Botanical Studies (2011) 52: 337-357.

SYSTEMATICS

Classifier modeling and numerical taxonomy of Actinidia (Actinidiaceae) in Taiwan

Tung-Yu HSIEH¹, Shin-Ming KU², Ching-Te CHIEN³, and Yun-Tsong LIOU⁴ *

¹Department of Horticulture, National Chung-Hsing University, 250 Kuo-Kuang Road, Taichung 402, Taiwan
²Herbarium (HAST), Research Center for Biodiversity, Academia Sinica, Nangang, Taipei 115, Taiwan
³Division of Silviculture, Taiwan Forestry Research Institute, Zhongzheng District, Taipei 100, Taiwan
⁴Miaoli District Agricultural Research and Extension Station, Council of Agriculture 261 Kung-Nan Kung-Kuan Miaoli 36344, Taiwan

(Received December 18, 2009; Accepted July 15, 2010)

ABSTRACT. Although kiwifruit (Actinidia) is popular worldwide, its complex morphology has resulted in long-standing confusion regarding its nomenclature, classification, and identification. In an attempt to resolve this issue, we conducted a phylogenetic analysis on 72 wild Actinidia accessions in Taiwan. We used 60 morphological characters as taxonomic traits to construct a seriated heat map that revealed A. callosa var. ephippioidea and A. rufa (sensu Flora of Taiwan, 2^nd Edition) as introgressive hybrids between A. chinensis var. setosa and A. callosa (sensu Flora of Taiwan, 2^nd Edition), as well as five significant groups of Actinida in Taiwan. Based on these results, we coded the indicator response matrix for the logistic regression models as dichotomizers and used a Bayesian discriminant model as a polychotomizer. After classifier modeling, the two classifiers were combined to identify Actinidia specimens in domestic and international herbaria. As a result, a taxonomic revision was made: A. callosa var. callosa and A. callosa var. ephippioidea were revised as A. rufa; A. rubricaulis was revised as A. callosa var. discolor; A. chinensis var. setosa was elevated to A. setosa; and A. tetramera was a misidentification of A. arguta. Of these, only A. setosa is endemic to Taiwan.

Keywords: Actinidia; Bayesian discriminant analysis; Flora of Taiwan; Heat map; Logistic regression classifier; Introgressive hybridization; Phylogenetic analysis.

INTRODUCTION

Kiwifruit, of the genus Actinidia, also known as Chinese gooseberry or mihoutao (in Chinese), enjoys worldwide popularity. Wild Actinidia is distributed throughout East and Southeast Asia from Siberia to Sumatra. Classical Chinese texts such as the Book of Odes (BC. 1046637), Bencao Gangmu (1596), and Zhiwu Mingshi Tukao-Changbian (1848), mentioned the genus and its primary uses as herbal medicine or edible fresh fruits (Warrington and Weston, 1990).

As Ferguson (1984) summarized, the first record of Actinidia is dated 1821, when Western botanist Wallich collected it in Nepal, assigning it number 6,634 in a catalog bearing his name. In 1836 Lindley named the genus Ac-tinidia (Greek aktis, a ray) based on its stylar arrangement and described the first species, A. callosa. After several years, additional species and varieties were discovered and published, including A. chinensis, published by Plan-chon in 1847, and A. eriantha and A. strigosa published by Bentham in 1860. Early classification of the genus,

however, was extremely confusing; many Actinidia species were initially placed in different genera. Actinidia latifolia was first placed in Heptaca (a doubtful genus in Tiliaceae) by Bentham in 1849, then in Kadsura (Schisan-draceae) by Miquel in 1861. Actinidia rufa, A. arguta, and A. polygama were first placed in Trochostigma in 1843, then transferred to Actinidia several years later. Actinidia kolomikta was variously placed in Prunus, Kalomikta, and Trochostigma before finally being identified as Actinidia by Maximowicz in 1859.

Dunn first revised the genus Actinidia in 1911, establishing two sections, Leiocarpae and Maculatae, and recognizing 24 species and almost 40 varieties or forms worldwide. Li (1952) carried out the second revision of Actinidia, establishing the sections Stellatae and Strigosae, and describing 36 species and over 50 varieties or forms. Since then, A. deliciosa has been domesticated in New Zealand and named "kiwifruit," after an endemic wingless bird there. An increasing number of botanists and horticulturists have since studied the physiology, biochemistry, cytology, biology, etc. of Actinidia, and kiwifruit's popularity has grown (Hsieh et al., 2004). In 1984, Liang completed a revision of Chinese Actinidia, recognizing over 50 species, about the same number of varieties, and more than a dozen

*Corresponding author: E-mail: liuyc@mdais.gov.tw.

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forms. Numerous cross-disciplinary studies indicated that the classification and placement of Actinidia remains confused, most likely due to its morphological complexity

(Warrington and Weston, 1990). Li et al. (2007) recently

revised the genus in China and Taiwan, describing 52 species, 44 of which are endemic.

The first preliminary report of Actinidia (A. callosa and A. championii) in Taiwan was published by Augustine Henry in 1896. Several years later, Dunn (1911) recorded a Taiwanese variety, A. callosa var. formosana, which was said to be originally published in Bulletin de la Societe Botanique de France 52(4): 20 by Finet and Gagnepain in 1905 (-1907). Hayata (1914) also recorded this taxon but treated it as A. championii. In 1919, Hayata described three new species in Taiwan, i.e., A. remoganensis, A. rankanensis, and A. arisanensis, and elevated A. cal-losa var. formosana to species rank, designating the type with a Latin diagnosis. The following year, he described another new species, A. gnaphalocarpa, which he had previously placed with A. championii. In 1936, Kanehira first recorded A. chinensis in Taiwan, which Li (1952) considered as a new variety, A. chinensis var. setosa, based on distinguishable features in leaf shape and hair types. For the same reason, Liang and Ferguson (1985) further elevated it to species rank, but other botanists continued to treat it as a variety of A. chinensis (Nee and Tsay, 1992; Peng and Lu, 1996; Li et al., 2007). Chou et al. (2008) in their paper on the characterization of the physicochemical and antioxidant properties of Taiwanese kiwifruit, chosed specific rank, A. setosa, as the scientific name. In 1984, Liang recorded four species and a new variety, A. callosa var. discolor, in Taiwan. Peng and Lu (1996) described seven species and one variety of Actinidia in the Flora of Taiwan, 2nd ed. (abbreviated as 'FOT2' below), four spe-cies and the variety being new records. Li et al. (2007), in their treatment of Actinidiaceae for Flora of China, record-ed five species and one variety in Taiwan, excluding previ-ous records of A. tetramera, A. callosa var. ephippioidea, and A. rubricaulis.

The examples above demonstrate the controversy in the classification of Actinidia. Although recent molecular studies have helped resolve phylogenetic and identification problems concerning Actinidia (Li et al., 2002; Zhao et al., 2007), they have not helped much in resolving Actinidia s classification and nomenclature problems. In 1989, Tang and Xiang first tried to use statistical methods to resolve the classification of the Clematoclethra complex (Actinidi-aceae) in China and pointed out that numerical taxonomy and statistics were effective ways to resolve classification issues of plant taxa, especially those with complex morphological characters. Xu et al. (1998) employed 11 leaf characters of A. chinensis cv. Tong-Shan no. 5 in conducting discriminant and cluster analyses to identify male and female plants. Discriminant analysis yielded a high rate of sexual identification of cultivars and cluster analysis implied that it was difficult to distinguish female from male plants by leaf characters. He et al. (2000a, b) selected

micromorphological characters of foliar trichomes and performed quantitative taxonomic analyses to study the classification and phylogenetic relationships of 27 Actinid-ia species and two varieties in China. The results indicated that genus Actinidia is a monophyletic group. Phylogenetic analysis of 22 morphological characters revealed two monophyletic groups within Actinidia in China (Li et al.,2000). Yang (2001) used 19 fruit traits from 12 taxa of Actinidia in China in a Q and R-type cluster analyses. The results showed that many traits of Actinidia are closely related. In 2006, Guo and Zhang first tried to use dielectric properties of kiwifruit and a back propagation network of an artificial neural network to clear up the classification of two A. deliciosa cultivars, 'Hayward' and 'Qinmei', which are hardly distinguishable from each other. The study only used 20 samples of each variety and got recognition rates of 100% on training samples and up to 90% on test samples. Furthermore, Cuong et al. (2007) employed a principal component analysis and cluster analysis to re-solve the nomenclature of the Actinidiaceae in Vietnam, and produced a taxonomic revision. Subsequently, Chen et al. (2008) selected 10 characters of the fruits and leaves of 11 Actinidia species to carry out a cluster analysis. The results implied that morphological traits of Actinidia are important for phylogenetic studies.

