.10 0.47
Leaf apex shape -0.74 -0.02 -0.26
Leaf base shape 0.14 -0.02 0.60
Leaf serration density 0.34 0.73 0.03
Leaf pubescence on abaxial surface 0.67 -0.23 0.36
Abaxial midrib pubescence 0.90 -0.32 0.06
Pairs of lateral veins -0.79 -0.27 0.13
Lateral vein angle at base -0.39 0.16 0.64
Lateral vein angle at middle -0.64 0.13 0.41
Petiole length -0.50 -0.35 -0.18
Petiole pubescence 0.91 -0.30 0.04
Eigenvalue 6.90 2.03 1.51
Variance explained (%) 43.20 12.70 9.40
Variance cumulative (%) 43.20 55.90 65.30
Cronbach's Alpha 0.91 0.54 0.36
210-1-2
Component 1
3
2
1
0
-1
-2
-3
Component 2
210-1-2
Component 1
3
2
1
0
-1
-2
-3
Component 2
Fig. 4. Ordination plot of NLPCA based on vegetative characters.
: central and southern C. sinensis f. formosensis. : eastern C.
sinensis f. formosensis. {: C. sinensis var. assamica. c: C. sinensis
var. sinensis.
March, 2007 Su et al.: Morphological comparisons of Taiwan wild tea plant and related taxa 77
Floral characters
Cluster analysis of OTUs based on floral
characters revealed two main groups (i.e. Group II-1
and Group II-2; Fig. 5), similar to the result obtained
based on vegetative characters (Fig. 3). However, the
members within each cluster were different. The
OTUs of C. sinensis f. formosensis from eastern
Taiwan, previously clustered within Group I-2, are
now located amongst Group II-1. Consequently,
Group II-1 encompassed all samples of C. sinensis f.
formosensis, while Group II-2 consisted of a mix of
C. sinensis var. sinensis and C. sinensis var.
assamica. A further subdivision of the two main
groups into geographical or taxonomic subgroups
could not be made.
The results of NLPCA on floral characters are
shown in Table 4. The first three principal
components accounted for 39.1%, 13.2% and 10.8%
of the total variance, respectively. Only the first
principal component was meaningful (Cronbach's
alpha = 0.84) for grouping samples discretely.
Characters with high loadings on the first principal
component were petal length, petal width and ovary
pubescence (over 0.8), followed by sepal width and
style length (over 0.7). C. sinensis f. formosensis also
can be separated by a line on the plot of the first two
components (Fig. 6), a result just the same with that
of the cluster analysis.
II-1
II-2
A-BU1
S-TP21
A-NT1
S-CY1
S-TP4
S-TP22
S-TP38
S-TP37
A-NT8
S-TP2
S-TP10
S-TP31
S-TP44
A-CN2
S-TP13
A-CN17
S-TP5
S-NN1
S-ML1
A-CN8
A-CN12
A-CN19
A-CN11
A-CN3
A-CN18
S-TP8
S-TY8
A-CN9
A-CN21
A-CN22
A-CN6
A-CN23
S-TP11
A-CN14
S-TP27
S-TP20
A-CN10
S-TP25
A-CN15
S-TP17
S-TP32
S-CN8
A-CN4
A-CN24
S-TY2
A-CN7
A-CN13
A-IN1
F-CY1
F-NT15
F-PT6
F-PT7
F-TT4
F-PT13
F-KH11
F-PT2
F-KH1
F-PT12
F-TT3
F-CY2
F-NT6
F-NT11
0.6 0.68 0.76 0.84 0.92 1
A-BU1
S-TP21
A-NT1
S-CY1
S-TP4
S-TP22
S-TP38
S-TP37
A-NT8
S-TP2
S-TP10
S-TP31
S-TP44
A-CN2
S-TP13
A-CN17
S-TP5
S-NN1
S-ML1
A-CN8
A-CN12
A-CN19
A-CN11
A-CN3
A-CN18
S-TP8
S-TY8
A-CN9
A-CN21
A-CN22
A-CN6
A-CN23
S-TP11
A-CN14
S-TP27
S-TP20
A-CN10
S-TP25
A-CN15
S-TP17
S-TP32
S-CN8
A-CN4
A-CN24
S-TY2
A-CN7
A-CN13
A-BU1
S-TP21
A-NT1
S-CY1
S-TP4
S-TP22
S-TP38
S-TP37
A-NT8
S-TP2
S-TP10
S-TP31
S-TP44
A-CN2
S-TP13
A-CN17
S-TP5
S-NN1
S-ML1
A-CN8
A-CN12
A-CN19
A-CN11
A-CN3
A-CN18
S-TP8
S-TY8
A-CN9
A-CN21
A-CN22
A-CN6
A-CN23
S-TP11
A-CN14
S-TP27
S-TP20
A-CN10
S-TP25
A-CN15
S-TP17
S-TP32
S-CN8
A-CN4
A-CN24
S-TY2
A-CN7
A-CN13
A-IN1
F-CY1
F-NT15
F-PT6
F-PT7
F-TT4
F-PT13
F-KH11
F-PT2
F-KH1
F-PT12
F-TT3
F-CY2
F-NT6
F-NT11
0.6 0.68 0.76 0.84 0.92 1
II-1
II-2
II-1II-1
II-2II-2
A-BU1
S-TP21
A-NT1
S-CY1
S-TP4
S-TP22
S-TP38
S-TP37
A-NT8
S-TP2
S-TP10
S-TP31
S-TP44
A-CN2
S-TP13
A-CN17
S-TP5
S-NN1
S-ML1
A-CN8
A-CN12
A-CN19
A-CN11
A-CN3
A-CN18
S-TP8
S-TY8
A-CN9
A-CN21
A-CN22
A-CN6
A-CN23
S-TP11
A-CN14
S-TP27
S-TP20
A-CN10
S-TP25
A-CN15
S-TP17
S-TP32
S-CN8
A-CN4
A-CN24
S-TY2
A-CN7
A-CN13
A-IN1
F-CY1
F-NT15
F-PT6
F-PT7
F-TT4
F-PT13
F-KH11
F-PT2
F-KH1
F-PT12
F-TT3
F-CY2
F-NT6
F-NT11
0.6 0.68 0.76 0.84 0.92 1
A-BU1
S-TP21
A-NT1
S-CY1
S-TP4
S-TP22
S-TP38
S-TP37
A-NT8
S-TP2
S-TP10
S-TP31
S-TP44
A-CN2
S-TP13
A-CN17
