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T. Arikawa, H. Tsubota, H. Deguchi, N. Nishimura & M. Higuchi (2008). Phylogenetic Analysis of the Family 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 family Hypnaceae. Nucleotide sequences of rbeL were newly determined in 71 samples. Neither the family Hypnaceae nor the type came together in a monophyletic group. Most of the relationships in the clade of hypnalean 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 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 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 (Ochyra, 1982), (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 . 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 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 Hed w., the type adnatum. Gollania taxiphylloides has been species of the genus Hypnum, is not known to be a species endemic to Japan closely related to most of the other long before, but it has puzzled Japanese

217 N I--' 0.01 WbSlitutions/site (J:) ~'--'~-AYS53t)74Racopilulll-Cllbpidig~rum .. . AY5244-.'9R""piicm,,",pid,g,.mm IRacopllum ' (out group) .----~_,_~ -f lOOllCu AY853973R~,coplillm-ferril'.1 >, . _ ,\/lJJ229/Racnp,lflm-flr/\11IIwn ,c(](,C'-, ~Yk53?75Br:'lth\:'aite,l.-:,ukAa AY)24441Dra\lhwi\lti;:Hll!cata DQIt.J6098Plychomniella-plychoc

I AYh311f'i1Glyphmhedum-~Litlr()i DQ1%07RDichelo:lontium-nitidu Hypnanae ~, 1!67 9{f92 , ---I , , DQI960RSGarovaglia-plicata DQ196090Garovagha , -zanterniI , I... ___ ~, Ptychomniieae , ,

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1---'~---lr_-_-_-_-_-_"_~'_--_-:!.f}~!,:I1~;2,,===:=jAY6311H5l-AY3~~~~~~:~:~~~~~~~;~U,m-sutinamell DQ467871Crmsonutnmn paln,l OJ 1-1 ~i287il'~k"i'h'Mii.",i" Hookert'ales ~-~I ~ AAI03.bIBenlhllama-ehmhata '-< : -1L______-""= _____ -j~Bl()3353IAY631!75~~(~~~~~!~~~~~~I~~4AddothEdum~bogotenso : %.,~q....--- ABJ.1227imichclyma-jrrponicllm , AY8539S2Bryobrolhcra-crl:"TIulal oqo c------t_ r=====AJ2;~~~~~~2~i:;;~~i~~~;:~~~~~~yreti Fontlnalaceae '-< ~. Cj --- ~)!:------l sum AF232695Hypopluygium-tnmans s- I ID ]~-_:r-~---- DQ46310S~:~~~~!~~nl~~I~~~~rgia_m~d'1ga",a I I jr,(_l DQ,t67S74f

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1 I)\.l4bt:)71iI_~HC(jdun-'Cl[J[Oldes , b... __ ~ AFX\]fJ75Le'Jwdun-Jlllac.eu, Portion II Portion I Fig. 1, The h'ee searched with the local rearrangement method from the neighbor-joining (NJ) tree of the aligned rbcL gene sequences (1,161 bp; TN93 model; 2a/~= 5.969; In L = -22478J7 ± 11.80,69 by NucML), The horizontal length of each branch is proportional to the estimated number of nucleotide substitutions, The root is arbih-arily placed on the branch leading to Racopilul11spp. and Braithwaitea 8pp. The numbers above the nodes are the local bootstrap probabilities (LBPs; in %) and bootstrap probabilities calculated with NJ method (BPs; in %). LBPs 70% or higher and BPs 50% or higher are surely shown, The nodes supported with both LBPs 70% or higher and BPs 50% or higher are shown as solid lines; the other nodes are shown as broken lines, Sequences newly determined in this study are in Bold-italic, Phylogenetic Analysis of Hypnaceae Based on rbeL Portion, I , 0.01 substitutions/site , , ,

, , _Jl..l2.L _: , , j----,~w· I , , ' ,

, , L~_:

: : I",J!!:. L--i , L_J , w-: , , ' L~

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, Portion II Portion III

219 Bryology in the New Millennium

Portion II

0.01 substitutions/site

~, , , ,

1~2i.! ;-1 ' I..,IW---,,t IGollania

Pylaisiaceae Portion III

ICampyliaceae

ITaxiphyllum

, , u 7SI-:

