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Geographic Distribution of Ploidy Levels and Chloroplast Haplotypes in Japanese Clerodendrum Trichotomum S

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Geographic Distribution of Ploidy Levels and Chloroplast Haplotypes in Japanese Clerodendrum Trichotomum S ISSN 1346-7565 Acta Phytotax. Geobot. 70 (2): 87–102 (2019) doi: 10.18942/apg.201823 Geographic Distribution of Ploidy Levels and Chloroplast Haplotypes in Japanese Clerodendrum trichotomum s. lat. (Lamiaceae) 1,* 2 3 4 5 Leiko Mizusawa , Naoko ishikawa , okihito YaNo , shiNji Fujii aNd Yuji isagi 1Faculty of Human Development and Culture, Fukushima University, 1 Kanayagawa, Fukushima 960-1296, Japan. * [email protected] (author for correspondence); 2Graduate School of Arts and Sciences, The University of Tokyo, 3-8-1 Komaba, Tokyo 153-8902, Japan; 3Faculty of Biosphere-Geosphere Science, Okayama University of Science, 1-1 Ridai-cho, Kita-ku, Okayama 700-0005, Japan; 4Faculty of Human Environments, University of Human Environments, 6-2 Kamisanbonmatsu, Motojuku-cho, Okazaki, Aichi 444-3505, Japan; 5Graduate School of Agriculture, Kyoto University, Kitashirakawa-oiwake-cho, Sakyo-ku, Kyoto 606-8502, Japan Clerodendrum trichotomum s. lat., under which many infraspecific taxa have been recognized, includes both tetraploid and diploid individuals, although chromosome numbers and geographic variation in ploi- dy levels have not been investigated in the Japanese archipelago. The geographic distribution of ploidy levels and chloroplast haplotypes of four Japanese taxa of C. trichotomum s. lat., based on chromosome counts, flow cytometry, and genotyping of five microsatellite loci is reported. It was determined that Japanese C. trichotomum var. trichotomum and var. yakusimense are tetraploid (2n = 104), while var. es- culentum and C. izuinsulare are diploid (2n = 52). The diploid taxa are distributed only on the southern edge of the Japanese archipelago, while tetraploid C. trichotomum is distributed widely. Such distribu- tion patterns may be formed by temperate forest shrinkage during, and tetraploid expansion after, glacial periods. Thirteen haplotypes were detected, and were divided into the following three clades: (1) Japa- nese C. trichotomum var. trichotomum and C. izuinsulare, (2) C. trichotomum var. yakusimense and var. esculentum, and (3) Chinese C. trichotomum. Two haplotypes were shared between diploid and tetra- ploid lineages, suggesting multiple polyploidization events in C. trichotomum s. lat.. Inconsistency be- tween nuclear and chloroplast phylogenetic trees suggests a past inter-lineage hybridization event in C. trichotomum s. lat. Key words: chloroplast phylogeny, Clerodendrum izuinsulare, Clerodendrum trichotomum, Cleroden- drum trichotomum var. esculentum, Clerodendrum trichotomum var. fargesii, Clerodendrum trichoto- mum var. yakusimense, diploid, polyploid, Sino-Japanese region, tetraploid Clerodendrum trichotomum Thunb. s. lat. is a Ohwi with glabrous and thick leaves; var. escul- deciduous pioneer shrub that grows to 5 m tall. It entum Makino with acute apex and cordate base is common in disturbed secondary forests in both of the leaves, and var. izuinsulare (K. Inoue, M. cool temperate and warm temperate climates and Haseg. et S. Kobay.) H. Ohba et S. Akiyama with is widely distributed in the Sino-Japanese region: short style and glabrous leaves (Ohwi 1953, Walk- southern China, Korean Peninsula, Taiwan, and er 1976, Yamazaki 1993, Ohba & Akiyama 2002). the Japanese archipelago (Yamazaki 1993). It is Although Ohba & Akiyama (2002) proposed highly variable in morphology and many infra- treating populations on the Izu islands as C. tri- specific taxa have been recognized (Ohwi 1953, chotomum var. izuinsulare, here we treat them as Yamazaki 1993, Zhuang 1999). In the Japanese C. izuinsulare Inoue, Haseg. et Kobay. following archipelago, at least four infraspecific taxa were Inoue et al. (1997). The diagnostic characteristics recognized: var. trichotomum with pubescent of the infraspecific taxa are variable and appear leaves and long style; var. yakusimense (Nakai) to be continuous between the taxa (Mizusawa & 88 Acta Phytotax. Geobot. Vol. 70 Miyajima unpublished), and have caused confu- the findings with the chloroplast genetic structure sion in their identity in local floras (Hatushima in the wider Japanese archipelago to better under- 1975, Yamazaki 1993, Shimabuku 1997, Shiroka- stand the morphological polymorphism and di- wa 2001, Miyata 2003, Fujii 2006). versification process in C. trichotomum s. lat., a Polyploidization is known to be a key factor highly variable species that includes many infra- that contributes to plant speciation and differen- specific taxa. tiation (Ramsey & Schemske 1998, Soltis et al. 2004). Previous studies of the chromosomes of Clerodendrum trichotomum s. lat. have reported Materials and Methods that C. trichotomum s. lat. includes