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Phylogenetic Analysis of the Family Hypnaceae Based on Rbel Gene Sequences

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Phylogenetic Analysis of the Family Hypnaceae Based on Rbel Gene Sequences T. Arikawa, H. Tsubota, H. Deguchi, N. Nishimura & M. Higuchi (2008). Phylogenetic Analysis of the Family Hypnaceae Based on rbeL Gene Sequences. In: Mohamed 1-1, Baki BB, Nasrulhaq-Boyee A, Lee PKY, cds. Bryology in the New Millenniuln. Kuala Lumpur: University of Malaya, pp. 215-225. PHYLOGENETIC ANALYSIS OF THE FAMILY HYPNACEAE BASED ON rbcL GENE SEQUENCES TOMOTSUGU ARIKAWAI*, HIROMI TSUBOTA2, HIRONORI DEGUCHI3, NAOKI NISHIMURM AND MASANOBU HIGUCHI5 1 Department of Biology, Keio University, 4-1-1 Hiyoshi, Kohoku-ku, Yokohama 223-8521 Current address: Tottori Prefectural Museum, 2-124 Higashi-machi, Tottori 680-0011 2 Miyajima Natural Botanical Garden, Graduate School of Science, Hiroshima University, 1156-2 Mitsumaruko-yama, Miyajima-cho, Hatsukaichi 739-0543 3 Department of Biological Science, Graduate School of Science, Hiroshima University, 1-3-1 Kagamiyama, Higashi-hiroshima-shi 739-8526 4 The Botanical Garden, Okayama University of Science, 1-1 Ridai-cho, Okayama 700-0005 5 Department of Botany, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba 305-0005, Japan * Corresponding author: [email protected] SUMMARY The molecular phylogenetic analysis of 365 OIUs was carried out using nucleotide sequences of the chloroplast gene rbeL, focusing on the most diversified moss family Hypnaceae. Nucleotide sequences of rbeL were newly determined in 71 samples. Neither the family Hypnaceae nor the type genus Hypnum came together in a monophyletic group. Most of the relationships in the clade of hypnalean mosses corresponding especially to families remain obscure. However, some new findings shown in the following were obtained: (1) Hypmmz plumaeforme, H. sakuraii, and H. oldhamii make a phylogenetic group with Eetropotheeiun1 spp. and Vesicularia spp., (2) Gollania taxiphylloides makes a monophyletic clade with Taxiphyllum spp. and Hondaella braehytheeiella, and G. splendens makes a clade with HOlnomallium japonico­ adnatum, while other species of Gollrmia come together in a monophyletic group. KEYWORDS: Hypnaceae, Pylaisiaceae, Hypnun1, Stereodon, Gollania INTRODUCTION Since Schimper (1856) established the (e.g., Andrews, 1954; Seki, 1969; family Hypnaceae, some characteristic Noguchi, 1972; Buck, 1980; Nishimura et members have been isolated from the al.,1984). Recently described hypnalean Hypnaceae and grouped into new genera with double costa and smooth families. Nishimura (1999) reviewed leaf laminal cells were placed in the the history of the Hypnaceae. Several Hypnaceae; Horridohypnum (Buck, 1981), authors who revised neighboring Andoa (Ochyra, 1982), Mahua (Buck, families have transferred many genera 1983), Caribaeohypnum (Ando & Higuchi, to the Hypnaceae without due discus ions 1984), Irelandia (Buck, 1984), Podperaea 215 Bryology in the New Millennium (Iwatsuki & Glime, 1984), Elharveya MATERIALS AND METHODS (Crum, 1986), Pseudotaxiphyllurn This study consists of two steps: (1) (Iwastsuki, 1987), Taxiphyllopsis (Higuchi obtaining sequence data (DNA & Deguchi, 1987), Fallaciella (Crum, extraction, peR amplification, DNA 1991), etc. The Hypnaceae has thus been sequencing, and download from DNA enlarged and heterogeneous, comprising database), and (2) data analysis more than 60 genera and 1000 species. (construction of phylogenetic trees). A Although the Hypnaceae have been total of 71 hypnalean samples and one variously circumscribed (e.g., Nishimura sample of Racopilum aristatum Mitt. et al., 1984), they remain as a rather were obtained from the fields, and the heterogeneous group. Hedenas (1989) voucher specimens are depOSited in the mentioned that the Hypnaceae is a kind herbaria of the National Museum of of catchall for genera taxonomically Nature and Science, Japan (TNS) or ill defined. Hiroshlraa University (HIRO). Methods Delimitation of families of of obtaining rbcL sequence data were hypnalean mosses and their arrangement essentiaU y the same as our previous iT: phylogenetic sequence are extremely studies (Arikawa & Higuchi, 1999, 2002, challenging. These families often 2003j Tsubota et al., 1999, 2000, 2001a, b, received moderate support by molecular 2002; Arikawa et al., 2006). In addition phylogenetic analyses, but inferences to the newly obtained sequences, 288 regarding their mutual relationships are rbcL sequences of hypnalean species and rather weak (e.g., Tsubota et aI., 1999, five sequences of Racopilum spp. were 2002a; Buck et al., 2000j Maeda et al., downloaded from the DNA database of 2000; Arikawa & Higuchi, 2002j Goffinet the DDBJ/EMBL/GenBankInternational & Buck, 2004). The Hypnaceae are Nucleotide Sequence Database consistently resolved as a polyphyletic Collaborations. A total