S-TP5
S-NN1
S-ML1
A-CN8
A-CN12
A-CN19
A-CN11
A-CN3
A-CN18
S-TP8
S-TY8
A-CN9
A-CN21
A-CN22
A-CN6
A-CN23
S-TP11
A-CN14
S-TP27
S-TP20
A-CN10
S-TP25
A-CN15
S-TP17
S-TP32
S-CN8
A-CN4
A-CN24
S-TY2
A-CN7
A-CN13
A-BU1
S-TP21
A-NT1
S-CY1
S-TP4
S-TP22
S-TP38
S-TP37
A-NT8
S-TP2
S-TP10
S-TP31
S-TP44
A-CN2
S-TP13
A-CN17
S-TP5
S-NN1
S-ML1
A-CN8
A-CN12
A-CN19
A-CN11
A-CN3
A-CN18
S-TP8
S-TY8
A-CN9
A-CN21
A-CN22
A-CN6
A-CN23
S-TP11
A-CN14
S-TP27
S-TP20
A-CN10
S-TP25
A-CN15
S-TP17
S-TP32
S-CN8
A-CN4
A-CN24
S-TY2
A-CN7
A-CN13
A-IN1
F-CY1
F-NT15
F-PT6
F-PT7
F-TT4
F-PT13
F-KH11
F-PT2
F-KH1
F-PT12
F-TT3
F-CY2
F-NT6
F-NT11
0.6 0.68 0.76 0.84 0.92 1
A-BU1
S-TP21
A-NT1
S-CY1
S-TP4
S-TP22
S-TP38
S-TP37
A-NT8
S-TP2
S-TP10
S-TP31
S-TP44
A-CN2
S-TP13
A-CN17
S-TP5
S-NN1
S-ML1
A-CN8
A-CN12
A-CN19
A-CN11
A-CN3
A-CN18
S-TP8
S-TY8
A-CN9
A-CN21
A-CN22
A-CN6
A-BU1
S-TP21
A-NT1
S-CY1
S-TP4
S-TP22
S-TP38
S-TP37
A-NT8
S-TP2
S-TP10
S-TP31
S-TP44
A-CN2
S-TP13
A-CN17
S-TP5
S-NN1
S-ML1
A-CN8
A-CN12
A-CN19
A-CN11
A-CN3
A-CN18
S-TP8
S-TY8
A-CN9
A-CN21
A-CN22
A-CN6
A-CN23
S-TP11
A-CN14
S-TP27
S-TP20
A-CN10
S-TP25
A-CN15
S-TP17
S-TP32
S-CN8
A-CN4
A-CN24
S-TY2
A-CN7
A-CN13
A-IN1
F-CY1
F-NT15
F-PT6
F-PT7
F-TT4
F-PT13
F-KH11
F-PT2
F-KH1
F-PT12
F-TT3
F-CY2
F-NT6
F-NT11
0.6 0.68 0.76 0.84
A-CN23
S-TP11
A-CN14
S-TP27
S-TP20
A-CN10
S-TP25
A-CN15
S-TP17
S-TP32
S-CN8
A-CN4
A-CN24
S-TY2
A-CN7
A-CN13
A-IN1
F-CY1
F-NT15
F-PT6
F-PT7
F-TT4
F-PT13
F-KH11
F-PT2
F-KH1
F-PT12
F-TT3
F-CY2
F-NT6
F-NT11
0.6 0.68 0.76 0.84 0.92 1
A-BU1
S-TP21
A-NT1
S-CY1
S-TP4
S-TP22
S-TP38
S-TP37
A-NT8
S-TP2
S-TP10
S-TP31
S-TP44
A-CN2
S-TP13
A-CN17
S-TP5
S-NN1
S-ML1
A-CN8
A-CN12
A-CN19
A-CN11
A-CN3
A-CN18
S-TP8
S-TY8
A-CN9
A-CN21
A-CN22
A
0.92 1
A-BU1
S-TP21
A-NT1
S-CY1
S-TP4
S-TP22
S-TP38
S-TP37
A-NT8
S-TP2
S-TP10
S-TP31
S-TP44
A-CN2
S-TP13
A-CN17
S-TP5
S-NN1
S-ML1
A-CN8
A-CN12
A-CN19
A-CN11
A-CN3
A-CN18
S-TP8
S-TY8
A-CN9
A-CN21
A-CN22
A-CN6
A-CN23
S-TP11
A-CN14
S-TP27
S-TP20
A-CN10
S-TP25
A-CN15
S-TP17
S-TP32
S-CN8
A-CN4
A-CN24
S-TY2
A-CN7
A-CN13
A-BU1
S-TP21
A-NT1
S-CY1
S-TP4
S-TP22
S-TP38
S-TP37
A-NT8
S-TP2
S-TP10
S-TP31
S-TP44
A-CN2
S
-CN6
A-CN23
S-TP11
A-CN14
S-TP27
S-TP20
A-CN10
S-TP25
A-CN15
S-TP17
S-TP32
S-CN8
A-CN4
A-CN24
S-TY2
A-CN7
A-CN13
A-BU1
S-TP21
A-NT1
S-CY1
S-TP4
S-TP22
S-TP38
S-TP37
A-NT8
S-TP2
S-TP10
S-TP31
S-TP44
A-CN2
S-TP13
A-CN17
S-TP5
S-NN1
S-ML1
A-CN8
A-CN12
A-CN19
A-CN11
A-CN3
A-CN18
S-TP8
S-TY8
A-CN9
A-CN21
A-CN22
A-CN6
A-CN23
S-TP11
A-CN14
S-TP27
S-TP20
A-CN10
S-TP25
A-CN15
S-TP17
S-TP32
S-CN8
A-CN4
A-CN24
S-TY2
A-CN7
A-CN13
A-IN1
-TP13
A-CN17
S-TP5
S-NN1
S-ML1
A-CN8
A-CN12
A-CN19
A-CN11
A-CN3
A-CN18
S-TP8
S-TY8
A-CN9
A-CN21
A-CN22
A-CN6
A-CN23
S-TP11
A-CN14
S-TP27
S-TP20
A-CN10
S-TP25
A-CN15
S-TP17
S-TP32
S-CN8
A-CN4
A-CN24
S-TY2
A-CN7
A-CN13
A-IN1
F-CY1
F-NT15
F-PT6
F-PT7
F-TT4
F-PT13
F-KH11
F-PT2
F-KH1
F-PT12
F-TT3
F-CY2
F-NT6
F-NT11
0.6 0.68 0.76 0.84 0.92 1
Fig. 5. Phenogram of UPGMA cluster analysis based on floral
characters. The coefficient was defined as the Gower's similarity
coefficient.
Table 4. Loadings of the 11 floral characters on the first three
components from NLPCA. Eigenvalues, percentage of variance
explained and cumulated, and Cronbach's Alpha are given for each
component.
Component
Character 1 2 3
Pedicel length 0.66 0.23 -0.34
Petal number per flower 0.39 0.15 0.61
Petal length 0.85 -0.16 0.02
Petal width 0.80 -0.16 0.19
Sepal length 0.58 0.31 -0.09
Sepal width 0.71 0.50 -0.05
Sepal pubescence 0.14 -0.54 0.65
Filament length 0.47 -0.42 -0.23
Style length 0.78 -0.19 -0.14
Ovary pubescence 0.80 0.05 0.19
Flower number per cluster -0.15 0.70 0.39
Eigenvalue 4.30 1.46 1.19
Variance explained (%) 39.10 13.20 10.80
Variance cumulative (%) 39.10 52.30 63.10
Cronbach's Alpha 0.84 0.35 0.18
210-1-2-3
Component 1
4
2
0
-2
-4
Component 2
210-1-2-3
Component 1
4
2
0
-2
-4
Component 2
210-1-2-3
Component 1
4
2
0
-2
-4
Component 2
Fig. 6. Ordination plot of NLPCA based on floral characters.
: C. sinensis f. formosensis. {: C. sinensis var. assamica. c: C.
sinensis var. sinensis.