diploid and tetraploid individuals, although those reports Target taxa and sample identification have been restricted to studies in China and from In this study we included Clerodendrum tri- some locations in Japan. Diploids (2n = 52) are chotomum Thunb. var. trichotomum, C. trichoto- distributed widely in China (Zeng et al. 2011), mum var. yakusimense (Nakai) Ohwi, C. trichot- while Bowden (1940, 1945) reported 2n = 92 for omum var. esculentum Makino, and C. izuinsu- two individuals of C. trichotomum from Japan lare Inoue, Haseg. et Kobay. The geographic dis- and cultivated in Brooklyn and Chapel Hill, USA. tribution of each variety is summarized in Fig. 1 During microsatellite marker isolation, we (Makino 1917, Nakai 1924, Hatushima 1975, generated data that suggested C. izuinsulare is Ohwi 1953, Yamazaki 1993, Inoue et al. 1997, diploid whereas var. trichotomum is tetraploid, at Shimabuku 1997, Shirokawa 2001). In southern least in the Izu islands (Mizusawa et al. 2011). We Kyushu and the Nansei islands, varieties trichot- genotyped 36 and 42 individuals of var. trichoto- omum, esculentum, and yakusimense may be mum and C. izuinsulare, respectively, for 19 mic- sympatric. Although Walker (1976) treated var. rosatellite loci and detected three or four alleles yakusimense as a synonym of C. trichotomum in some loci in each individual of var. trichoto- var. fargesii (Dode) Rehder, var. fargesii is re- mum, but only one or two in each individual of C. ported to occur locally in the mountains of west- izuinsulare (Mizusawa et al. 2011). The microsat- ern China (Sargent 1917). Here, we focus on the ellite genotype of var. yakusimense, a naturalized populations in southern Kyushu to the Nansei is- population in the Izu islands, also suggested that lands, which we refer to as var. yakusimense, fol- it was a tetraploid (Mizusawa 2018). These previ- lowing Ohwi (1953). The distribution of C. tri- ous studies suggest that geographic variation in chotomum s. lat. has not been sufficiently investi- the ploidy levels of C. trichotomum s. lat. corre- gated in the southern portion of the region and on late with the infraspecific taxa recognized. How- Taiwan. ever, ploidy levels have not been considered in re- Identification of the samples was based on the gard to the infraspecific taxa of C. trichotomum following diagnostic features. Variety trichoto- s. lat. mum has pubescent leaves and style protruding Ploidy and DNA variation is sometimes effec- ca. 40 mm beyond the pink corolla tube with tive at elucidating the taxonomy of infraspecific white petal. Clerodendrum var. yakusimense has taxa. For example, a combined approach of ploidy flowers similar to those of var. trichotomum, but and DNA variation improved the understanding can be identified by the thick, glabrous leaves. of infraspecific taxa in Aucuba, Damnacanthus, Clerodendrum var. esculentum has pubescent and Cayratia in the Sino-Japanese region, as well leaves with acute apex and cordate base, some- as in Clerodendrum trichotomum s. lat. (Ohi et what shorter inflorescences, and slender and acu- al. 2003, Naiki & Nagamasu 2004, Ishikawa et minate sepals (Makino 1917), but its floral mor- al. 2014). phology has not been described in detail. In Shi- Here, we report the ploidy levels of the Japa- koku, Kyushu, and the Nansei islands, we identi- nese taxa of C. trichotomum s. lat. and correlate fied potentially sympatric populations of var. tri- June 2019 Mizusawa & aL.– Distribution of Ploidy and Haplotypes in Clerodendrum trichotomum s.l. 89 chotomum and var. esculentum based on the larg- of staining, the fluorescence intensity of each cell est leaf on a shoot. Clerodendrum izuinsulare can was measured using a BD Accuri C6 Flow Cy- be easily distinguished from the other taxa by its tometer (BD Biosciences, CA, USA). In the histo- glabrous leaves, short style and white corolla tube gram of the fluorescence intensity, only the peak (Inoue et al. 1997). region data was used to calculate the mean fluo- Voucher specimens have been deposited in rescence intensity. one of the following herbaria: Kyoto University (KYO), Osaka Museum of Natural History Estimation of geographic distribution of ploidy (OSA), Tokyo Metropolitan University (MAK), levels in C. trichotomum s. lat. and Faculty of Symbiotic Systems Science, Fuku- The geographic distribution of ploidy levels in shima University (FKSE) (Tables 1, 2). Japanese C. trichotomum s. lat. was estimated us- ing five microsatellite markers (ct028, ct041, Determination of ploidy level of each taxon ci111, ci141, and ci144; Mizusawa et al. 2011) The ploidy level of each taxon was estimated based on the methods of Yagi et al. (2009) and by counting the chromosomes, and/or flow cyto- Vergilino et al. (2009). The microsatellite mark- metric analysis (FCA). The number of individu- ers are highly polymorphic and co-dominant als tested in each assay and the results
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