Previous research shows that both statistical methods and morphological characters are crucial to the taxonomic study of Actinidia. Thus far, all numerical taxonomic work on Actinidia has been on the native species of China and Vietnam. The purpose of this study is to use numerical taxonomy to first resolve Actinidia's classification and nomenclature problems in Taiwan, then complete a taxo-nomic revision of the genus.

MATERIALS AND METHODS

This study was divided into two stages, as follows:

1. Investigation of Actinidia field populations and numerical classification

This stage refers to previous taxonomic studies of Ac-tinidia, the field observations of the first author, and 60 selected botanical characters as investigated traits (Table 1), 34 of these traits being qualitative and 26, quantitative. A total of 72 wild Actinidia accessions were surveyed throughout Taiwan from 1999 to 2006, each of these comprises at least one mature male and female plant. All quantitative traits were measured in 100 samples of each accession, and average values to the second decimal place were considered representative of the operation values of each accession. All Actinidia accessions were identified according to FOT2; then coded, using the first two syllables of the Actinidia specific epithet as acronyms, followed by serial numbers. All species in the FOT2 but A. tetramera were represented in the 72 investigated accessions. These consisted of A. callosa var. callosa (coded Callo-01-19), A. callosa var. ephippioidea (one accession was coded Callo.

HSIEH et al. ― Classifier modeling and numerical taxonomy of Actinidia in Taiwan 339


Character	Character	Character states (code) for qualitative characters or unit for quantitative
code		characters
X1	Average length of leaf blade	cm
X2	Average width of leaf blade	cm
X3	Length/width index of leaf blade	cm/cm
X4	Symmetry of leaf blade base	asymmetrical (0); symmetrical (1)
X5	No. of pairs of secondary veins	number of pairs
X6	Deciduousness	evergreen (0); semi-deciduous (1); deciduous (2)
X7	Maculate of leaf blade	absent (0); present (1)
X8	Dorsal color of leaves	green (0); yellowish-green (1); pale-green (2)
X9	Degree of leaf upper-surface vestiture	hairless (0); sparsely haired (1); densely haired (2); very densely haired (3)
X10	Under-surface vestiture features	hairless (0); pubescent (1); villous (2); strigose (3); hispid (4); stellate (5)
X11	Degree of leaf under-surface vestiture	hairless (0); sparsely haired (1); densely haired (2); very densely haired (3)
X12	Length of leaf under-surface vestiture	hairless (0); short (1); intermediate (2); long (3)
X13	Leaf domatia	absent (0); present (1)
X14	Leaf blade texture	membranous (0); mesophytic (1); coriaceous (2); chartaceous (3); cartilaginous (4)
X15	Leaf blade margin	entire with sparse serrate (0); dentate and serrate (1); serrate (2); spinose (3); dentate (4); doubly serrate (5); spinose and dentate (6); spinose and serrate (7)
X16	Leaf blade apex	caudate (0); acuminate (1); acute (2); obtuse (3); truncated (4); emargin-ated (5)
X17	Leaf blade base	attenuate (0); cuneate (1); narrowly cuneate (2); obtuse (3); cordate (4); obliquely cordate (5)
X18	Petiole length	cm
X19	Petiole thick	cm
X20	Degree of petiole vestiture	hairless (0); sparsely haired (1); densely haired (2); very densely haired (3)
X21	Length of petiole surface vestiture	hairless (0); short (1); intermediate (2); long (3)
X22	Petiole vestiture features	hairless (0); pubescent (1); villous (2); strigose (3); hispid (4); stellate (5)
X23	Petiole color	green (0); yellowish-green (1); yellowish-red (2); light-red (3)
X24	Pith type of 1-year-old branch	solid (0); white lamellate (1); brown lamellate (2)
X25	Degree of young shoot vestiture	hairless (0); sparsely haired (1); densely haired (2); very densely haired (3)
X26	Young shoot vestiture features	hairless (0); pubescent (1); villous (2); strigose (3); hispid (4); stellate (5)
X27	Degree of 1-year-old branch vestiture	hairless (0); sparsely haired (1); densely haired (2); very densely haired (3)
X28	Length of 1-year-old branch vestiture	hairless (0); short (1); intermediate (2); long (3)
X29	One-year old branch vestiture features	hairless (0); pubescent (1); villous (2); strigose (3); hispid (4); stellate (5)
X30	No. of petals of staminate flower	number of petals
X31	Petal length of staminate flower	cm
X32	Petal width of staminate flower	cm
X33	Petal color of staminate flower	white (0); light-red (1); yellow (2)
X34	Sepal length of staminate flower	cm
X35	Sepal width of staminate flower	cm
X36	Sepal length/width index of staminate flower	cm/cm
X37	No. of sepals of staminate flower	number of sepals
X38	Filament length of staminate flower	cm

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Table 1. (Continuation)



Character		Character states (code) for qualitative characters or unit for quantitative
code		characters
X39	Anther color of staminate flower	purplish-black (0); yellow (1)
X40	No. of petals of pistillate flower	number of petals
X41	Petal length of pistillate flower	cm
X42	Petal width of pistillate flower	cm
X43	Petal length/width index of pistillate flower cm/cm
X44	Petal color of pistillate flower	white (0); light-red (1); yellow (2)
X45	Sepal length of pistillate flower	cm
X46	Sepal width of pistillate flower	cm
X47	No. of sepals of pistillate flower	number of sepals
X48	No. of styles of pistillate flower	number of styles
X49	Ovary length of pistillate flower	cm
X50	Anther color of pistillate flower	purplish-black (0); yellow (1)
X51	Fruit length	cm
X52	Fruit width	cm
X53	Length/width index of fruit	cm/cm
X54	Fruit color	green (0); greenish-brown (1); brownish-yellow (2)
X55	Degree of fruit vestiture	hairless (0); sparsely haired (1); densely haired (2); very densely haired (3)
X56	Length of fruit vestiture	hairless (0); short (1); intermediate (2); long (3)
X57	Fruit vestiture features	hairless (0); pubescent (1); villous (2); strigose (3); hispid (4); stellate (5)
X58	Fruit apex	obtuse (0); round (1)
X59	Fruit spot	absent (0); with minor spots (1); with obvious spots (2)
X60	Persistent calyx on fruit	absent (0); reflexed (1); reflexed and unreflexed (2); unreflexed (3)

ephip-01), A. latifolia (coded Lati-01-14), A. rufa (coded Rufa-01-03), A. arguta (coded Argu-01-10), A. chinensis var. setosa (coded Seto-01-12), and A. rubricaulis (coded Rubri-01-13), for a total of six species and one variety of Actinidia in Taiwan.