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Neckeraceae

IHylocomiaceae

220 Phylogenetic Analysis of Hypnaceae Based on rbcL bryologists for the solution of its generic Yoshida, for providing materials. We are identity since it has been found only in also grateful to Ms. Y. Hirayama sterile condition and has several (National Museum of Nature and characters overlapping with these allied Science, Japan) for strenuous laboratory genera: Gollania, Hypnum, and work for obtaining sequence data. Taxiphyllum (Ando & Higuchi, 1981). The of G. splendens also have REFERENCES not been found yet either. Further Adachi J, Hasegawa M. 1996. Computer morphological studies and taxonomic Science Monographs, 28. MOLPHY treatments are needed for stabilizing version 2.3. Programs for molecular their taxonomic positions. phylogenetics based on maximum Subgeneric taxa have been proposed likelihood. Tokyo: The Institute of in the genus Hypnum (e.g., Ando, 1973). Statistical Mathematics. Although all subgeneric taxa were Ando H. 1973. Studies on the genus Hypnum included in this study, their segregation Hedw. (II). Journal of Science of the Hiroshima University. Series B, Div. 2 is not supported and the relationship 14: 165-207. remained obscure. Ando H, Higuchi M. 1981. Gollania Buck (2007) pointed out that a taxiphylloides Ando et Higuchi, sp. nov. combination of multiple genes with and Gollania japonica (Card.) Ando et carefully observed morphology should Higuchi, comb. nov. (Hypnaceae, provide us with a stable classification. Musci) from Japan. Hikobia, Supplement He also mentioned, "to date, though, the 1: 189-195. largest group among the mosses lacking Ando H, Higuchi M. 1984. Caribaeohypnum, phylogenetic resolution is the Hypnales." a new genus for a northern Latin Not only exhaustive taxon sampling American mosses, Hypnum polypterum but also application of more suitable (Mitt.) Broth. Cryptogamie, Bryologie, Lichenologie 5: 7-14. genes than rbcL would be needed for Andrews AL. 1954. Taxonomic notes. XII. constructing a phylogenetic tree with The families Rhytidiaceae and higher resolution. In order to . The Bryologist 57: circumscribe the family Hypnaceae, 251-26l. further molecular phylogenetic analysis Arikawa T, Higuchi M. 1999. Phylogenetic based on multiple genes and careful analysis of the Plagiotheciaceae reexaminations of morphological (Musci) and its relatives based on rbcL characters are needed. gene sequences. Cryptogamie, Bryologie 20: 231-245. ACKNOWLEDGEMENT Arikawa T, Higuchi M. 2002. Phylogenetic position of the genera lsopterygiopsis This study was supported by a Grant­ and Herzogiella (Musci) based on rbcL in-Aid from the Ministry of Education, gene sequences. Bryological Research 8: Culture, Sports, Science and Technology, 137-147. Japan (no. 18570097 to Higuchi, Arikawa T, Higuchi M. 2003. Preliminary "Phylogenetic and systematic study to phylogenetic analysis of Pylaisia redefine the family Hypnaceae") and the (Hypnaceae, Musci) and its relatives General Project of National Museum of based on rbcL gene sequences. Journal Nature and Science, Japan (Integration of the Hattori Botanical Labara tory 94: of molecular phylogeny and 87-106. Arikawa T, Tsubota H, Higuchi M. 2006. classification for all organisms). Our A reappraisal of Pylaisiopsis gratitude is due to Ms. M. Chishiki, Mr. (). The Bryologist 109: K. Kawai, Mr. K. M. Kellman, Mr. H. 381-390. Kiguchi, Mr. Y. Tateishi, and Dr. T.