of 365 sequences entity from molecular phylogenetic treated in this study were aligned analyses (Arikawa & Higuchi, 1999, 2003; manually. The undetermined sites Tsubota et aI., 1999, 2002a, b; Buck et al., were excluded from the analysis and 2000). Even the type genus Hypnum is determined sites were concatenated shown to be polyphyletic. to a single data matrix. Bootstrap The present paper is the first step probabilities (BPs) were calculated with toward the refinement of the family neighbor-joining (NT) method using Hypnaceae to be a natural group. In Clustal W 1.83 (Thompson et al.I 1994). order to redefine the Hypnaceae, we A maximum-likelihood (ML) tree were need to construct a reliable tree covering searched with the local rearrangement aU genera in the order Hypnales. The method from the neighbor-joining (NJ) ribulose 1,5-bisphosphate carboxylase/ tree generated by the MOLPHY version oxvgenase large subunit gene (rbcL) is a 2.3b3 package (Adachi & Hasegawa, oao~d "molecular marker for constructing 1996; NucML 2.3b3 and NJdist 1.25). phylogenetic tree, because so many sequences are already available. The aim this is to construct a preliminary phylogenetic tree of the Hypnaceae and I"elated faimiIies using a large number of rbcL sequences, to delimit the and to make clear the Voucher among hypnaceous taxa. information 216 Phylogenetic Analysis of Hypnaceae Based on rbeL the sequences were listed in Appendix species commonly included in the genus. A. A total length of 1,161 sites was used This conclusion was supported by all for the following phylogenetic analyses. molecular phylogenetic analyses that Some identical sequences over the 1,161 involved at least several species of sites were integrated into one sequence Hypnum sensu lato (e.g., Goffinet et al., each. A total of 333 sequences among 365 2001; Tsubota et al., 2002a; Gardiner et sequences were then analyzed for al., 2005). The present analysis also constructing a phylogenetic tree. TN93 confirmed this result. Gardiner et al. model was used for the analysis. The (2005) resurrected the family Pylaisiaceae log-likelihood of the tree searched with based on the phylogenetic analysis the local rearrangement method from the deduced from ITS, trnL-trnF region and neighbor-joining (Nn tree was -22478.17 morphological data. The Pylaisiaceae ± 1180.69 (Fig. 1). The nodes supported sensu Gardiner et al. (2005) consists of with both local bootstrap probabilities Homomallium, Pseudohygrohypnum, (LBPs; in %) 70% with ML method or Pylaisia, and Stereodon (the main part of higher and bootstrap probabilities (BPs; Hypnum sensu lata except its type species; in %) calculated with NJ method 50% or see also Ignatov & Ignatova, 2004). The higher are shown as solid lines in Fig. 1; Pylaisiaceae clade is recognizable in the the other nodes are shown as broken present result (Fig. 1). Although the BP lines.. Most of the relationships in the calculated with NJ method is not high clade of hypnalean mosses corresponding (46%), Hypnum plumaeforme, H sakuraii, especially to families remain obscure. As and H. oldhamii make a phylogenetic also shown in the recent studies (e.g., group with Ectropotheciu111 spp. and Arikawa & Higuchi, 1999; Goffinet Vesicularia spp. This group makes a weak et al'f 2001; Tsubota et al., 2002), neither clade with Pylaisia, H01110111allium, the family Hypnaceae nor the type Foreauella, Hypnum fauriei Cardot, and genus Hypnum came together in a H. calcicolum Ando. This clade is monophyletic group. However, some recognizable as the Pylaisiaceae clade. new findings were obtained: (1) Hypnum Gardiner et al. (2005) did not include any plumaefor111e Wilson, H sakuraii (Sakurai) species of Ecrtopothecium, Vesicularia, Ando, and H. oldhamii (Mitt.) A.Jaeger and Foreauella in their analyses. In order & Sauerb. made a phylogenetic group to delimit the ranges of the family with Ectropotheciu111 spp. and Vesicularia Pylaisiaceae and the genera Hypnum and spp., (2) Gollania taxiphylloides Ando & Stereodon respectively, further samplings Higuchi made a monophyletic clade with of additional taxa would be necessary. Taxiphyllum spp. and Hondaella The genus Gollania is one of the brachytheciella (Broth. & Paris) Ando, and members of the Hypnaceae and is most G. splendens (lisiba) Nog. made a clade closely related to Hypnu111 (Higuchi, with Homomallium japonico-adnatum 1985). Most species of Gollania examirled (Broth.) Broth., while other species of came together in a monophyletic group Gollania came together in a monophyletic with strong statistical support (100% LBP group near the clade of Ctenidium and and 98% BP with NJ), while G. the Pylaisiaceae. taxiphylloides made a monophyletic clade with Taxiphyllum spp. and Hondaella DISCUSSION brachytheciella, and G. splendens made a Hedenas (1989) pointed out that clade with Homomalliu111 japonico­ Hypnu111 cupressiforme
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