All characters
Cluster analysis of all the floral and vegetative
characters produced a phenogram with two groups of
OTUs (Group III-1 and Group III-2, Fig. 7) that
corresponded to the separation based on floral
characters. OTUs of C. sinensis f. formosensis were
78 TAIWANIA Vol. 52, No. 1
III-1
III-2
A-BU1
S-TP21
A-NT1
S-CY1
S-TP2
A-NT8
S-TP4
S-TP38
S-TP22
S-TP37
S-TP44
S-TP31
A-CN2
A-CN11
A-CN24
A-CN10
A-CN23
A-CN13
A-CN19
A-CN8
S-TP17
A-CN9
A-CN22
S-CN8
S-TP5
S-NN1
A-CN6
S-ML1
S-TP13
S-TP27
S-TP20
A-CN12
A-CN17
A-CN14
A-CN21
A-CN15
S-TP25
S-TP8
S-TY8
A-CN4
S-TY2
S-TP32
A-CN7
A-IN1
S-TP10
S-TP11
A-CN3
A-CN18
F-CY1
F-CY2
F-NT6
F-KH1
F-PT2
F-KH11
F-NT11
F-NT15
F-PT7
F-PT6
F-PT13
F-TT3
F-TT4
F-PT12
0.520.6 0.680.760.840.92 1
A-BU1
S-TP21
A-NT1
S-CY1
S-TP2
A-NT8
S-TP4
S-TP38
S-TP22
S-TP37
S-TP44
S-TP31
A-CN2
A-CN11
A-CN24
A-CN10
A-CN23
A-CN13
A-CN19
A-CN8
S-TP17
A-CN9
A-CN22
S-CN8
S-TP5
S-NN1
A-CN6
S-ML1
S-TP13
S-TP27
S-TP20
A-CN12
A-CN17
A-CN14
A-CN21
A-CN15
S-TP25
S-TP8
S-TY8
A-CN4
S-TY2
S-TP32
A-CN7
A-IN1
S-TP10
S-TP11
A-CN3
A-BU1
S-TP21
A-NT1
S-CY1
S-TP2
A-NT8
S-TP4
S-TP38
S-TP22
S-TP37
S-TP44
S-TP31
A-CN2
A-CN11
A-CN24
A-CN10
A-CN23
A-CN13
A-CN19
A-CN8
S-TP17
A-CN9
A-CN22
S-CN8
S-TP5
S-NN1
A-CN6
S-ML1
S-TP13
S-TP27
S-TP20
A-CN12
A-CN17
A-CN14
A-CN21
A-CN15
S-TP25
S-TP8
S-TY8
A-CN4
S-TY2
S-TP32
A-CN7
A-IN1
S-TP10
S-TP11
A-CN3
A-CN18
F-CY1
F-CY2
F-NT6
F-KH1
F-PT2
F-KH11
F-NT11
F-NT15
F-PT7
F-PT6
F-PT13
F-TT3
F-TT4
F-PT12
0.520.6 0.680.760.840.92 1
III-1III-1
III-2
III-2
A-BU1
S-TP21
A-NT1
S-CY1
S-TP2
A-NT8
S-TP4
S-TP38
S-TP22
S-TP37
S-TP44
S-TP31
A-CN2
A-CN11
A-CN24
A-CN10
A-CN23
A-CN13
A-CN19
A-CN8
S-TP17
A-CN9
A-CN22
S-CN8
S-TP5
S-NN1
A-CN6
S-ML1
S-TP13
S-TP27
S-TP20
A-CN12
A-CN17
A-CN14
A-CN21
A-CN15
S-TP25
S-TP8
S-TY8
A-CN4
S-TY2
S-TP32
A-CN7
A-IN1
S-TP10
S-TP11
A-CN3
A-CN18
F-CY1
F-CY2
F-NT6
F-KH1
F-PT2
F-KH11
F-NT11
F-NT15
F-PT7
F-PT6
F-PT13
F-TT3
F-TT4
F-PT12
0.520.6 0.680.760.840.92 1
A-BU1
S-TP21
A-NT1
S-CY1
S-TP2
A-NT8
S-TP4
S-TP38
S-TP22
S-TP37
S-TP44
S-TP31
A-CN2
A-CN11
A-CN24
A-CN10
A-CN23
A-CN13
A-CN19
A-CN8
S-TP17
A-CN9
A-CN22
S-CN8
S-TP5
S-NN1
A-CN6
S-ML1
S-TP13
S-TP27
S-TP20
A-CN12
A-CN17
A-CN14
A-CN21
A-CN15
S-TP25
S-TP8
S-TY8
A-CN4
S-TY2
S-TP32
A-CN7
A-IN1
S-TP10
S-TP11
A-CN3
A-BU1
S-TP21
A-NT1
S-CY1
S-TP2
A-NT8
S-TP4
S-TP38
S-TP22
S-TP37
S-TP44
S-TP31
A-CN2
A-CN11
A-CN24
A-CN10
A-CN23
A-CN13
A-CN19
A-CN8
S-TP17
A-CN9
A-CN22
S-CN8
S-TP5
S-NN1
A-CN6
S-ML1
S-TP13
S-TP27
S-TP20
A-CN12
A-CN17
A-CN14
A-CN21
A-CN15
S-TP25
S-TP8
S-TY8
A-CN4
S-TY2
S-TP32
A-CN7
A-IN1
S-TP10
S-TP11
A-CN3
A-CN18
F-CY1
F-CY2
F-NT6
F-KH1
F-PT2
F-KH11
F-NT11
F-NT15
F-PT7
F-PT6
F-PT13
F-TT3
F-TT4
F-PT12
0.520.6 0.680.760.840.92 1
A-BU1
S-TP21
A-NT1
S-CY1
S-TP2
A-NT8
S-TP4
S-TP38
S-TP22
S-TP37
S-TP44
S-TP31
A-CN2
A-CN11
A-CN24
A-CN10
A-CN23
A-CN13
A-CN19
A-CN8
S-TP17
A-CN9
A-CN22
S-CN8
S-TP5
S-NN1
A-CN6
S-ML1
S-TP13
S-TP27
S-TP20
A-BU1
S-TP21
A-NT1
S-CY1
S-TP2
A-NT8
S-TP4
S-TP38
S-TP22
S-TP37
S-TP44
S-TP31
A-CN2
A-CN11
A-CN24
A-CN10
A-CN23
A-CN13
A-CN19
A-CN8
S-TP17
A-CN9
A-CN22
S-CN8
S-TP5
S-NN1
A-CN6
S-ML1
S-TP13
S-TP27
S-TP20
A-CN12
A-CN17
A-CN14
A-CN21
A-CN15
S-TP25
S-TP8
S-TY8
A-CN4
S-TY2
S-TP32
A-CN7
A-IN1
S-TP10
S-TP11
A-CN3
A-CN18
F-CY1
F-CY2
F-NT6
F-KH1
F-PT2
F-KH11
F-NT11
F-NT15
F-PT7
F-PT6
F-PT13
F-TT3
F-TT4
F-PT12
0.52 0.6 0.68 0.76 0.84
A-CN12
A-CN17
A-CN14
A-CN21
A-CN15
S-TP25
S-TP8
S-TY8
A-CN4
S-TY2
S-TP32
A-CN7
A-IN1
S-TP10
S-TP11
A-CN3
A-CN18
F-CY1
F-CY2
F-NT6
F-KH1
F-PT2
F-KH11
F-NT11
F-NT15
F-PT7
F-PT6
F-PT13
F-TT3
F-TT4
F-PT12
0.520.6 0.680.760.840.92 1
A-BU1
S-TP21
A-NT1
S-CY1
S-TP2
A-NT8
S-TP4
S-TP38
S-TP22
S-TP37
S-TP44
S-TP31
A-CN2
A-CN11
A-CN24
A-CN10
A-CN23
A-CN13
A-CN19
A-CN8
S-TP17
A-CN9
A-CN22
S-CN8
S-TP5
S-NN1
A-CN6
S-ML1
S-TP13
S-TP27
S
0.92 1
A-BU1
S-TP21
A-NT1
S-CY1
S-TP2
A-NT8
S-TP4
S-TP38
S-TP22
S-TP37
S-TP44
S-TP31
A-CN2
A-CN11
A-CN24
A-CN10
A-CN23
A-CN13
A-CN19
A-CN8
S-TP17
A-CN9
A-CN22
S-CN8
S-TP5
S-NN1
A-CN6
S-ML1
S-TP13
S-TP27
S-TP20
A-CN12
A-CN17
A-CN14
A-CN21
A-CN15
S-TP25
S-TP8
S-TY8
A-CN4
S-TY2
S-TP32
A-CN7
A-IN1
S-TP10
S-TP11
A-CN3
A-BU1
S-TP21
A-NT1
S-CY1
S-TP2
A-NT8
S-TP4
S-TP38
S-TP22
S-TP37
S-TP44
S-TP31
A-CN2
A-CN11
A
-TP20
A-CN12
A-CN17
A-CN14
A-CN21
A-CN15
S-TP25
S-TP8
S-TY8
A-CN4
S-TY2
S-TP32
A-CN7
A-IN1
S-TP10
S-TP11
A-CN3
A-BU1
S-TP21
A-NT1
S-CY1
S-TP2
A-NT8
S-TP4
S-TP38
S-TP22
S-TP37
S-TP44
S-TP31
A-CN2
A-CN11
A-CN24
A-CN10
A-CN23
A-CN13
A-CN19
A-CN8
S-TP17
A-CN9
A-CN22
S-CN8
S-TP5
S-NN1
A-CN6
S-ML1
S-TP13
S-TP27
S-TP20
A-CN12
A-CN17
A-CN14
A-CN21
A-CN15
S-TP25
S-TP8
S-TY8
A-CN4
S-TY2
S-TP32
A-CN7
A-IN1
S-TP10
S-TP11
A-CN3
A-CN18
-CN24
A-CN10
A-CN23
A-CN13
A-CN19
A-CN8
S-TP17
A-CN9
A-CN22
S-CN8
S-TP5
S-NN1
A-CN6
S-ML1
S-TP13
S-TP27
S-TP20
A-CN12
A-CN17
A-CN14
A-CN21
A-CN15
S-TP25
S-TP8
S-TY8
A-CN4
S-TY2
S-TP32
A-CN7
A-IN1
S-TP10
S-TP11
A-CN3
A-CN18
F-CY1
F-CY2
F-NT6
F-KH1
F-PT2
F-KH11
F-NT11
F-NT15
F-PT7
F-PT6
F-PT13
F-TT3
F-TT4
F-PT12
0.520.6 0.680.760.840.92 1
Fig. 7. Phenogram of UPGMA cluster analysis based on vegetative
and floral characters. The coefficient was defined as the Gower's
similarity coefficient.