An Actinidia accession was taken as an operational taxonomic unit (OTU). The survey locations of Actinidia accessions are shown in Figure 1, excluding some that are endangered. Subsequently, all of the OTU data were filed in R language (Ihaka and Gentleman, 1996), measurements of similarity used Gower's similarity coefficient (Gower, 1971). R language was employed to analyze phylogenetic relationships among all Actinidia accessions (Paradis, 2006; Wiens, 2000). In order to understand the gradient relationships of all accessions in this study, we employed a simulated annealing algorithm and R language to create a Q-Q-type seriated heat map to elucidate all groups of Actinidia species. We then combined the results with the phylogenetic and gradient relationships to clarify the classification of Actinidia in Taiwan (Claude, 2008).

response matrix was coded and combined with the character variables of previous OTUs as training data to conduct model variable selection and classifier modeling. All data computing and analysis at this stage were programmed in R language. We selected logistic regression models as dichotomizers and a Bayesian discriminant model as a polychotomizer to be the classifiers of Actinidia in Taiwan. In the dichotomizers, the dependent variable of the logistic regression was designated to have a binomial distribution and employed the forward selection of a greedy algorithm and a stepwise regression for variable selection. We then used Akaike's information criterion to select the best variable combinations of the classifier regression models for each Actinidia species. In the polychotomizer, we took the Bayesian discriminant model as a multi-class classifier, then used a non-subjective prior for Bayesian discriminant modeling and variable selection until the iterations reached a 100% recognition rate for each taxa of Actinidia

(cf. Albert, 2009; Hastie et al., 2001). Consequently, the

dichotomizers and polychotomizer were combined to identify Actinidia specimens or images of specimens in the herbaria. Based on the results of the discriminant analysis, the correct scientific names of all specimens in this study were confirmed, and we completed a taxonomic revision of Actinidia in Taiwan.

2. Specimen identification and discriminant analysis

Based on the results of the previous stage, the indicator

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RESULTS

Classification of Actinidia accessions in Taiwan

Figure 2 shows the phylogenetic relationships between Actinidia accessions in this study. From the phylogenetic tree, A. chinensis var. setosa and A. rufa in the FOT2 were combined to form a separate branch. Although all phylo-genetic relationships between A. chinensis var. setosa accessions were very consistent, those between accessions of A. rufa were highly variable. Actinidia arguta, A. latifolia, A. callosa var. calloa, A. callosa var. ephippioidea, and A. rubricaulis formed another highly diversed branch; the branch with the highest diversity was found between A. callosa accessions, and the most consistent relationships were between accessions of A. arguta. On all branches, A. callosa var. callosa and A. callosa var. ephippioidea showed the closest phylogenetic relationship, followed by A. chinensis var. setosa and A. rufa. The remaining species, A. latifolia, A. arguta, and A. rubricaulis, were located on more independent branches. Furthermore, there were no significant phylogenetic correlations between Actinidia accessions and geographic distribution.

Figure 3 shows the Q-Q type seriated heat map of Ac-tinidia accessions, where 72 accessions were divided into 5 observable groups. On the map, accessions of A. rufa and A. callosa var. ephippioidea were between accessions of A. callosa var. callosa and A. chinensis var. setosa; thus creating a slightly fuzzy boundary for the A. callosa var. callosa group. During our investigations A. rufa and A. callosa var. ephippioidea were found only in the areas where A. chinensis var. setosa and A. callosa var. cal-losa accessions overlapped. Comparing all characters of the four taxa, we found that most characters of A. callosa var. ephippioidea and A. rufa were intermediate between those of A. chinensis var. setosa and A. callosa var. cal-losa, creating a series gradient phenomenon. However, A. rufa has some traits that also belong to A. chinensis var. setosa, so it was combined into a separate branch with A. chinensis var. setosa on the phylogenetic tree. In contrast, in terms of overall similarity, A. rufa and A. callosa var. ephippioidea were closer to A. callosa var. callosa, not A. chinensis var. setosa, which affected the boundary of the A. callosa var. callosa group on the seriated heat map. A comparison of the characters, wild habitats, population locations, previous studies (cf. Mallet, 2007; Peng and Ku, 2009; Suezawa, 1989) and the results of the seriated heat map of the aforementioned four taxa suggests that A. rufa and A. callosa var. ephippioidea represent natural hybrids between A. chinensis var. setosa and A. callosa var. cal-losa, which show a one-way introgression hybridization trend toward A. callosa var. callosa (cf. Wiens, 2000).

Natural hybrids led to the incongruency between the phylogenetic tree and seriated heat map. Actinidia callosa var. callosa was closer to A. rubricaulis on the phyloge-netic tree, but was next to A. chinensis var. setosa on the heat map because of hybridization between Actinidia cal-losa var. callosa and A. chinensis var. setosa. This also led

Figure 1. Distribution map of wild Actinidia accessions in this study. Some endangered accessions are not shown on this map. The code of accession names are based on identifications in the Flora of Taiwan 2nd ed. Lati-01-14 are A. latifolia. Callo-01-19 are A. callosa var. callosa. Argu-01-10 are A. arguta. Seto-01-12 are A. chinensis var. setosa. Rubri-01-13 are A. rubricaulis. Rufa-01-03 are A. rufa. Callo.ephip-01 is A. callosa var. ephip-pioidea.

Figure 2. Phylogenetic tree of Actinidia accessions in Taiwan. Actinidia accessions in the phylogenetic tree were coded based on the abbreviations of specific epithets from the Flora of Taiwan 2nd ed.,

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Table 2. Identification results of important Actinidia specimens in international herbaria. For the polychotomizers, 1 is for A. latifolia, 2 is for A. callosa, 3 is for A. arguta, 4 is for A. chinensis var. setosa, and 5 is for A. rubricaulis as per the Flora of Taiwan 2nd edition.


Specimen number in the
herbarium (Herbarium code or collector's number)	Polychotomizer	Dichotomizers	Description of specimens

50939 (HAST)	2	c	Voucher specimen of A. callosa Lindl. var. ephippioidea C. F. Liang in the FOT2
759 (HAST)	3	c	Voucher specimen of A. tetramera Maxim. in the FOT2
17446 (HAST)	2	c	Voucher specimen of A. rufa (Sieb. & Zucc.) Planch. ex Miq. in the FOT2
761 (HAST)	5	*	Voucher specimen of A. rubricaulis Dunn. in the FOT2
760 (HAST)	3	c	Specimen of A. callosa var. formosana Finet. & Gagnep.
46836 (HAST)	1	c	Voucher specimen of A. latifolia (Gardn. & Champ.) Merr. in the FOT2
758 (HAST)	2	c	Voucher specimen of A. callosa Lindl. in the FOT2
103514 (HITBC)	5	c	Specimen of A. callosa Lindl. of China
106761 (HITBC)	2	X	Specimen of A. callosa Lindl. of China
009839 (HITBC)	2	c	Specimen of Actinidia callosa Lindl. of China
106761 (HITBC)	2	X	Specimen of Actinidia callosa Lindl. of China
00122814 (IBK)	5	_*	Paratype of A. callosa var. discolor of China
00122790 (IBK)	5	c	Paratype of A. callosa var. discolor of China
00190557 (IBK)	5	c	Holotype of A. callosa var. discolor of China
00123032 (IBK)	4	X	Specimen of A. deliciosa (A. Chev.) C. F. Liang & A. R. Ferguson of China
00123033 (IBK)	4	X	Specimen of A. deliciosa (A. Chev.) C. F. Liang & A. R. Ferguson of China
00124359 (IBK)	1	c	Specimen of A. zhejiangensis C. F. Liang
00123011 (IBK)	4	X	Specimen of Actinidia chengkournsis C. Y. Chang of China
00122800 (IBK)	5	c	Specimen of Actinidia callosa var. discolor of China.
000229481 (K)	4	X	Type of A. chinensis Planch.
000229484 (K)	4	X	Type of A. chinensis Planch.
000442641 (K)	1	c	Type of Heptaca latifolia Gardn. & Champ. of Hong Kong
000442642 (K)	1	c	Type of Heptaca latifolia Gardn. & Champ. of Hong Kong
00428787 (NY )	2	_*	Type specimen of A. callosa Lindl.
00428824 (NY)	2	X	Type of A. tetramera Maxim.
222799 (TAI)	4	c	Specimen of A. chinensis var. setosa Li
183057 (TAI)	5	c	Specimen of A. rubricaulis Dunn of Taiwan
12718 (TAIE)	3	c	Specimen of A. arguta (Sieb. & Zucc.) Planch. ex Miq.
16684 (TAIF)	5	_*	Type of A. rankanensis Hayata.
16685(TAIF)	5	_*	Type of A. rankanensis Hayata.
083484 (TAIF)	2	c	Specimen of A. rufa (Sieb. & Zucc.) Planch. ex Miq. of Japan
16697 (TAIF)	1	c	Isotype of A. gnaphalocarpa Hayata.
083643 (TAIF)	1	c	Voucher specimen of A. latifolia (Gardn. & Champ.) Merr. in the FOT2
094191 (TAIF)	3	c	Specimen of A. arguta of Taiwan
080645 (TAIF)	3	c	Voucher specimen of A. arguta (Sieb. & Zucc.) Planch. ex Miq. in the FOT2
140149(TAIF)	2	c	Specimen of A. callosa Lindl. of Taiwan

