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Based on Molecular Phylogenetic Evidence

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Based on Molecular Phylogenetic Evidence J. Jpn. Bot. 91(6): 337–344 (2016) Two New Combinations of Leptopetalum (Rubiaceae) Based on Molecular Phylogenetic Evidence a, b b b Akiyo NAIKI *, Yoshiteru KOMAKI , Keiko MIZUNASHI and Tetsuo OHI-TOMA aIriomote Station, Tropical Biosphere Research Center, University of the Ryukyus, 870, Uehara, Yaeyama-gun, Okinawa 907-1541 JAPAN; bBotanical Gardens, Graduate School of Science, The University of Tokyo, 3-7-1, Hakusan, Bunkyo-ku, Tokyo 112-0001 JAPAN *Corresponding author: [email protected] (Accepted on October 11, 2016) As a result of our phylogenetic analysis, two new combinations, Leptopetalum pachyphyllum (Tuyama) Naiki & Ohi-Toma, and L. strigulosum (Bartl. ex DC.) Neupane & N. Wikstr. var. luxurians (Hatus.) Naiki & Ohi-Toma, are proposed for taxa formerly in Hedyotis s.l., which was recently divided into 13 genera recircumscribed on the basis of the phylogenetic and morphological evidences. Leptopetalum pachyphyllum is closely related to L. mexicanum Hook. & Arn., the type of the genus, and L. grayi (Hook. f.) Hatus., shrub species endemic to the Ogasawara Islands, Japan. Leptopetalum strigulosa var. luxurians is nested in the clade of L. strigulosum. Key words: Bonin Islands, Hedyotis, Kazan Islands, Leptopetalum pachyphyllum, Leptopetalum strigulosum var. luxurians, Ogasawara Islands, Rubiaceae, Volcano Islands. Hedyotis L. s.l. (including Oldenlandia: W. H. Lewis, Oldenlandia L., and Scleromitrion Rubiaceae) comprises 500–600 species of (Wight & Arn.) Meisn. herbs, subshrubs and shrubs from tropical, The genus Leptopetalum was described in subtropical to warm temperate regions in the 1838 by W. J. Hooker and G. A. Walker-Arnott world (Neupane et al. 2015). Since the genus based on L. mexicanum Hook. & Arn. (Hooker is a highly heterogeneous assembly, many and Walker-Arnott 1830–41). The original separate genera have been recognized. Neupane material was collected from the Ogasawara et al. (2015) accepted the following 13 genera (Bonin) Islands, Japan, but mistakenly supposed based on previous, and their own, detailed to have been collected from Mexico (Fosberg morphological and/or phylogenetic analyses and Sachet 1991, Ohba 2003). For many (Terrell et al. 2005, Groeninckx et al. 2010, years, Leptopetalum was treated as a subgroup Guo et al. 2013, Wikström et al. 2013): Debia of Hedyotis or Oldenlandia, cf. Hedyotis Neupane & N. Wikstr., Dentella J. R. Forst. [unranked] Leptopetalum (Hook. & Arn.) Hook. & G. Forst., Dimetia (Wight & Arn.) Meisn., f., Hedyotis subgenus Leptopetalum (Hook. Edrastima Raf., Exallage Bremek., Hedyotis & Arn.) Fosberg & Sachet, or Oldenlandia s.str., Involucrella (Benth. & Hook. f.) Neupane, sect. Leptopetalum (Hook. & Arn.) K. Schum., Kadua Cham. & Schltdl., Kohautia Cham. & although some Japanese botanists published Schltdl., Leptopetalum Hook. & Arn., Neanotis new species and new combinations under —337— 338 植物研究雑誌 第 91 巻 第 6 号 2016 年 12 月 Leptopetalum (Hatusima 1936, Hosokawa (Fig. 1)). Regardless of the mistake of the 1938). In Flora of Japan, Yamazaki (1993) citation, the publication by Fosberg and Sachet published Hedyotis subgenus Leptopetalum (1991) can be regarded as a valid one because (Hook. & Arn.) T. Yamaz. as a new rank, but the they designated one of the two ‘type specimens’ name was a later isonym of Hedyotis subgenus by Tuyama (1968) as the type. Leptopetalum (Hook. & Arn.) Fosberg & Sachet. Although Hedyotis pachyphylla is thought Recent phylogenetic studies indicated that to be closely related to the shrub species in two species of Leptopetalum (L. foetidum (G. the Mariana Islands rather than Leptopetalum Forst.) Neupane & N. Wikstr. and L. mexicanum) grayi and L. mexicanum, which are endemic formed a distinct clade together with three small to the Ogasawara (Bonin) Islands (Tuyama annual or perennial herbs of Thecagonum Babu: 1968, Ono and Kobayashi 1985, Fosberg and T. biflorum (L.) Babu, T. pteritum (Blume) Babu Sachet 1991), its phylogenetic relationship and T. strigulosum (Bartl. ex DC.) Terrell & H. has not been investigated. In the present study, Rob, all of which were once treated as species of we investigated the phylogenetic position Hedyotis and Oldenlandia (Wikström et al. 2013, of H. pachyphylla based on the nucleotide Neupane et al. 2015). Neupane et al. (2015) sequences of nuclear ribosomal ITS regions, and recircumscribed Leptopetalum that comprises chloroplast rps16 intron and petB–petD spacer. L. biflorum (L.) Neupane & N. Wikstr., L. Consequently, Leptopetalum pachyphyllum foetidum, L. pteritum (Blume) Neupane & N. (Tuyama ex Fosberg & Sachet) Naiki & Ohi- Wikstr., L. strigulosum (Bartl. ex DC.) Neupane Toma, a new combination, is here published. & N. Wikstr., L. mexicanum