contained entirely within Group III-1, while C.
sinensis var. sinensis and C. sinensis var. assamica
were dispersed throughout Group III-2.
The NLPCA character loadings, percentage, and
variance explained and cumulated for the first three
components are given in Table 5. The first component
accounted for 36.8% of the total variance observed,
and was highly interpretable (Cronbach's alpha =
0.93). It had high contributing component loadings
from bud pubescence, young branchlet pubescence,
abaxial midrib pubescence, petiole pubescence and
ovary pubescence (over 0.88), and leaf length, leaf
apex shape, leaf pubescence on abaxial surface, pairs
of lateral veins, petal length, petal width and sepal
width (over 0.65). Figure 8 shows that the first
principal component effectively separates C. sinensis
f. formosensis from C. sinensis var. sinensis and C.
sinensis var. assamica. The second component was
just on the level of reliability (Cronbach's alpha =
0.71), with negative loading on leaf width, and
positive loading on leaf serration density. The results
of this study also showed that characters with high
component loadings based on all characters were in
agreement with those only based on vegetative or
floral characters. The plot of the first two components
based on all characters (Fig. 8) shows a clearer
separation of C. sinensis f. formosensis from the other
Table 5. Loadings of the 27 vegatative and floral characters on the
first three components from NLPCA. Eigenvalues, percentage of
variance explained and cumulated, and Cronbach's Alpha are given
for each component.
Component
Character 1 2 3
Bud pubescence -0.95 0.19 0.10
Young branchlet pubescence 0.89 -0.21 -0.15
Leaf length -0.67 -0.62 -0.08
Leaf width -0.46 -0.81 0.05
Leaf thickness 0.15 -0.18 -0.38
Leaf shape -0.55 -0.05 -0.13
Leaf apex shape -0.65 -0.03 -0.06
Leaf base shape -0.26 -0.16 -0.20
Leaf serration density 0.26 0.71 0.11
Leaf pubescence on abaxial surface 0.73 -0.40 0.03
Abaxial midrib pubescence 0.89 -0.21 -0.15
Pairs of lateral veins -0.70 -0.49 0.07
Lateral vein angle at base -0.25 -0.25 0.69
Lateral vein angle at middle -0.52 -0.19 0.63
Petiole length -0.48 -0.35 -0.32
Petiole pubescence 0.89 -0.21 -0.15
Pedicel length 0.59 -0.23 0.30
Petal number per flower 0.27 -0.52 -0.02
Petal length 0.75 0.00 0.24
Petal width 0.71 -0.09 0.17
Sepal length 0.49 -0.20 0.25
Sepal width 0.66 -0.29 0.03
Sepal pubescence -0.20 -0.11 0.49
Filament length 0.29 -0.02 0.51
Style length 0.63 0.01 0.24
Ovary pubescence 0.91 -0.17 -0.14
Flower number per cluster -0.21 -0.49 -0.15
Eigenvalue 9.95 3.11 2.12
Variance explained (%) 36.80 11.50 7.90
Variance cumulative (%) 36.80 48.30 56.20
Cronbach's Alpha 0.93 0.71 0.55
related taxa than only on vegetative or floral
characters. It suggests that both vegetative and floral
characters should be taken into consideration for
distinguishing these taxa.
DISCUSSION
Morphological distinctiveness of C. sinensis f.
formosensis
The results of clustering analyses based on the
floral (Fig. 5) and all characters (Fig. 7) showed
almost a similar clustering pattern. In both
phenograms all C. sinensis f. formosensis samples
were grouped into a single cluster and clearly
separated from C. sinensis var. sinensis and C.
sinensis var. assamica. Although the phenogram
based on vegetative characters alone showed an
inconsistency position of C. sinensis f. formosensis
from eastern Taiwan (Group I-2, Fig. 3), the plot of
the first two components based on vegetative
characters indicated the eastern C. sinensis f.
formosensis is intermediate morphologically rather
March, 2007 Su et al.: Morphological comparisons of Taiwan wild tea plant and related taxa 79
10
-1-2
Component 1
2
1
0
-1
-2
Component 2
10
-1-2
Component 1
2
1
0
-1
-2
Component 2
10
-1-2
Component 1
2
1
0
-1
-2
Component 2
Fig. 8. Ordination plot of NLPCA based on vegetative and floral
characters. : C. sinensis f. formosensis. {: C. sinensis var.
assamica. c: C. sinensis var. sinensis.
than closed to the other two taxa (Fig. 4). Taken
together, the present study has shown that clear
morphological differences existed between C.
sinensis f. formosensis and two other closely related
taxa, and it seems not proper to treat it as the same as
C. sinensis var. sinensis as considered by Ming
(2000).
The present results show that reproductive
organs provide more informative characters for the
classification of tea plant than do vegetative
structures. This is in close agreement with that
previously reported by Banerjee (1992a). In general,
reproductive characters have been considered more
useful than vegetative features in plant systematics
(Stuessy, 1990).
In both cluster analysis and NLPCA, the
currently recognized varieties C. sinensis var.
sinensis and C. sinensis var. assamica integrated
considerably. Economic tea plants are heterogeneous
with many overlapping morphological attributes.
Most vegetative characters show a continuous
variation and a high degree of plasticity, and hence,
cannot be separated into discrete groups to identify
various taxa (Banerjee, 1992a). For the improvement
of tea quality, it did happen that artificial
hybridizations on the two taxa in the history
(Banerjee, 1992b). In this study, some materials of C.
sinensis var. sinensis and C. sinensis var. assamica
were collected from the wild in China or the
germplasm banks, which were thought to be
genetically independent. Other materials were
sourced from tea gardens which might possibly be
hybrids. These hybrids have intermediate
characteristics that may confuse their identification.
However, the individuals of C. sinensis f.
formosensis formed a clearly defined group, and were
never embedded in the group of C. sinensis var.
sinensis and C. sinensis var. assamica.
Hu (2004) used 15 leaf characters measured on
132 tea germplasms to evaluate inter-taxa variation
among C. sinensis var. sinensis, C. sinensis var.
assamica and C. sinensis f. formosensis. In the
scatterplot of PCA (Hu, 2004, Fig. 4), all individuals
of the three taxa showed two distinct groups.
Individuals from C. sinensis f. formosensis were
dispersed throughout both groups. This is
incongruent with present study. Two reasons may
explain these inconsistent results. First, characters
considered as diagnostic in the present study such as
bud pubescence, young branchlet pubescence,
abaxial midrib pubescence and petiole pubescence
were not used by Hu. Only 7 out of the 15 characters
(Hu, 2004, Table 10) adopted by Hu were used in the
present study, but these characters were not
significantly different among taxa in both studies.