HSIEH et al. ― Classifier modeling and numerical taxonomy of Actinidia in Taiwan 343

Table 2. (Continuation)


Specimen number in the
herbarium (Herbarium code or collector's number)	Polychotomizer	Dichotomizers	Description of specimens

099736(TAIF)	2	c	Specimen of A. callosa Lindl. of Taiwan
126875 (TAIF)	3	c	Specimen of A. callosa Lindl. of Taiwan
094740 (TAIF)	2	c	Specimen of A. callosa Lindl. of Taiwan
077374 (TAIF)	1	c	Specimen of A. latifolia (Gardn. & Champ.) Merr.
101270 (TAIF)	1	c	Specimen of A. latifolia (Gardn. & Champ.) Merr.
117529 (TAIF)	1	c	Specimen of A. latifolia (Gardn. & Champ.) Merr.
075479 (TAIF)	4	c	Specimen of A. chinensis var. setosa Li
096937 (TAIF)	1	c	Voucher specimen of A. latifolia (Gardn. & Champ.) Merr. in the FOT2
03141 (TI)	5	_*	Holotype of A. arisanensis Hayata.
03142 (TI)	5	_*	Holotype of A. rankanensis Hayata.
03143 (TI)	5	_*	Holotype of A. remoganensis Hayata.
T. Soma s.n. (TI)	2	c	Voucher specimen of A. formosana in Icones Plantanum Formosa-narum, Vol. VIII
Hayata s.n. (TI)	2	c	Voucher specimen of A. formosana in Icones Plantanum Formosa-narum, Vol. VIII (Ochobi)
Faurie s.n. (TI)	2	c	Voucher specimen of A. formosana in Icones Plantanum Formosa-narum, Vol. VIII
Hayata s.n. (TI)	1	c	Type of A. gnaphalocarpa Hayata. (Suisha)
Hayata s.n. (TI)	1	c	Type of A. gnaphalocarpa Hayata. (Uraisha)
S56391 (TNM)	3	c	Specimen of A. arguta of China
S10113 (TNM)	3	X	Specimen of A. chinensis Planch. var. setosa Li
S11128 (TNM)	5	_*	Voucher specimen of A. callosa Lindl. in the FOT2
S43479 (TNM)	2	c	Specimen of A. callosa Lindl. of Taiwan
S10031 (TNM)	2	c	Specimen of A. callosa Lindl. var. formosana Finet. & Gagnep.
S13435 (TNM)	5	c	Voucher specimen of Actinidia rubricaulis Dunn. in the FOT2
S5990 (TNM)	2	c	Specimen of A. callosa Lindl. var. formosana Finet. & Gagnep.
S12005 (TNM)	5	c	Voucher specimen of Actinidia rubricaulis Dunn. in the FOT2
S17543 (TNM)	5	c	Specimen of A. callosa Lindl. var. formosana Finet. & Gagnep.
S16534 (TNM)	5	c	Specimen of A. callosa Lindl. var. formosana Finet. & Gagnep.
S4902 (TNM)	2	c	Specimen of A. callosa Lindl. var. formosana Finet. & Gagnep.
S76931 (TNM)	1	c	Specimen of Actinidia latifolia (Gardn. & Champ.) Merr.
S085954 (TNM)	4	c	Specimen of Actinidia chinensis var. setosa Li
S072915 (TNM)	4	c	Specimen of Actinidia chinensis var. setosa Li
L0012511 (U)	2	X	Type specimen of Kadsura pubescens Miq.
L0012506 (U)	3	c	Type of Trochostigma arguta Siebold & Zucc.
L0012509 (U)	2	c	Type of Trochostigma rufa Siebold & Zucc.
L0012507 (U)	3	c	Type of A. cordifolia Miq.
1052327 (US)	4	c	Holotype of A. chinensis var. setosa Li
00458008 (US )	2	X	Isosyntype of A. rubricaulis Dunn

For the dichotomizers, a circle "o" indicates that the results match the polychotomizer and an "x" indicates that they do not match. An asterisk indicates that no characters were selected by the model for the specimens. The list is arranged alphabetically by herbarium acronym.

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to the placement of A. latifolia between A. callosa var. cal-losa and A. rubricaulis in the heat map. The irregular color gradations surrounding A. callosa on the heat map show its natural hybridization with A. chinensis var. setosa. Without the presence of hybrids, the map and the tree reflect comparable relationships.

As a result of this study, we divided Actinidia accessions into five groups in Taiwan: A. latifolia, A. callosa var. callosa, A. arguta, A. chinensis var. setosa and A. rubricaulis. We found no evidence of reproductive or geographical isolation between putative parents and the hybrids in the field.

X18, and X19 for the polychotomizer of Actinidia; then X1, X2, and X24 for the dichotomizer of A. callosa, X13 for A. arguta, X29 for A. chinensis var. setosa, X59 for A. rubricaulis, and X10 for the A. latifolia model. All the identification results of Actinidia specimens are shown in

Table 2.

From Table 2, all A. latifolia specimens were identified as "1" by the polychotomizer and accepted by the dichoto-mizer of A. latifolia, including the types of A. latifolia and A. gnaphalocarpa. These results imply that the scientific name of A. latifolia in the FOT2 (Peng and Lu, 1996) is the correct name, and that A. gnaphalocarpa is a synonym

of A. latifolia (used by the ICBN: McNeill et al., 2006).

The identification results of A. callosa specimens were very complex. The polychotomizer assigned the same group to the type specimens of A. rufa, A. callosa, and A. rubricaulis and the cited specimens of A. callosa, A.

Identification of Actinidia specimens and numerical nomenclature

From the results of classifier modeling, this study selected model variable combinations of X1, X2, X3, X5,

Figure 3. Seriated heat map of Actinidia accessions in this study. The codes are based on the abbreviations of specific epithets from the Flora of Taiwan 2nd ed.

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345

callosa var. ephippioidea, and A. rufa in the FOT2; The exception being TNM S11128, a specimen of A. callosa cited by the FOT2. Moreover, we were not able to investigate character X24 (pith type of 1-year-old branch) on the types of A. callosa because we had only type images on hand. Other specimens, including the type of A. rufa and cited specimens of A. callosa var. ephippioidea, A. rufa and A. callosa var. callosa in the FOT2, were accepted by the dichotomizer, except the type of A. rubricaulis. The results suggest that the names A. callosa var. callosa and A. callosa var. ephippioidea in the FOT2 should be revised to A. rufa.

The type specimens of A. arguta and A. cordifolia and the cited specimens of A. arguta and A. tetramera in the FOT2 were placed into the same group, and all were accepted by the dichotomizer of A. arguta, except the type of A. tetramera. The results imply that the scientific name of A. arguta in the FOT2 is the correct name, and that A. cordifolia is a synonym of A. arguta. In addition, the cited specimen of A. tetramera in the FOT2 should be corrected as A. arguta.

The type specimens of A. chinensis var. setosa and A. chinensis var. chinensis were classified into the same group by the polychotomizer, but only the A. chinensis var. setosa holotype was accepted by the dichotomizer of A. chinensis var. setosa. These results imply that A. chinensis var. setosa should be elevated to the species rank, but that A. setosa may be close to A. chinensis.

