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A Morphometric Analysis of Infraspecific Taxa Within the Ixeris Chinensis Complex (Asteraceae, Lactuceae)

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A Morphometric Analysis of Infraspecific Taxa Within the Ixeris Chinensis Complex (Asteraceae, Lactuceae) WhangBot. Bull. et Acad.al. — Sin.A morphometric (2002) 43: 131-138 analysis within the Ixeris chinensis complex 131 A morphometric analysis of infraspecific taxa within the Ixeris chinensis complex (Asteraceae, Lactuceae) Sung Soo Whang1, Kyung Choi2,5, Robert S. Hill3, and Jae-Hong Pak4,* 1Division of Science Education, Chonbuk National University, Chonju 561-756, Republic of Korea 2Department of Biology, Yeungnam University, Keongbuk 712-749, Republic of Korea 3Department of Environmental Biology, University of Adelaide, SA 5005, Australia 4Department of Biology, Kyungpook National University, Taegu 702-701, Republic of Korea (Received July 17, 2001; Accepted October 3, 2001) Abstract. Morphometric methods were used with 40 morphological characters from each of 110 individuals col- lected from 11 populations of the Ixeris chinensis complex to test hypotheses regarding infraspecific taxa. Individu- als of the nearest relative I. tamagawaensis were included as an outgroup. To utilize structual associations of the original data and the cumulative values of character variation, a modified data set was generated by matching factor scores with the large original data set. As a result, the original data was reduced to about 10% of its original size. The modified data also emphasized the cumulative values of character variations. Structural associations among char- acters measured, however, were not significant for most factors extracted even though some obvious trends appeared. Relationships between cumlative values of variation and taxa in the complex are discussed based on the results of the canonical discriminant function analysis, ANOVA, multiple comparison test, and cluster analysis, using the modified data. This study supports Kitamura’s (1956) classification of infraspecific taxa in the I. chinensis complex. Given that there is no overlap of cumulative values of variation between the species complex and I. tamagawaensis, there is probably a 75% character overlap within the complex. Keywords: Ixeris chinensis complex; Infraspecific taxa; Morphometric analysis. Introduction several morphological characters, especially if other forms of data are not available. The term “species complex” has frequently been applied The Ixeris chinensis complex (Asteraceae, Lactuceae) to flowering plants, but its origin is obscure. It has often is distributed from Siberia to Japan, including Korea, been used to describe species aggregations made up of Taiwan, and China (Pak et al., 1997). This complex has a infraspecific taxa such as microspecies, agamospecies, and basic chromosome number of eight (Pak and Kawano, semispecies (e.g. Fosberg, 1942; Sylvester-Bradley, 1952; 1990), but also contains diploids (2n = 16), triploids (2n = Davis and Heywood, 1963; Northington, 1976; Grant, 1981). 24), and tetraploids (2n = 32) (Ishikawa, 1921; Babcock et Several criteria have been used to establish groups of in- al., 1937; Chuang et al., 1962; Hsu, 1967, 1970; Peng and fraspecific taxa, including their morphological discreteness, Hsu, 1978; Pak and Kawano, 1990) and exhibits sympatry ecological differences, genetic divergence, and natural re- and/or allopatry among the ploidy (e.g. Pak et al., 1995, productive isolation (e.g. Davis and Heywood, 1963; 1997). The diploid plants observed are sexual while trip- Gilmartin, 1974; Grant 1981; Stuessy, 1990). Morphologi- loids and tetraploids are asexual. Many complicated varia- cal discreteness plays a major role in the early stage rec- tions in external morphology have been reported in hybrids ognition of infraspecific taxa. A practical difficulty here between sexual and usually asexual plants, which can re- is that almost all morphological characters and/or charac- vert to sexual reproduction under some conditions (e.g. ter states used for evaluating infraspecific taxa are only Harlan and de Wet, 1963; Yahara, 1983, 1990). Within this slightly differentiated from one another and usually show complex a complicated morphological overlap, without any considerable overlap. Therefore the identification of in- discontinuities, has led to taxonomic difficulty (Kitamura, fraspecific taxa and their hierarchical arrangement within 1956; Pak and Kawano, 1992; Kim et al., 1999). Kitamura a complex of species usually requires a combination of (1956) classified this complex into the three subspecies Ixeris chinensis (Thunb.) Nakai subsp. chinensis, subsp. 