Second, Hu transformed nominal characters to
ordinal variables for PCA analysis, and this would
produce results different from those derived from
NLPCA with the same characters. A comparison
between PCA and NLPCA showed that the NLPCA
would gain more loadings and led to a better
performance than PCA (Ellis et al., 2006).
Taxonomic rank of C. sinensis f. formosensis
As mentioned before, there has been controversial
regarding the appropriate taxonomic rank of C.
sinensis f. formosensis. Current study showed that C.
sinensis f. formosensis could be clearly discerned
from C. sinensis var. sinensis and C. sinensis var.
assamica in both cluster analysis and NLPCA. In
contrast, extensive overlap was found between C.
sinensis var. sinensis and C. sinensis var. assamica,
even though these two varieties have long been
recognized as distinct species (Chang, 1984) or
infraspecies (Kitamura, 1950; Sealy, 1958; Ming,
2000). Therefore, it is quite probable that C. sinensis
f. formosensis might deserve the species rank.
Further work, perhaps including molecular
approaches, may be necessary to resolve these
taxonomic questions.
80 TAIWANIA Vol. 52, No. 1
Morphological identification of C. sinensis f.
formosensis
Based on the results of morphological study and
NLPCA, several characters were found useful to
distinguish among these tea plants (Table 6). The
surface features of buds and ovaries with the highest
component loadings clearly separated C. sinensis f.
formosensis from C. sinensis var. sinensis and C.
sinensis var. assamica by the first principal
component. The buds of C. sinensis var. sinensis and
C. sinensis var. assamica are densely covered with
silver-yellowish hairs, whereas those of C. sinensis f.
formosensis are glabrous or partly covered with
sparse hairs. The surface features of buds have been
previously used to identify C. sinensis f. formosensis
by Suzuki (1937) and Kitamura (1950). The surface
of ovaries was also a reliable and stable character.
Ovaries of C. sinensis f. formosensis are glabrous
while those of C. sinensis var. sinensis and C.
sinensis var. assamica are pubescent. The importance
of reproductive characters in the taxonomy of
Camellia has been previously reported (Hsieh et al.,
1996).
Based on herbarium specimens, the flowering
time of C. sinensis f. formosensis extends from
September to January. After that period identification
can only be based on vegetative characters. In most
cases, young branches, abaxial midribs and petioles
of C. sinensis f. formosensis are glabrous, while those
of C. sinensis var. sinensis and C. sinensis var.
assamica are hairy. There are, however, some
inconsistencies that do not fit with the above
delineation. The samples of C. sinensis f. formosensis
from eastern Taiwan share some characteristics with
C. sinensis var. sinensis and C. sinensis var. assamica
such as sparsely hairy young branches, abaxial
midribs and petioles.
Other characters with higher component
loadings included leaf pubescence on abaxial surface,
pairs of lateral veins and petal size. However,
variation of these characters was continuous with
some degree of overlap across taxa and was not
considered to be the most taxonomically
discriminating. The angles between midrib and
lateral veins have been considered by local technical
personals to be useful characters for distinguishing.
They feel the angles were usually wider in C. sinensis
f. formosensis. However, component loadings of
these characters were low, indicating that the use of
these characters to discriminate among taxa is not
reliable.
Phytogeography of C. sinensis f. formosensis
The remarkable floristic similarity between
Taiwan and southeastern China has long been
recognized (Li, 1957; Hsieh, 2003). Migrations
between Taiwan and mainland China were facilitated
by the presence of the Taiwan Strait land bridge that
had connected Taiwan and mainland China several
times during the glacial ages. The close
morphological affinity between C. sinensis f.
formosensis and C. sinensis implied that this forma
may originated in mainland China.
There was a marked difference in distribution
patterns between the western and eastern populations
of C. sinensis f. formosensis (Fig. 1). C. sinensis f.
formosensis is distributed almost continuously
throughout the western side of Taiwan, while there is
only one population on the southeastern flank of the
Central Mountain Range. The eastern population was
likely the result of post-glacial range expansion of the
populations in western Taiwan. The little separation
between the western and eastern populations of C.
sinensis f. formosensis in vegetation characters
indicated that the vegetative characters of this plant
displayed a stronger response to environmental
variables than did floral characters. Future research
should explore different approaches, including the
use of different molecular markers, to establish the
post-glacial migration patterns of the Taiwan wild tea
plant and determine its origin.
Table 6. Useful characters to distinguish C. sinensis f. formosensis from C. sinensis var. sinensis and C. sinensis var. assamica. An asterisk (*)
denotes characters that are (nearly) decisive.
Character C. sinensis f. formosensis C. sinensis var. sinensis and C. sinensis var.
assamica
Bud pubescence* glabrous or sparsely covered with hairs at margins densely covered with silver-yellowish hairs
Young branchlet pubescence glabrous except for eastern populations nearly all hairy
Abaxial midrib pubescence glabrous except for eastern populations nearly all hairy
Petiole pubescence glabrous except for eastern populations nearly all hairy
Leaf pubescence on abaxial surface glabrous mostly hairy
Pairs of lateral veins 8-14, mostly > 10 5-12, mostly < 10
Ovary pubescence * glabrous pubescent
Petal length 0.7-1.3 cm 1.1-2.1 cm
Petal width 0.6-1.1 cm 0.7-2.0 cm
March, 2007 Su et al.: Morphological comparisons of Taiwan wild tea plant and related taxa 81
ACKNOWLEDGEMENTS
We greatly appreciate TAI, TAIF, PPI, Tea
Research and Extension Station and Prof. Tzen-Yuh
Chiang for their support and permission to examine
the related specimens. Financial support was
provided by the National Science Council of the
Republic of China (NSC-95-2422-H-002-002).
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March, 2007 Su et al.: Morphological comparisons of Taiwan wild tea plant and related taxa 83
利用數值分類方法比較臺灣野生茶與兩近緣分類群之形態特徵關係
蘇夢淮(1)、鄒稚華(2)、謝長富(1,3)
(收搞日期:2006 830 日;接受日期:2006 12 22 )
摘 要
臺灣野生茶 (Camellia sinensis f. formosensis) 乃泛指自生於臺灣中海拔山區的茶類植
物,臺灣植物誌將其中名稱為臺灣山茶。在分類史上,臺灣野生茶曾經被處理成數個不
同分類階層的歸屬。為了提供分類學者一個更客觀的看法,我們利用數值分類方法中的
群聚分析 (cluster analysis) 與非線性主成份分析 (nonlinear principal component analysis)
計算了 165 份標本中的 16 個營養及 11 個花部特徵的測量值,並依據分析結果來探究臺
灣野生茶與關係密切的茶 (C. sinensis var. sinensis) 和阿薩姆茶 (C. sinensis var. assamica)
的區別。將營養及花部特徵獨立或合併進行群聚分析的結果幾乎一致地指出臺灣野生茶
在形態特徵上的獨立性。相反地,茶與阿薩姆茶的形態差異卻無法被解析出來。非線性
主成份分析則顯現休眠芽與子房的毛被狀態是區分臺灣野生茶以及茶與阿薩姆茶的最
有效特徵。參照目前分類學家對茶與阿薩姆茶這兩群植物的分類處理,我們建議臺灣野
生茶應該被處理成種的階層會比較恰當。
關鍵詞:野生茶、臺灣山茶、臺灣、數值分類。
___________________________________________________________________________
1. 國立臺灣大學生態學與演化生物學研究所,106台北市羅斯福路41號,臺灣。
2. 中央研究院植物暨微生物學研究所,115 台北市研究院路 2128 號,臺灣。
3. 通信作者。Email: tnl@ntu.edu.tw
... 2001). Mong et al. (2006) studied the taxonomic position of the 'Taiwanese wild tea' has been called the "tea of the gods" and explored relationship with two closely related taxa, C. sinensis var. sinensis and C. sinensis var. ...
... Cluster-I i.e., Cluster-1a and Cluster-1b are also show homology at genetic distance of 3 wards, while group I and group II also show homology and genetic distance of their homology is 4.5 GD (Mohanan & Sharma, 1981;Wei et al., 2005). These taximetrics findings regarding tea cultivars are congruently in line with the previous research work of Mong (Mong et al., 2006). Hence, it proves that morphological features are not only of paramount significance in assessing tea yield but also they are key informative markers to identify and assign taxonomic status to certain species or varieties. ...