The types of A. callosa var. discolor, A. rankanensis, A. arisanensis, A. remoganensis, and the cited specimens of A. rubricaulis in the FOT2 were classified into the same group by the polychotomizer and accepted by the dichotomizer, but the type of A. rubricaulis was classified into another group and rejected by its dichotomizer. These results imply that A. rubricaulis does not exist in Taiwan; all cited specimens of A. rubricaulis in the FOT2 fell into the same group with types of A. callosa var. discolor, A. rankanensis, A. arisanensis and A. remoganensis. Liang (1984) and Li et al. (2007) synonymized A. rankanensis, A. arisanensis, and A. remoganensis under A. callosa var. discolor, which was taken up in this study. All specimens not mentioned above are discussed in detail below.

and Hayata, A. callosa var. formosana may be any of the following taxa: A. callosa var. discolor, A. rufa, A. arguta, and A. latifolia. The major obstacle to the clarifying this issue is the unavailability of protologues and authentic specimens of A. callosa var. formosana. Hayata (1919) elevated A. callosa var. formosana to the species rank and designated a type with Latin diagnosis. The specimens of A. formosana cited therein by Hayata, however, were classified into A. rufa by the polychotomizer and accepted by the dichotomizer. As a result, we have synonymized A. formosana under A. rufa in this paper.

The result of applying classifiers to A. setosa showed that all specimens of A. chinensis (including the types) and A. deliciosa were rejected by the dichotomizer. Our field investigations indicated that the flowering and fruiting periods of A. setosa differ from those of A. chinensis and A. deliciosa, when they were cultivated at the same locations (Chou et al., 2008; Hsieh, 2011). The geographic and phe-nological isolations and morphological distinctions among them uphold Liang and Ferguson's (1985) ranking of A. setosa as a species and not variety.

There has been long-standing confusion regarding the classification and nomenclature of A. rufa, A. callosa, and A. arguta. Actinidia callosa is the type of the genus, but the brief description of this species made it difficult to distinguish from other members of Actinidia. Actinidia arguta and A. rufa were published by Siebold and Zuc-carini in 1843, but their descriptions were also too brief to be useful. Furthermore, the authors mistakenly labeled their drawing of A. arguta fruit as that of A. rufa. Many botanists were thus even more confused about A. rufa, A. arguta, and A. callosa. Maximowicz advocated that A. rufa was a variety of A. arguta (Li et al., 2007). Makino (1901) treated A. arguta and A. rufa as varieties of A. cal-losa. In contrast, Dunn (1911) treated A. arguta as a variety of A. rufa. Such controversy on the classification and nomenclature of these species continued for a century. Taiwan has all three of these Actinidia species (Peng and Lu, 1996), thus providing an opportunity to clarify this issue. A cluster analysis conducted in the present study confirms that they belong in separate groups and the discriminant analysis clarified the correct names, resolving this long-standing issue.

What our results lack, however, is a model-selected character for type specimens and an investigation of accessions of A. callosa. Accepting the identification of foreign specimens without checking the dichotomizer is a highly risky matter (Hastie et al., 2001). Specimens labeled as A. callosa were a complex group that included some mis-identifications. Further studies are needed to determine the proper rank and phylogenetic relationships of A. callosa and its varieties.

Specimens in Table 2 showed some interesting features. Specimen S10113 (TNM), the sole specimen labeled A. chinensis var. setosa, was determined as A. arguta by the polychotomizer but then rejected by the dichotomizer of A. arguta. After re-examining the specimen, we found it to

DISCUSSION

Only two species of Actinidia were reported in Taiwan at the end of the nineteenth century: A. championii and the complex taxon A. callosa (Henry, 1895). Several years later, Dunn (1911) recorded a variety, A. callosa var. for-mosana, in Taiwan, which was originally published in Bulletin de la Societe Botanique de France 52(4): 20 by Finet and Gagnepain in 1905(-1907). We were unable, however, to find the published protologue of this variety despite a thorough search of that journal from the year 1896 to 1911. Hayata (1911) recorded A. championii in Taiwan, which he later considered it a misapplied name for A. callosa var. formosana (Hayata, 1914). From the descriptions of Dunn

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be Schisandra arisanensis (Schisandraceae). The classifiers of this study seem capable not only of identifying new species in Actinidia but also of picking out erroneously determined entities that do not belong to the genus.

Testolin et al. (1997) and Cipriani et al. (1998) studied some plastids and mitochondria sequences of 21 Actinidia taxa and pointed out that most Actinidia taxa, including A. arguta, A. rufa, A. callosa, and A. latifolia, can be separated by the chloroplast and mitochondrial DNA sequence. Subsequently, the phylogenetic study of Li et al. (2002) and Huang et al. (2002) based on nuclear ribosomal DNA internal transcribed spacers, chloroplast matK gene, and RAPD analysis showed that A. arguta are clearly distinguishable from the other species, including A. callosa var. discolor, A. latifolia and A. rufa inferred from either matK gene sequences, nrDNA ITS/5.8s region, or RAPD data. The latter three species appeared to be separable based on nrDNA ITS/5.8s region and RAPD data. Chat et al. (2004) studied the evolutionary relationships within Actinidia based on chloroplast, mitochondrial restriction site, and sequence data. Their study revealed reticulate evolution resulted from hybridization/introgression events and that A. arguta, A. callosa var. discolor, A. setosa, A. latifolia, and A. rufa can be separated based on 41 se-quences (rbcl and trnL-trnF) and restriction sites (matK and psbC-trnS). We believe that introgression may have contributed to the morphological variation as observed in these studies. By comparison with previous phylogenetic studies, our work revealed the same results on the phylo-genetic pattern and classification among all Actinidia at the species level. In comparison to the molecular methods, our methods are far more cost-efficient and simple. The heat map, a visual tools, is an ideal way to display introgres-sive hybridization. Most taxonomic studies of east Asian plants in recent years (e.g., Boyce and Wong, 2009; Chang et al., 2011; Chen and Chou, 2008; Chen et al., 2008,

2009; Chung et al., 2008, 2010; Cong et al., 2008; Dong, 2010; Gao and Yang, 2009; He and Zhang, 2010, 2011; Hou et al., 2009; Hsieh, 2002; Hsieh et al., 2002, 2007; Hsu et al., 2011; Hughes et al., 2011; Ku, 2006; Ku et al., 2008; Lammers and Klein, 2010; Lin et al., 2010; Liu et al., 2007, 2009, 2010, 2011; Liu and Yang, 2010, 2011a,b; Lu and Wang, 2009; Mou and Zhang, 2010; Peng et al., 2007a,b, 2008a,b, 2010; Sam et al., 2009; Sheue et al., 2009, 2010; Su et al., 2009; Tian et al., 2010; Tsai et al., 2003; Wang et al., 2010; Wong and Boyce, 2010a, b; Wu et al., 2009; Yang, 2009; Yu et al., 2009; Yuan and Yang, 2009; Yuan et al., 2011; Zhang and He, 2009a,b; Zhang et al., 2008, 2010) have not combined classification and nomenclature with statistics. Considering the diversity and complexity of plants, more models and methods should be developed and utilized for future studies of important and complex taxa (Cuerrier et al., 1998).

1. Leaf abaxially glabrous, with hairy domatia in axils of lateral veins ...................................................3. A. arguta

1. Leaf abaxially glabrous or hairy, axils of lateral veins without domatia.

2. Pith of 1-year-old branchlets not lamellated ..............

........................................... 5. A. callosa var. discolor

2. Pith of 1-year-old branchlets lamellated.

3. Branchlets densely hispid .................... 4. A. setosa

3. Branchlets not hispid.

4. Mature leaf densely stellate abaxially ...............

..................................................... 1. A. latifolia

4. Mature leaf not or rarely stellate abaxially........

........................................................... 2. A. rufa

1. Actinidia latifolia (Gardner & Champ.) Merr., J. Straits

Branch Roy. Asiat. Soc. 86: 330. 1922; Li, J. Arnold

Arbor. 33: 49. 1952; Li, Woody Fl. Taiwan 571. 1963; Li, Fl. Taiwan 2: 588. 1976; Liu et al., Tr. Taiwan (2nd ed.) 444. 1994; Peng & Lu, Fl. Taiwan (2nd ed.) 2: 659. 1996; Li et al., Fl. China 12: 343. 2007. Figure 4

Heptaca latifolia Gardner & Champ., Hooker's J. Bot.