Hook. & Arn. (= Leptopetalum strigulosum (Bartl. ex DC.) H. hookeri (K. Schum.) Fosberg), and L. grayi Neupane & N. Wikstr. var. luxurians (Hatus.) (Hook. f.) Hatus., based on their morphological Naiki & Ohi-Toma is also published as a new and phylogenetic evidences. Leptopetalum is combination because the variety was not treated characterized by being glabrous throughout in the newly recircumscribed genus by Neupane plants and the ovoid or obtusely angulate seeds et al. (2015). appearing pitted due to shallow depressions bordered by thick and sinulate walls (Neupane et Materials and Methods al. 2015). Nucleotide sequences of ITS, rps16 intron, Hedyotis pachyphylla was published in and petB–petD spacer were used for the 1968 by Tuyama from the Volcano (Kazan) present phylogenetic analysis. In addition to Islands, Japan (Tuyama 1968). This publication, Leptopetalum samples with which the three however, was invalid because two specimens regions were analyzed in Guo et al. (2013), were cited as types (‘H. Huzikawa, Aug. 20, Wikström et al. (2013), and Neupane et al. 1930–typus fructus in TI’ and ‘M. Okabe, Oct. (2015), nucleotide sequences of Japanese 22, 1935–typus floris in TI’), contrary to the Leptopetalum and H. pachyphylla were newly Art. 40.1 and 40.2 of the Melbourne Code determined (Appendix 1). Total genomic DNA (McNeill et al. 2012). The name was validly was extracted from silica-gel-dried leaf tissue, published when Fosberg and Sachet (1991: 204) using the modified HEPES-CTAB method designated ‘Okabe in 1935 (TI)’ as the lectotype described by Ohi-Toma et al. (2010). Nucleotide with a direct reference to Tuyama (1968). They sequences were determined using polymerase cited the collection locality as ‘Iwoto’ (Sulphur chain reaction (PCR) and direct bi-directional Island) whereas ‘Okabe in 1935 (TI)’ was sequencing. PCR amplification was conducted actually collected from ‘Kita-Iwoto’ (as ‘San using TaKaRa Ex Taq (TaKaRa Bio, Shiga, Alessandro Island’ on the label of the specimen Japan), at the following cycling conditions: December 2016 The Journal of Japanese Botany Vol. 91 No. 6 339 Fig. 1. Holotype of Hedyotis pachyphylla Tuyama ex Fosberg & Sachet (M. Okabe s.n., TI00044065). denaturation at 96°C for 45 sec, followed by 33 and PIpetB1365F and PIpetD738R for petB– cycles at 96°C for 45 sec, annealing at 50°C for petD (Löhne and Borsch 2005). Amplification 45 sec, extension at 72°C for 1 min, and a final products were purified using illustra ExoProStar extension at 72°C for 10 min. For amplifying (GE Healthcare UK Ltd., Buckinghamshire, and sequencing, we used the following published England, UK). The purified PCR fragments were universal primers: ITS5 and ITS4 (White et al. amplified using the Big Dye Terminator v3.1 1990), F and 2R for rpl16 (Oxelman et al. 1997), Cycle Sequencing Kit (Applied Biosystems, 340 植物研究雑誌 第 91 巻 第 6 号 2016 年 12 月 Fig. 2. Strict consensus tree of 180 most parsimonious trees of Leptopetalum, based on the combined data of nuclear ribosomal ITS regions, and chloroplast rps16 intron and petB– petD spacer. Values above the nodes represent the number of nucleotide changes, and values below the nodes represent bootstrap support. Foster City, California, USA). Cycle-sequencing basis of the molecular phylogenetic study of reactions were performed using the same Neupane et al. (2015) (Appendix 1). primers for the PCR amplification. Sequencing The nucleotide sequences, including was performed using Applied Biosystems® sequences from GenBank, were manually 3130 Genetic Analyzer. All newly generated aligned and gaps, except the length nucleotide sequences were deposited in the polymorphisms caused by mononucleotide DNA Data Bank of Japan (DDBJ). Two Kadua repeats, were binary coded as a fifth character. species were selected as outgroup taxa on the The incongruence length difference (ILD) December 2016 The Journal of Japanese Botany Vol. 91 No. 6 341 among ITS, rps16, and petB–petD was tested clades were recognized in Leptopetalum, using a partition-homogeneity test in PAUP* although a few branches were weakly supported. v.4.0a150 (Swofford 2002). A three-way test The clade of L. pteritum in Southeast Asia (BP was run using a heuristic search of 10,000 98%) was the first to branch off in the genus. replicates, with 10 random stepwise additions, Leptopetalum biflorum was not monophyletic tree bisection and reconnection (TBR) branch and was divided into two clades, 1) Nepal and swapping, MulTrees in effect, and no limit of China (BP 47%) and 2) Singapore and Ryukyu MaxTrees limit. Since the test indicated that the in Japan (BP 100%). The shrub species in the three regions were not significantly incongruent, Volcano Islands, Hedyotis pachyphylla (shown their combined sequences were used as a final as Leptopetalum pachyphyllum in Fig. 2), formed
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