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Tea (Camellia sinensis (L.) O. Kuntze is the most consumed beverage in the world. Six genotypic cultivars of tea, namely Indonesian, Sri Lankan, Ruopi, Qi men, Chuye and Japanese were taxonomically characterized by using morphogenetic approach. Randomized Completely Block Design (RCBD) technique was applied in the analysis and results were presented in form of phenogram. Taxonomically genotypes were divided into two clusters with genetic distance (GD) 4.5. The cluster-I contained four cultivars bifurcating into two sub-linages with difference of GD 3. Intra-cluster GD of between genotypes Japanese and Sir Lankan was 2.0 GD and Chuye and Ruopi genotypes were closely associated with 1.8 GD. The cluster-II indicated the trait homology between Indonesian and Qi Men populations was 1.4 GD. This demonstrates that Qi men and Indonesian genotype are genetically more closely related than other genotypes, may be originated from one ancestor. The qualitative character evaluation was conducted to explore adaptability of these genotypes to the environment. The vein pairs per leaf was highest in Indonesian (13.60), followed by Qi men (12.08). The internode distance was highest in Qi men (3.7 cm) succeeded by Ruopi (3.6 cm). The branch angle to stem value was 35° in Qimen followed by Japanese with 38°. This morphogenetic analysis shows that Qimen and Sri Lankan genotypes are congruently adapted to the environment which depicts that area is appropriate for the tea plant growth and cultivation. This analysis also reflects that although these tea cultivars are phenetically similar to each other but can be differentiated by use of numerical analysis.
... Germplasm can be characterized using morphological, biochemical and molecular features descriptors that conformed to a set of standards specified in the descriptors for tea (Institute, 1997). In China (Chen and Zhou, 2005), China Taiwan (Su et al., 2007), India (Karthigeyan and Sud, 2010) and Sri Lanka (Piyasundara et al., 2009), morphological characteristics have been commonly used to evaluate the genetic variation of tea plants. In comparison to other crops, tea is known as a leavesusage beverage crop, and its distinctive properties are based on the chemical compounds in tea leaves. ...
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Many attentions have been previously focused to identify the multiple biochemical components related to tea quality and health benefits, however, the natural variation of biochemical components present in tea germplasm has not been adequately evaluated. In this study, the main biochemical components, leaf morphological and yield characteristics were evaluated for four rounds of tea leaves in a panel of 87 elite tea cultivars suitable for black, green, or oolong tea. Significant variations were observed among the tea cultivars, as well as seasonal differences in the levels of the free amino acid (FAA), caffeine (CAF), tea polyphenols (TP), water extract (WE) and TP to FAA ratio (TP/FAA). Results showed that the average levels of FAA showed a seasonal change, with the highest level of 4.0% in the 1st spring tea in the cultivars suitable for green tea and the lowest of 3.2% in summer tea in the cultivars suitable for black tea. The average CAF content was highest 3.2% in the cultivars suitable for oolong tea in the 1st spring and the lowest 2.5% in the cultivars suitable for green tea in summer. Limited seasonal and varietal variations were noticed in the average levels of WE among the three categories of tea. In addition, significant natural variation of the morphological characteristics, bud length varying from 2.5 cm to 8.7 cm, bud density from 190.3 buds · m-2 to 1 730.3 buds · m-2, mature leaves biomass from 128.4 kg · hm-2 to 2 888.4 kg · hm-2, and yield component traits of 100 buds (one bud with two leaves) dry weight from 3.7 g to 37.7 g, tea yield/round from 444.6 kg · hm-2 to 905.3 kg · hm-2, were observed. The aim of our evaluation was not only to identify the advantages of seasonal and clonal variations but also to provide a new viewpoint for their further application. Representative accessions were selected from the germplasm to promote the establishment of an inherent biochemical constituent expressing the quality of black, green, and oolong tea. The findings might be utilized to establish early selection criteria to enhance the tea breeding and production program.
... Naturally it is spread more in the tropical Asia (Martinez et al., 2003). Genus Camellia having 82 species belongs to family Theaceae, generally native to the high ground of south India (Lingaiah et al., 2011).Plant is medium sized, evergreen having branches in common condition (Mong and Hsieh, 2007). Tea is absolutely influenced because a permanent monoculture harvest (Jin, 2005). ...
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The present study was conducted at National Tea and High Value Crops Research Institute Shinkiari during 2020. Three different varieties of Camellia assamica processed black tea include P3, P5 and P9 were collected from PARC-National Tea and High Value Crops Research institute (P-NTHRI), Shinkiari, Mansehra, Pakistan. Presences of secondary metabolities in aqueous, methanolic and Ethanolic black tea extract was recognized by its colour intensity using standard chemical tests. Evaluation of tea samples was conducted to establish preference rating of tea for flavor, taste and color. Secondary metabolities like tannin, phenol, glycosides, terpenoids and protein are present in three extract of all tea samples while carotenoid are absent in all three solvent. Flavonoids and saponins are absent in all verities in Ethanolic and methanolic solvent while present in aqueous solvent while steroids and alkaloids are absent in P3 verities of aqueous solvent. Tea appearance in P3 is narrow while broad in P5 and P9 and fiber appearance of P3 is low while P5 is medium and P9 is higher. Aroma is more in all verities and Colour of P5 and P9 is dark brown and P3 is light brown. Tea without milk showed that the aroma of P5 is very strong as compared to other varieties of black tea P3 and P9. The taste of P3 is medium, P5 is better and P9 is highly better and colour of P3 is yellowish brown and P5 is brownish while P9 is dark brown and variation in color was seen from 5-30 mins. Yellowish color of P3 changed to dark brown after 20 minutes and dark brown of P9 changed into blackish brown after 20 mins while no color changed occurred in P5 variety. Tea with milk showed that the aroma of P3 was light and P5 and P9 was strong. Light yellow color was present in P3 and P5 and P9 showed bright yellow color. Taste of P9 was very strong and P3 was lighter while P5 was strong.
... Morphological traits such as leaf characters, fruit characters, flower characters etc., are used regularly for determination of genetic diversity among germplasm accessions. In tea, morphological characters have been widely used to study genetic diversity in China , Taiwan (Su et al. 2007), India (Rajkumar et al. 2010) and Sri Lanka (Gunasekara et al. 2001;Piyasundara et al. 2009). Phenotypic variation is positively associated with genetic diversity, yet also depends on environmental factors and the interaction between genotypes and environment (Wachira et al. 2002). ...
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Tea (Camellia sinensis L.) is the most widely-consumed beverage in the world. The biochemical components of tea leave include polyphenols (catechins and flavonoides), alkaloids (caffeine, theobromine, theophylline, etc.), volatile compounds, polysaccharides, amino acids, lipids and vitamins show a variety of bioactivities. Prolong cross-pollination nature of tea plants have produced considerable heritable variation, resulting in a high level of genetic diversity. The collection and conservation of the cultivars, landraces and wild relatives of the tea plant provides breeders with fundamental materials from which new cultivars are to be developed. The major role of tea breeding is to improve productivity, enhance tolerant to biotic and abiotic stress, and increase tea flavor and quality. Dissection of the genetic basis of these traits provides the potential for accelerating the breeding process by developing new tools such as marker-assisted selection. Therefore present review provides an overview of the biochemical and metabolite diversity of the global tea germplasm and its characterization and utilization.
... lasiocalyx (intermediate leaves), referred to as China, Assam, and Cambod varieties, respectively, are the sources of almost all the teas produced in the world [15]. Most of the commercially grown tea plant varieties in Taiwan include the small-leaf variety, which was previously imported from mainland China and is used for making green tea and some fermented teas, and the large-leaf variety, which was imported from India and is employed for making black tea [16,17]. There are also new varieties (large-, medium-, and small-leaf varieties) obtained through crossbreeding and seed selection using imported varieties at the Taiwan Tea Research and Extension Substation (TRES) and the native large-leaf wild tea from Taiwan (C. ...