Kew Gard. Misc. 1: 243. 1849.

Actinidia championii Benth., Fl. Hongk. 26. 1861; Dunn.,

J. Linn. Soc., Bot. 39: 407. 1911.

Actinidia gnaphalocarpa Hayata, Icon. Pl. Formos. 9: 7.

1920.

Large climbing woody vine, deciduous to semi-evergreen. Branchlets glabrous, slightly puberulent or densely tomentose when young; pith white lamellate or hollow when old. Petiole 3-6 cm; leaf blade abaxially pale-green, adaxially green, usually broadly ovate to obovate, 7-14 X 4.5-9 cm, abaxially densely appressed stellate tomentose, glabrescent when old, base broadly cuneate to rounded or reniform, margin minutely and remotely callose-serrulate, apex acute to acuminate. Inflorescences 2-4 branched, 8 or more flowered, densely brownish tomentose. Flowers yellowish-brown. Sepals 3, ovate, 4-5 mm, reflexed after anthesis, both surfaces tomentose. Petals 5, oblong to obo-vate-oblong, 6-8 mm, reflexed after anthesis. Ovary globose, ca. 2 mm, densely pilose. Fruit brown, subglobose to ovoid, 1.2-2.1 X 0.7-1.5 cm with lenticels, glabrous when mature or sparse tomentose, especially both base and apex of fruit.

Specimens examined. TAIPEI COUNTY (CO.): Wulai Township, Uraisha, B. Hayata s.n. (TAIF 16697; TI). TAOYUAN CO.: Fuhsing Township, Lalashan, ca. 1,500 m elev., 19 June 1994, Wen-Pen Leu 2045 (HAST 46836). ILAN CO.: Tatong Township, Mingchih, ca. 1,100-1,200 m elev., 22 June 1994, Her-Long Chiang s.n. (TAIF 101270). TAICHUNG CO.: Hoping Township, Tahsuehshan, ca. 1,675 m elev., 15 Aug 2001, C. M. Wang 5234 (TNM S76931). NANTOU CO.: Yuchih Township, Lienhuachih, ca. 600 m elev., 10 Oct 1995, Kuoh-Cheng Yang 4731 (TAIF 077374); Lienhuachih, ca. 650 m elev., 6 June 1985, Sheng-You Lu 16482 (TAIF 096937); Smsha,

Taxonomic treatment of Actinidia in Taiwan

Key to the taxa of Actinidia in Taiwan (excluding natural hybrids).

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347

Hayata s.n. (TI). KAOHSIUNG CO.: Maolin Township,

Fengkang logging road, ca. 1,600 m elev., 4 July 2000,

Her-Long Chiang 1299 (TAIF 117529). TAITUNG CO.:

Taimali Township, Taimali working station, ca. 500-1,000 m elev., 15 Aug 1993, Jenn-Che Wang et al. 8762 (TAIF

083643).

Etymology. The specific epithet 'latifolia, means "broadly leaved"

Distribution and habitat. China, Taiwan, Cambodia, Laos, Thailand, and Vietnam. In Taiwan, occurs in forests and thickets, on slopes, and along roads throughout Taiwan at 300-2,200 m.

Pin Cheng s.n. (TAIF 140149). TAICHUNG CO.: Hoping Township, Chingshan-Techi, ca. 1,320 m elev., 28 July

1997, C. M. Wang 2749 (TNM S043479). KAOHSIUNG

CO.: Liouguei Township, Shanping-Nanpengshan, 6 Apr. 1987, C. H. Ou et al. s.n. (TNM S5990). PINGTUNG

CO.: Chunrih Township, Tahanlintao, 29 Sept. 1985, C. H. Ou et al., s.n. (TNM S4902). TAITUNG CO.: Tajen Township, Kueitien, 12 Aug 1994, Sheng-You Lu s.n. (TAIF

094740).

Etymology. The specific epithet'rufa, means "reddish".

Distribution and habitat. Japan and Taiwan. In Taiwan, occurs in forests and thickets, and along streams and roads throughout Taiwan at 150-2,200 m.

2. Actinidia rufa (Sieb. & Zucc.) Planch. ex Miquel, Ann. Mus. Bot. Lugduno-Batavi 3: 15. 1867; Dunn., J. Linn.

Soc., Bot. 39: 402. 1911; Peng and Lu, Fl. Taiwan (2nd ed.) 2: 660. 1996; Li et al., Fl. China 12: 337. 2007.

Figure 5

Trochostigma rufum Sieb. & Zucc., Abh. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. 3(2): 727. 1843. ；

Actinidia arguta var. rufa (Sieb. et Zucc.) Maxim., Bull. Acad. Sci. St. Petersb. 31. 19. 1886; Li, J. Arnold Arbor. 33: 34. 1952.

Actinidia callosa var. rufa (Sieb. et Zucc.) Makino, Bot. Mag. (Tokyo) 15: 147. 1901.

Actinidia callosa auct. non Lindl.: Peng & Lu, Fl. Taiwan

(2nd ed.) 2: 657. 1996.

Actinidia callosa var. ephippioidea auct. non C. F. Liang:

Peng & Lu, Fl. Taiwan (2nd ed.) 657. 1996.

Actinidia formosana Hayata, Icon. Pl. Formos. 8: 12.

1919.

Deciduous to semi-evergreen climbing woody vine. Branches glabrous; pith brown lamellate; branchlets reddish, brownish puberulent. Petiole 2-7 cm, glabrous; leaf blade ovate to broadly ovate or orbicular, 4.5-13 X 3.38.5 cm, papery, lateral veins 5-8 pairs, base rounded to truncate or cordatulate, oblique or not, margin shallowly mucronate-serrate, sometimes glandular, apex obtuse to long acuminate. Inflorescences cymose, axillary, brownish velutinous. Male inflorescences multi-flowered. Female inflorescences with fewer flowers then male. Flowers white, often reddish at base. Sepals 4-5, ovate, ca. 4.8-5.7 mm, apex acute to round. Petals 5, obovate, ca. 1 cm. Ovary globose, ca. 4.5-5.7 mm, densely tomentose. Fruit oblong to ovoid, 2.3-4.5 cm, densely or sparsely tomentose to glabrous when mature, lenticels obscure.

Specimens examined. TAIPEI CO.: Kelung, S. Soma s.n. (TI); Daiton, 1903, U. Faurie s.n. (TI). TAIPEI CITY: Matsao, ca. 600-650 m elev., 6 Oct. 1985, Ching-I Peng

8644 (HAST 758). ILAN CO.: Su-ao Town, Wushihpi, 17 Sept. 1992, C. K. Lin s.n. (TNM S10031); Ochobi, May

1916, B. Hayata s.n. (TI); Tatong Township, no. 100 logging road, ca. 1500 m elev., 17 Sept. 1996, C. M. Wang 2219 (TAIF 099736). HUALIEN CO.: Sioulm Township, Hoping logging trail, ca. 1,200 m elev., 12 June 2001, Yu-

3. Actinidia arguta (Sieb. & Zucc.) Planch. ex Miquel,

Ann. Mus. Bot. Lugduno-Batavi 3: 15. 1867; Li, J. Arnold Arbor. 33: 31. 1952; Liang, Fl. Reipubl. Popularis

Sin. 49(2): 205. 1984; Peng and Lu, Fl. Taiwan (2nd ed.) 2: 657. 1996; Li et al., Fl. China 12: 337. 2007.

Figure 6

Trochostigma argutum Sieb. & Zucc., Abh. Math.-Phys.

Cl. Konigl. Bayer. Akad. Wiss. 3(2): 727. 1843.

Actinidia cordifolia Miq., Ann. Mus. Bot. Lugduno-Batavi

3: 15. 1876.

Actinidia callosa var. arguta (Sieb. et Zucc.) Makino, Bot.

Mag. (Tokyo) 15: 148. 1901.

Actinidia rufa var. arguta (Sieb. & Zucc.) Dunn, J. Linn.

Soc., Bot. 39: 402. 1911.