5Current address. Division of Life Science & Graduate School strigosa (H. Lév. et Vaniot) Kitamura, and subsp. versi- of Biotechnology, Korea University, Seoul 136-701, Republic color (Fischer) Kitamura. Tzvelev (1964) classified it as of Korea. three different species in a different genus, Ixeridium *Corresponding author. Tel: +82-53-950-5352; Fax: +82-53- chinensis (Thunb.) Tzvelev, I. strigosum (H. Lév. et Vaniot) 953-3066; E-mail: [email protected] Tzvelev and I. gramineum (Fisch.) Tzvelev. Pak and 132 Botanical Bulletin of Academia Sinica, Vol. 43, 2002 Kawano (1992) classified it as two subspecies of one spe- cies and one different species in the same genus; Ixeris chinensis subsp. chinensis and subsp. versicolor and I. strigosa (H. Lév. et Vaniot) Pak et Kawawo. The taxo- nomic resolution of this complex, therefore, remains uncertain. In an attempt to resolve difficulties caused by morpho- logical overlap, multivariate methods have increasingly been applied, especially in species complexes of flower- ing plants (e.g. Sokal and Sneath, 1963; Gilmartin, 1967; Jensen and Eshbaugh, 1976; McNeil, 1984; Jensen et al., 1993). Several multivariate analyses are applicable to this problem, and the adoption of different statistical proce- dures in dealing with such problems is common. The clas- sificatory schemes proposed for the Ixeris chinensis complex based on morphological characters can be tested using multivariate statistical analysis of the huge amount of raw data that can be collected from such a morphologi- cally variable group. As a first step in such a process, a factor analysis capable of regenerating reduced data might Figure 1. Map showing collecting localities of the Ixeris be effective in order to yield a clear output in later chinensis complex and its nearest relative I. tamagawaensis. !=I. treatments. chinensis subsp. chinensis (Mt. Keumo, Janghang-ri and Mokye- ri in Kyungbuk); !=I. chinensis subsp. strigosa (triploid) This study aims to establish a theoretical model for the (Moraejae and Malgoljae in Chonbuk, Damyang-up in categories of infraspecific taxa and to provide taxonomic Chonnam); "=I. chinensis subsp. strigosa (tetraploid) (Is. Nami resolution of the Ixeris chinensis complex based on mor- and Taebong-ri in Kyunggi, Sanchong-up in Kyungnam), #=I. phometric analyses. We will not only discuss categories tamagawaensis (near Tamagawa in Tokyo). at the level of both species and infraspecies, but we will also test the taxonomic schemes previously proposed for this complex. lections and from the following herbaria: KYO, SKK, SNAU, TI, TNS, KNU, Chonnam National University, and Materials and Methods Kangwon National University (Figure 1). Forty characters were scored for the morphometric Specimens of the Ixeris chinensis complex were se- analysis (Table 1). They were measured using a digimatic lected from among 81 populations, chosen on the basis caliper and hand lens. Of these characters, radical leaf of ploidy and geographic distribution (Pak et al., 1997, Fig- blade length (RLL) was measured from three leaves in or- ure 1). Twenty individuals from two populations of I. der of size among those attached at the base; internode Tamagawaensis (Makino) Kitamura were collected near length (INL) was measured at the second node up from Tamagawa, Ozaku, Oome-shi, Tokyo, Japan and were the base; length and width of inner-phyllary (IIL and IIW) added to the analysis because this species is regarded as were measured on three randomly selected leaves; length the nearest relative of I. chinensis (Kitamura, 1956; Pak and and width of the outer-phyllary (OIL and OIW) were mea- Kawano, 1990, 1992, Figure 1). The complex has gener- sured on three replicates; and length and width of the ally been recognized as comprising three taxa although achene (ACL and ACW) were measured on one mature their taxonomic category and nomenclature differs among specimen. researchers (Kim et al., 1999). To begin with, Kitamura’s Factor analysis was used to interpret structural asso- (1956) two subspecies of I. chinensis were recognized, with ciations among the 40 morphological characters, as well subspecies chinensis only being diploid and subspecies as to reduce the large amount of data measured. Factors strigosa being both triploid and tetraploid. Individuals of >1 in eigenvalue were selected and scored. An analysis the triploid and tetraploid populations of I. chinensis of variance of the modified data, generated by matching subsp. strigosa were identified and separated for morpho- the factor score (Âfactor score coefficients of character logical analysis. Kitamura’s (1956) Ixeris chinensis subsp. measurement to factor 1 × standard scores of taxa to versicolar, distinguished within the complex by having a characters) of the original data, was performed to deter- different corolla color and branched roots, is restricted to mine whether significant differences among members of the North Korea, Manchuria, and Russia, and was omitted I. chinensis complex and its nearest relative I. here due to lack of sufficient material for comparison. tamagawaensis exist or not. Multiple comparison
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