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The major aim of made tea identification is to identify the variety and provenance of the tea plant. The present experiment used 113 tea plants [Camellia sinensis (L.) O. Kuntze] housed at the Tea Research and Extension Substation, from which 113 internal transcribed spacer 2 (ITS2) fragments, 104 trnL intron, and 98 trnL-trnF intergenic sequence region DNA sequences were successfully sequenced. The similarity of the ITS2 nucleotide sequences between tea plants housed at the Tea Research and Extension Substation was 0.379–0.994. In this polymerase chain reaction-amplified noncoding region, no varieties possessed identical sequences. Compared with the trnL intron and trnL-trnF intergenic sequence fragments of chloroplast cpDNA, the proportion of ITS2 nucleotide sequence variation was large and is more suitable for establishing a DNA barcode database to identify tea plant varieties. After establishing the database, 30 imported teas and 35 domestic made teas were used in this model system to explore the feasibility of using ITS2 sequences to identify the varieties and provenances of made teas. A phylogenetic tree was constructed using ITS2 sequences with the unweighted pair group method with arithmetic mean, which indicated that the same variety of tea plant is likely to be successfully categorized into one cluster, but contamination from other tea plants was also detected. This result provides molecular evidence that the similarity between important tea varieties in Taiwan remains high. We suggest a direct, wide collection of made tea and original samples of tea plants to establish an ITS2 sequence molecular barcode identification database to identify the varieties and provenances of tea plants. The DNA barcode comparison method can satisfy the need for a rapid, low-cost, frontline differentiation of the large amount of made teas from Taiwan and abroad, and can provide molecular evidence of their varieties and provenances.
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Thirty six tea genotypes (M1-M36) along with three check genotypes (Qi men, P3 and Indonesian) treated with gamma irradiation were evaluated with help of Random Amplified Polymorphic DNA (RAPD) markers. All the genotypes were treated with 10 Kr gamma radiations and grown by using augmented design. A total of 8 RAPD markers were used for PCR amplification of all 39 genotypes. Maximum polymorphic bands were recorded. However 16 unique extra alleles were found in the experimental samples. The genetic similarity values varied among genotypes. The phylogenetic analysis classified all the genotypes into six diverged groups (I-VI). The groups (I-VI) contained 17, 8, 4, 2, 7 and 1 genotype, respectively. Maximum variability in the allelic pattern was observed in treated samples. The variability in band patterns might be due to the mutation. The 3D analysis identified 4 elite tea genotypes (Qi men, M19, M28 and M31). The important tea genotypes treated with gamma radiations were characterized at molecular level. However further characterization through SSRs or SNP markers are needed to check further genomic variability at different gamma radiation treatments.
Chapter
In addition to the shortfalls described earlier in the “Chap. 2,” progress of tea breeding had also been slowed down due to the lack of reliable selection criteria (Kulasegaram 1980). Various morpho-biochemical markers had been reviewed in past (Wachira 1990; Singh 1999; Ghosh-Hazra 2001; Bandyopadhyay 2011; Mukhopadhyay et al. 2016), and it had been seen that they had marginally improved the efficacy of selection for desired agronomic traits in tea. This was mainly due to the fact that most of the morphological markers defined so far were influenced greatly by the environmental factors, a fact which is known for a long time, and hence showed a continuous variation with a high degree of plasticity. Therefore, these markers could not be separated into discrete groups for identification (Wickramaratna 1981). Recently, development of the molecular biology had resulted in alternative DNA-based markers for crop improvement of tea (Bansal et al. 2014). These markers can assist the process of traditional breeding with several efficacies. The greatest advantages of molecular markers are (1) free from the environmental influences and (2) detection of polymorphisms at an early stage of growth, and (3) accuracy of detection is much higher than the morphological markers. The different markers, which have been employed for varietal improvement of tea and its wild relatives, are reviewed below.
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Despite applications of breeding in tea are difficult, entire varietal development in tea and other Camellia species has been done through conventional breeding which started way back in 1939. Since then, several developments of genetics and breeding have taken place, with objectives of developing high different types of tea variety, increase yield, improvement cup tea quality, and resistance to biotic and abiotic stress. This is through tea genetic improvement mechanisms like conventional selection, hybridization, marker-assisted selection, mutation breeding, polyploidy, genetic engineering, and micro propagation. In this paper, (1) the achievements of tea genetic improvement and breeding, (2) the current situation of collection, conservation, appraisal and evaluation of tea germplasms, (3) Genetic introgression to enhance germplasm innovation (4) the establishment and development of tea breeding system and (5) the main research emphases of tea genetics and breeding soon were reviewed.
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The role of tea germplasm in crop improvement, though well recognized, yet lacks sufficient information depriving its optimum use. About 600 accessions are conserved as tea germplasm in Sri Lanka and only 4% have been frequently utilized in breeding. Floral morphological characters are useful descriptors for preliminary characterization of genetic resources and particularly pistil traits are considered as reliable criteria in taxonomical studies of higher plants. The objectives of the present study were to conduct a comprehensive analysis on floral diversity of tea germplasm to determine the nature and extent of genetic structure of tea germplasm and to categorize accessions into major taxa. Eighty-nine accessions from the tea germplasm were characterized using 16 floral traits. Results indicated presence of considerable variation among germplasm accessions. Accessions were categorized into five different groups based on the diversity of floral traits and highly discriminating accessions were identified based on the grouping pattern. Among the traits, pistil traits were highly variable compared to other traits. Tea germplasm is predominantly represented by Cambod type accessions (68%) followed by Assam types (20%). Availability of China type accessions is low. Gaps in the germplasm collection were identified and information generated can be used for decision making in future germplasm exploration missions and breeding programme.
Chapter
Among the several factors described in chapter 2, progress of tea breeding had also been slowed down due to the lack of reliable selection criteria. Thus various morpho-biochemical, digital markers such as artificial neural network, metallic markers, isozymes, are described. However, it had been seen that they had marginally improved the efficacy of selection for desired agronomic traits in tea. This was mainly due to the fact that most of these markers defined so far were influenced greatly by the environmental factors and hence showed a continuous variation with a high degree of plasticity. Recently, development of the molecular biology had resulted in alternative DNA-based markers for crop improvement of tea. These markers can assist the process of traditional breeding with several efficacies. The greatest advantages of molecular markers are: (1) free from the environmental influences and (2) detection of polymorphisms at an early stage of growth. The different DNA besed markers, which have been employed for varietal improvement of tea and its wild relatives, are reviewed in this chapter.
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The Taiwan vascular flora is exceptionally interesting not only because it is rich and diversified, but because it is of great phytogeographic significance owing to its geographic location. The flora of Taiwan, including naturalized plants, comprises 233 families and 1389 genera with 4216 species. In terms of major growth forms, there are 588 trees, 426 shrubs, 249 lianas, 177 vines, and 2776 herbs or ferns. Approximately 234 species are exotics typically associated with pastures, road clearings and other human disturbances. An extremely large percentage of these naturalized plants are of tropical New World origin. Among the native flora, the Orchidaceae (331 species), Gramineae (249), Compositae (194), Leguminosae (176), Cyperaceae (174), Rosaceae (105), Rubiaceae (93) and Euphorbiaceae (76) rank highest in numbers of species. Clearly, the greatest part of Taiwan's floristics richness comes from a wealth of species in primarily lowland (0–600 m asl.) taxa. A total of 2571 species were recorded in the lowlands, whereas only about 251 species occur between 3100-3950 m. Endemic genera are extremely scarce in Taiwan, with only four, namely Sinopanax (Araliaceae), Hayatella (Rubiaceae), Kudoacanthus (Acanthaceae), and Haraella (Orchidaceae). In contrast to the low percentage of generic endemism, there is a remarkably higher specific endemism. About 1041 species (26.1% of indigenous plants) are known only from Taiwan. A detailed examination of these species shows that there is a distinct trend of increasing endemism with increasing altitude (r² = 0.99). A survey of indigenous non-endemic species on the basis of their geographical distribution outside Taiwan shows that they can be classified into 6 major categories: 1. pantropical and palaeotropical species (1029 species); 2. species distributed in eastern Asia, from Himalayas through southern & eastern China to Taiwan, with some extending to the Ryukyus and Japan (1075 species); 3. widespread species extending from tropical Asia to eastern Asia (232 species); 4. species distributed in Japan and the Ryukyus (189 species); 5. species distributed in temperate and subboreal regions (221 species); and 6. cosmopolitan species (72 species). The main theme of western affinity of Taiwan flora is clearly indicated by almost 52% of the total flora (2069 species) which are also represented in China. The alliances of the flora are also pronounced with the Ryukyus and southern Japan (1618 species in common). However, the close relationship between Taiwan and Japan is through their mutual relation to the lowland flora of southern and eastern China. The tropical elements, mostly ranging from Malaysia to the Philippines, are well represented in the lowlands of Taiwan and particularly in the southernmost Hengchun Peninsula and the Lanyu Island off the southeast coast of Taiwan.