Actinidia rufa var. cordifolia (Miq.) Dunn, J. Linn. Soc.,

Bot. 39: 403. 1911.

Actinidia arguta var. cordifolia (Miq.) Bean, Trees Shrubs

Brit. Isles 1: 162. 1914.

Actinidia tetramera auct. non Maxim.: Peng & Lu, Fl. Taiwan (2nd ed.) 2: 660. 1996.

Deciduous woody twiner. One-year-old branchlets glabrous or puberulent when young; 2nd-year branches grayish-brown, glabrous; pith white to brown, 1-year shoot lamellate. Petiole green or sometimes pinkish-yellow when young, 2-5 cm, glabrous; leaf blade abaxially green, adaxi-ally dark- or pale-green, ovate to broadly ovate, rarely ovate-oblong, 5-11 X 4-10 cm, papery, abaxially glabrous, with hairy domatia in axils of lateral veins, lateral veins 4-7 pairs, straight or arcuate-ascending, base rounded to cordate, symmetrical, margin with sharply serrate teeth, apex abruptly acuminate. Inflorescences cymose, axillary or lateral, 1-7-flowered. Flowers white, 1.4-2.3 cm in diam. Sepals 4-5, ovate to oblong, glandular-tomentose. Petals -5, obovate to orbicular, 7-9 mm. Ovary long bottle-shaped, 6-7 mm, glabrous. Fruit green when mature, globose to oblong, 1.6-2.7 cm, glabrous, without lenticels or persistent sepals.

Specimens examined. ILAN CO.: Yuanshan Township,

Ayushan, 10 July 1996, Yu-Pin Cheng s.n. (TAIF 092148);

Tatong Township, Taipingshan, ca. 1,950 m elev., 8 June 1985, Ching-I Peng 7872 (HAST 759). TAOYUAN CO.:

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Fusing Township, Lalashan, ca. 1,900-2,000 m elev., 5

Aug 1999, Su-Wen Chung 2078 (TAIF 126875). TAIC-

HUNG CO.: Hoping Township, Szuyuanyakou, ca. 1,9002,200 m elev., 13 Aug 1993, Chieh-Lin Huang et al. 36

(TAIF 080645); Szuyuanyakou, ca. 1,915 m elev., 27 July

1998, Tsai-Wen Hsu 9094 (TAIE 12718); Jiayang, no. 810

logging road, ca. 1,900 m elev., 26 June 1998, Ching-Kuoh

Liou et al. 977 (TAIF 094191). CHIAYI CO.: Alishan

Township, Tongpu, 26 May 1960, T. I. Chuang et al. 4097

(HAST 760).

Figure 4. Actinidia latifolia (Gardner & Champ.) Merr. A, Habit (Tengchi, Kaohsiung County); B, Flowering branch; C, Fruiting branch; D, Male flowers; E, Overy; F, Fruits, showing cross sections; G, Female flower; H, Developmental stages from flower buds to mature fruits; I, Leaf, abaxial surface; J, Pith of one-year old shoot.

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349

Etymology. The specific epithet arguta means sharp teeth" in reference to the leaf blade margin.

Distribution and habitat. Siberia, Japan, Korea, China, and Taiwan. In Taiwan, occurs in mountain forests and along streams of the northern and central parts at 1,300-

2,600 m.

4. Actinidia setosa (H. L. Li) C. F. Liang & A. R. Ferguson, Guihaia 5 (2): 72. 1985. Actinidia chinensis auct. non Planchon: Susuki in Masamune, Short Fl. Form.

Figure 5. Actinidia rufa (Sieb. & Zucc.) Planch. ex Miquel. A, Habit (Yangmingshan, Taipei County; Callo-05); B, Flowering branch; C, Fruiting branch; D, Male flower; E, Female flower; F, Different shapes of leaves; G, Domatia in axils of lateral veins; H, Fruit; I, Pith of one-year old shoot.

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137. 1936; Kanchira, Formos. Trees (rev. ed.) 449. pl.

406. 1936. Figure 7

Actinidia chinensis var. setosa H. L. Li, J. Arnold Arbor. 33: 56. 1952; Li, Woody Fl. Taiwan 573. 1963; Li,

Fl. Taiwan 2: 588. 1976; Liang, Act. Phytotaxon. Sin.

13(4): 33. 1975; Liu et al., Tr. Taiwan (2nd ed.) 444. 1994; Peng & Lu, Fl. Taiwan (2nd ed.) 2: 659. 1996; Li et al., Fl. China 12: 350. 2007.

Large climbing vine, deciduous. Branchlets reddish, young branchlets densely hispid; pith lamellate, whit-

Figure 6. Actinidia arguta (Sieb. & Zucc.) Planch. ex Miquel. A, Habit (Szuyuanyakou, Taichung County; Argu-02); B, Flowering branch; C, Fruiting stem; D, Female flower; E, Male flower; F, Five-sepal flower; G, Four-sepal flower; H, Fruit; I, Leaf, adaxial sur-fae; J, Leaf, abaxial surface; K, Leaf margin; L, Domatia in axils of lateral veins; M, Pith of one-year old shoot.

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ish to brown when mature. Petiole 3-5 cm, hispid; leaf blade abaxially pale-green, adaxially dark-green, broadly ovate to broadly obovate or suborbicular, 6-21 x 6-16 cm, chartaceous, abaxially brownish stellate tomentose, adaxially scabrid-hispid, lateral veins 5-8 pairs, base cor-datulate, margin setose-serrulate with teeth, apex acute or

shortly acuminate to acuminate. Inflorescences cymose, 1-4-flowered, white to yellowish-brown. Flowers white to orangish-yellow when mature. Sepals 5(-7), broadly ovate to oblong-ovate, 6-10 mm. Petals -5(-8), broadly obo-vate, 1-2 cm shortly clawed at base, apex rounded. Ovary globose, ca. 5 mm in diam. Fruit subglobose, cylindric to

Figure 7. Actinidia setosa (H. L. Li) C. F. Liang & A. R. Ferguson. A, Habit (Hehuan river, Taichung County; Seto-04); B, Flowering branch; C, Fruiting branch; D, Female flower; E, Male flower; F, Seven-petal flower; G, Female flower with six petals and sepals; H, Sections of fruits; I, Leaf margin; J, Leaf, abaxial surface; K, Pith of one-year old shoot.

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obovoid or ellipsoidal, 3.6-7.4 cm, densely hispid, with lenticels; persistent sepals reflexed.

Specimens examined. ILAN CO.: Tatong Township, Taipingshan, ca. 1,870 m elev., 14 May 1992, S. F. Huang

4779 (TAI 222799). HSINCHU CO.: Wufeng Township,

Kuanwu, ca. 2,000 m elev., 21 May 1994, Jenn-Che Wang

et al. 9224 (TAIF 075479). MIAOLI CO.: Taian Township,

Kuanwu, ca. 2,350 m elev., 9 May 2003, C. M. Wang et al.

6707 (TNM S085954). TAICHUNG CO.: Hoping Township, no. 710 logging track, ca. 1,800 m elev., 24 July 24, Y.

C. Lu 112 (HAST 23846). NANTOU CO.: Hsinyi Township, Yushankou, ca. 2,300 m elev., 6 May 2001, C. M.

Figure 8. Actinidia callosa var. discolor C. F. Liang. A, Habit (Lixing industrial road, Nantou County; Rubri-03); B, Flowering branch; C-D, Fruiting branch; E, Male flower; F, Female flower; G, Different shapes of fruits; H, Sections of fruits; I, Pith of one-year old shoot; J, Leaf, abaxial surface.

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Wang et al. 4974 (TNM S072915). CHIAYI CO.: Alishan

Township, Alishan, 18 Oct 1918, E. H. Wilson 10802 (US 1052327).

Etymology. The specific epithet 'setosa, means "bristly hairy", based on this plant being bristly hairy throughout.

Distribution and habitat. Endemic to Taiwan, in mountain forests, on slopes, and along roads throughout Taiwan

at (500-) 1,300-2,700 m.

leaf margin with callose teeth. The variety epithet 'discolor' refers to the leaf blade with different colors on the two sides.