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Geographic variation in morphology, leaf anatomy, and flavonoid chemistry is appraised in Hebe pubescens, H. bollonsii, and allied plants from islands of outer Hauraki Gulf, North Island, New Zealand. The circumscription of H. pubescens is modified to include plants from Great Barrier, Little Barrier, and the Mokohinau Islands, as well as the Coromandel Peninsula and immediately surrounding islands. A new infraspecific classification is proposed for H. pubescens, wherein three subspecies are recognised. Subspecies pubescens occurs on the Coromandel Peninsula and immediately surrounding islands, and also at Papanui Point on the western coast of the Firth of Thames. Subspecies rehuarum occurs on Great Barrier Island. Subspecies sejuncta occurs primarily on the Mokohinau Islands and Little Barrier Island, with a single plant also known on Great Barrier Island. Populations of H. pubescens probably form a cline of variation, and some forms of subsp. sejuncta share morphological features with H. bollonsii, which is here considered restricted to the Poor Knights Islands, the Hen and Chickens Islands, and nearby areas of the Northland coast. Possible relationships of H. pubescens and H. bollonsii are discussed. A key to the taxa of Hebe pubescens is provided.
Article
Tea (Camellia sinensis [L.] O. Kuntze) is an important beverage crop in Taiwan. Most of the cultivated clones were introduced from China and India though some native wild teas are distributed in the mountains of central and southern Taiwan. In this study, 37 tea samples were evaluated using RAPD and ISSR markers. The samples comprised 21 clones of China, 3 clones of Assam, 7 hybrid clones between China and Assam tea, and 6 individual samples of native Taiwanese wild tea. A total of 53 and 56 polymorphic RAPD and ISSR markers respectively, were scored. The results of cluster analysis based on RAPDs revealed that three major groups could be recognized, i.e., cultivars of China tea and the cultivars developed in Taiwan from hybridization and selection; Assam tea; native Taiwanese wild tea. The native Taiwanese wild teas were, however, most distant in the clustering tree. In the ISSR dendrogram, Taiwanese native wild teas clustered closely with Assam tea then with China tea and the Taiwanese hybrid cultivars. The population gene diversity of the native wild tea was found to be the highest among the three populations studied. Analysis of molecular variance (AMOVA) revealed that the variance component within groups was larger than that among groups. The correlation coefficient between similarity matrices based on RAPD and ISSR was 0.811. A Mantel test revealed that the correlation was highly significant (p<0.001), indicating good congruence between the results of these two molecular markers.
Chapter
The history of plant breeding in tea is as chequered and as old as the discovery of wild tea in Assam and elsewhere. From the very early days of tea growing, it was recognized that breeding of tea creates problems that are somewhat unique to the plant. This is so because, firstly, unlike other woody perennials, in tea only a part of the total biomass constitutes the harvest, and secondly the plant is highly heterogeneous and self-incompatible. The ease with which cut-off branches of tea sprouted when they were buried in the soil was, however, noted (Watt, 1907). It was also noted that tea sets better with pollen from another bush, the average set of the plant with its own pollen being about one quarter of that obtained by cross pollination (Wight and Barua, 1939). This apart, selfing results in smaller seeds with reduced germinability or no seeds at all (Mamedov, 1961; Sebastiampillai, 1963). Consequently, the earlier breeding strategy relied on artificial pollination between plants different in some morphological features as a means for producing superior planting materials.
Chapter
The genus Camellia includes some 82 species which are mostly indigenous to highlands of south-east India (Sealy, 1958). Tea is the most important of all Camellia spp. both commercially and taxonomically. Since all Camellia spp. do not produce the brew that goes into the cup that cheers (Banerjee, 1988a), taxonomy plays a major role in the identification of true teas among the Camellia spp. for commercial exploitation. Many non-tea species of Camellia are however used as ornamental plants.
Article
The assignment of collections belonging to the genus Trema has been reassessed using numerical taxonomy methods. Observation in the field in Togo, West Africa, confirmed high phenotypic variation but not the previously asserted lack of character concordance within the T. guineensis complex. This study was carried out to assess the circumscription of taxa in the area. Distance between specimens was computed using the Gower's general coefficient of similarity and the non-parametric MODECLUS cluster analysis was used to discover how the specimens segregated. The existence of three clusters in the 158 specimen dataset using 44 morphological characters was suggested by MODECLUS. Canonical discriminant analysis supported the recognition of those three clusters using 40 morphological characters. Classificatory discriminant analysis showed that all specimens except one are identified correctly by the discriminant function. We suggest that three species be recognized in Togo and the neighboring countries: T. orientalis, T. africana, and T. nitens. A key to the species is included and the advantages of the method suggested here are discussed.
Article
Seventy-nine individuals representing all species of Raoulia, Ewartia, and Leucogenes, and selected species of Anaphalis, Cassinia, Gnaphalium, Helichrysum, Mniodes, and Pseudognaphalium, were scored for 165 morphological characters. Numerical phenetic analysis showed that, on the basis of overall morphological similarity, Raoulia encompasses two distinct groups worthy of generic rank, as well as several species of uncertain status. Preliminary indications for other genera in New Zealand are that Cassinia, Pseudognaphalium, and Anaphalis (which latter should include Helichrysum bellidioides) are distinct entities, while Helichrysum includes species of several genera, all outside Helichrysum sensu Hilliard & Burtt. Raoulia, Leucogenes, Ewartia, and part of both Gnaphalium sect. Euchiton and Helichrysum sect. Ozothamnus, form a suite of closely related genera.
Article
A morphometric analysis was conducted of Echinacea Moench (Asteraceae) to measure variation between native populations for taxonomic purposes. Data were collected from living and herbarium specimens. From a matrix of 321 specimens by 74 characters, a pair-wise distance matrix was computed using Gower's coefficient. Cluster strategies were explored from the distance matrix. MODECLUS clustering separated the data into two clusters, and a flexible agglomerative clustering method separated the data into the same two clusters, which were broken into four sub-clusters. Canonical discriminant analysis gave significant support for the two- and the four-cluster solutions. Canonical discriminant analysis also showed support for eight smaller clusters identified using McGregor's 1968 classification. We recognize two subgenera and four species: Echinacea subg. Echinacea contains only E. purpurea; Echinacea subg. Pallida contains E. atrorubens, E. laevigata, and E. pallida. The revised varieties are as follows: E. atrorubens var. atrorubens, E. atrorubens var. neglecta, E. atrorubens var. paradoxa, E. pallida var. angustifolia, E. pallida var. pallida, E. pallida var. sanguinea, E. pallida var. simulata, and E. pallida var. tennesseensis. A cladistic analysis was done on the four species. In the most parsimonious solution, E. purpurea was basally divergent to a clade of the other three species (70% bootstrap value), and all four were distinguishable by at least one apomorphy. A key to Echinacea taxa is provided, which should be valuable given the pharmaceutical and horticultural importance of Echinacea. Communicating Editor: Paul Wilson