Distribution and habitat. China and Taiwan. In Taiwan, at forest margins, on slopes, in thickets and valleys, and along roads throughout Taiwan at 300-2,100 m.

Insufficiently known taxon:

5. Actinidia callosa var. discolor C. F. Liang in K. M.

Feng, Fl. Reipubl. Popularis Sin. 49(2): 315. 1984; Li et al., Fl. China 12: 343. 2007. Figure 8

Actinidia callosa auct. non Lindl.: Peng & Lu, Fl. Taiwan

(2nd ed.) 2: 657. 1996, pro part.

Actinidia arisanensis Hayata, Icon. Pl. Formos. 8: 11.

1919.

Actinidia rankanensis Hayata, Icon. Pl. Formos. 8: 13.

1919.

Actinidia remoganensis Hayata, Icon. Pl. Formos. 8: 13.

1919.

Actinidia rubricaulis auct. non Dunn: Peng & Lu, Fl. Taiwan (2nd ed.) 2: 660. 1996.

Deciduous to semi-evergreen twiner. Branchlets glabrous, lenticels conspicuous; pith of 1-year-old shoot solid, rarely lamellate; old branches grayish, pith solid or inconspicuously brown lamellate. Petiole glabrous; leaf blade abaxially pale-green, adaxially dark-green, elliptic or ovate to obovate, rarely elliptic, 3.6-10 x 3-6 cm, abaxially glabrous, lateral veins 4-7 pairs, base cuneate to obtuse, margin coarsely serrate or serrate to subentire, apex obtuse or acute. Inflorescences cymose, 1-5-flowered, glabrous; peduncles 0.7-1.5 cm; pedicels 1.1-1.7 cm. Flowers white. Sepals 5, ovate, 4-5 mm, glabrous. Petals 5, obovate, 8-10 mm. Ovary subglobose, densely pubescent; Fruit grayish-green, ovate to oblong, 1.2-2.7 cm, glabrous, lenticels white, conspicuous.

Specimens examined. TAIPEI CO.: Wulai Township, Kang-gu, 17 June 1955, Hsuen Kao and Muh-Tsuen Kao 2904 (HAST 761); Houkengtzechi, 9 Nov. 1980, H. N. Yang 3436 (TAI 183057); Remogan, 7 May 1916, B. Hayata s.n. (TI). ILAN CO.: Nan-ao Township, Rankan-zan, 12 May 1916, B. Hayata s.n. (TAIF 16685, 16684; TI). NANTOU CO.: Ren-ai Township, Meifen, ca. 2,100 m elev., 13 July 1993, C. M. Wang 80 (TNM S11128); Yuchih Township, Lienhuachih, ca. 576-925 m elev., 19 Sept. 1995, Liang Hung Wu 77 (TNM S17543); Luku Township, Fenghuangku, ca. 750-850 m elev., 11 Sept. 1994, Kuang-Yuh Wang 168 (TNM S16534). HUALIEN CO.: Chohsi Township, Yamagon to Huangma, ca. 800-1300 m elev., 1 Aug. 1993, Tseng-Pin Chiang 66 (TNM S13435). CHIAYI CO.: Alishan Township, Tatungshan, ca. 1,700 m, 6 Sept. 1993, C. M. Wang 222 (TNM S12005); Alishan, inter Taroyen et Heishana, 26 April 1912, B. Hayata s.n. (TI).

Etymology. The specific epithet ^ccallosa' refers to the

Actinidia callosa var. formosana Finet & Gagnep., Bull.

Soc. Bot. France, Mem. 4: 20. 1905.

Dunn (1911) referred to the variety, A. callosa var formosana Finet & Gagnep., in Taiwan but the original description of A. callosa var. formosana in Bulletin de la Societe botanique de France is not available. This entity is excluded in this treatment for lack of the protologue and type specimen.

Acknowledgments. The authors are greatly indebted to Yan Liu for information on Actinidia specimens in the Herbarium, Guangxi Institute of Botany, China; Hiroshi Ikeda and Akiko Shimizu for providing the type images of Actinidia in the Herbarium, University Museum, University of Tokyo; Chung-Bow Lee, Sen-Wei Huang, Mark C. K. Yang, Wen-Han Huang, Hong-Dr Isaac Wu, Tsung-I Lin, Ying-Lin Hsu, Tsung-Jen Shen, and Guo-Qing Zhang for valuable comments on statistics; and Teng-Chi Tsai and Yen-Lin Hsueh for accompanying us in the field. Re國 gretfully, my mentor Cheng-Chu Nee passed away before the publication of this paper. He had tirelessy dedicated his life to Actinidia research in Taiwan. We dedicate this article to his memory. This study was supported in part by grants from the Research Center for Biodiversity, Aca-demia Sinica (Taipei, Taiwan) to Ching-I Peng (HAST) and the Division of Silviculture, Taiwan Forestry Research Institute (Taipei, Taiwan) to Ching-Te Chien. We also thank Chung-Han Tsai, Tsan-Piao Lin, and several anonymous reviewers for improving the manuscript.

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357

台灣原生獼猴挑數值分類與分類定模之研究

謝東佑¹ 古訓銘² 簡慶德³ 劉雲聰⁴

¹國立中興大學園藝系

²中央研究院生物多樣性研究中心植物標本館(HAST)

³行政院農業委員會林業試驗所育林組
⁴行政院農業委員會苗栗區農業改良場

獼猴桃爲世界重要原生果樹之一，然而其形態性狀極爲複雜，長期以來造成分類、命名混亂與鑑
定上的困難。本硏究使用60個形態性狀，調查72個包含成熟雌、雄株在內的台灣野生獼猴桃族群，作
爲分類運算單元。以高爾相似性係數計算族群間相似度，進行親緣分析，並將熱量圖排序後，得出五個
明顯的類群斑塊，並發現介於台灣羊桃與硬齒獼猴桃之間的駝齒獼猴桃與腺齒獼猴桃漸滲雜交族群。依
此結果進行指示反應矩陣編碼，再進行各種分類模型的性狀選擇與定模，以邏輯式迴歸模型作爲二類分
類模型，貝氏判別模型作爲多類分類模型，並用這些分類模型對國內外各標本館的重要獼猴桃標本進行
結合性判別分析'以確認各分類群學名使用之適確性。分析結果顯示，台灣共有四種一變種之原生獼猴
桃，因此將台灣植物誌第二版中記載的硬齒獼猴桃(Actinidia callosa)訂正爲腺齒獼猴桃(A.rufa)，駝
齒獼猴桃(A. callosa var. ephppioidea )倂入腺齒獼猴桃中，而紅莖獼猴桃(A. rubricaulis)則訂正爲異
色獼猴桃（A. callosa var. discolor)，台灣羊桃（A. chinensis var. setosa)提升爲台灣特有種；另台灣產四
萼獼猴桃（A. tetramera)之紀錄，則爲軟棗獼猴桃（A. arguta )鑑定之誤，應予以更正。

關鍵詞：貝氏判別分析；台灣植物誌；熱量圖；邏輯式迴歸分類法；漸滲雜交；親緣分析。



HSIEH et al. ― Classifier modeling and numerical taxonomy of Actinidia in Taiwan		349

Etymology. The specific epithet arguta means sharp teeth" in reference to the leaf blade margin. Distribution and habitat. Siberia, Japan, Korea, China, and Taiwan. In Taiwan, occurs in mountain forests and along streams of the northern and central parts at 1,300-	2,600 m. 4. Actinidia setosa (H. L. Li) C. F. Liang & A. R. Ferguson, Guihaia 5 (2): 72. 1985. Actinidia chinensis auct. non Planchon: Susuki in Masamune, Short Fl. Form.



Figure 5. Actinidia rufa (Sieb. & Zucc.) Planch. ex Miquel. A, Habit (Yangmingshan, Taipei County; Callo-05); B, Flowering branch; C, Fruiting branch; D, Male flower; E, Female flower; F, Different shapes of leaves; G, Domatia in axils of lateral veins; H, Fruit; I, Pith of one-year old shoot.