221 Bryology in the New Millennium

Buck WR. 1980. A generic revision of the Higuchi M. 1985. A taxonomic revision of . Journal of the Hattori the genus Gollania Broth. (Musci). Botanical Laboratory 48: 71-159. Journal of the Hattori Botanical Laboratory Buck WR. 1981. Animadversions on 59: 1-77. Pterigynandrum with special Higuchi M, Deguchi H. 1987. Taxiphyllopsis, commentary on Forsstroemia and a new genus of the Hypnaceae from Leptopterigynandrum. The Bryologist Japan. Memoirs of the New York Botanical 83: 451-465. Garden 45: 528-532. Buck WR. 1983. A synopsis of the Ignatov MS, Ignatova EA. 2004. Moss Flora South American taxa of Fabronia of the Middle European Russia, vol. 2. (Fabroniaceae). Brittonia 35: 248-254. Moscow: KMK (In Russian) Buck WR. 1984. Taxonomic and nomen­ Iwatsuki Z. 1987. Notes on Isopterygium Mitt. clatural notes on West Indian (Plagiotheciaceae). Journal of the Hattori Hypnaceae. Brittonia 36: 178-183. Botanical Laboratory 63: 445-451. Buck WR. 2007. The history of pleurocarp Iwatsuki Z, Glime JM. 1984. Podperaea, a classification: two steps forward, one new genus of Hypnaceae (Musci). step back. In: Newton AE, Tangney RS, Journal of the Hattori Botanical Laboratory eds. Pleurocarpous Mosses: Systematics 55: 495-500. and Evolution. Boca Raton: CRC Press, Maeda S, Kosuge Kf Gonzalez D, De pp.1-18. Luna E, .Akiyama H. 2000. Buck WR, Goffinet Bf Shaw AJ. 2000. Molecular phylogeny of the suborder Testing morphological concepts of Leucodontineae (Musci; Leuco­ orders of pleurocarpous mosses dontales) inferred from rbcL sequence (Bryophyta) using phylogenetic data. Journal of Research 113: reconstructions based on trnL-trnF and 29-38. rps4 sequences. Molecular Phylogenetics Nishimura N. 1999. A historical review of and Evolution 16: 180-198. H ypnaceae (). B ryobrothera Crull H. 1986. Taxonomic and 5: 283-298. nomenclatural addenda to the Mexican Nishimura N, Higuchi M, Seki T, Ando H. moss flora. The Bryologist 89: 23-27. 1984. Delimitation and subdivision of Crum H. 1991. A partial clarification of the the moss family Hypnaceae. Journal of . Journal of the Hattori the Hattori Botanical Laboratory 55: Botanical Laboratory 69: 313-322. 227-234. Gardiner A, Ignatov M, Huttunen S, Noguchi A. 1972. On the delimination of Troitsky A. 2005. On resurrection of the the genera of Hylocomiaceae and families Pseudoleskeaceae Schimp. Rhytidiaceae. Journal of the Hattori and Pylaisiaceae Schimp. (Musci, Botanical Laboratory 35: 155-168. Hypnales). Taxon 54: 651-663. Ochyra R. 1982. New names for genera of Goffinet B, Buck WR. 2004. Systematics of mosses. Journal of Bryology 12: 31-32. the Bryophyta (mosses): from Schimper WP. 1856. Corollilrium Bryologiae molecules to a revised classification. Europaeae, conspectum diagnosticum In: Goffinet B, Hollowell V, Magill R, farniliarum, gencrUin et specierum, ed. Molecular Systematics of . aclnotationes novas alque emendationes St. Louis: Missouri Botanical Garden complectens. Stuttgart: E. Schweitzerbart. Press, pp. 205-239. Seki T. 1969 [1968]. A revision of the family Goffinet B, Cox C}, Shaw AJ, Hedderson Sematophyllaceae of Japan with TJ. 2001. The Bryopsida (mosses): special reference to a statistical systematic and evolutionary inferences demarcation of the family. Journal of from an rps4 gene (cpDNA) phylogeny. Science of the Hiroshima University. Series Annals of Botany 87: 191-208. B, Div. 2 12: 1-80. Hedenas L. 1989. Some neglected character distribution patterns among the pleurocarpous mosses. The Bryologist 92: 157-163.

222 Phylogenetic Analysis of Hypnaceae Based on rbcL

Thompson JD, Higgins DG, Gibson TJ. Tsubota H, Matsuda H, Mohamed H, 1994. CLUSTAL W: improving the Deguchi H. 2002b. A protocol for DNA sensitivity of progressive multiple isolation using small quantities of sequence alignment through sequence material from very small hepatic . weighting, positions-specific gap Bryological Research 8: 118-122. (In penalties and weight matrix choice. Japanese with English protocol.) Nucleic Acids Research 22: 4673-4680. Tsubota H, Nakao N, Arikawa 'I, Tsubota H, Akiyama H, Yamaguchi 'I, Yamaguchi 'I, Higuchi M, Deguchi H, Deguchi H. 2001a. Molecular phylogeny Seki T. 1999. A preliminary phylogeny of the Sematophyllaceae (Hypnales, of Hypnales (Musci) as inferred from Musci) based on chloroplast rbcL chloroplast rbcL sequence data. sequences. Journal of the Hattori Botanical Bryological Research 7:233-248. Laboratory 90: 221-240. Tsubota H, Nakao N, Yamaguchi 'I, Seki 'I, Tsubota H, Akiyama H, Yamaguchi 'I, Deguchi H. 2000. Preliminary Deguchi H. 2001b. Molecular phylogeny phylogenetic relationships of the of the genus Trismegistia and related genus Brotherella and its allied genera genera (Sematophyllaceae, Musci) (Hypnales, Musci) based on chloroplast based on chloroplast rbcL sequences. rbcL sequence data. Journal of the Hattori Flikobia 13: 529-549. Botanical Laboratory 88: 79-99. Tsubota H, Arikawa 'I, Akiyama H, De Luna E, Gonzalez 0, Higuchi M, Deguchi H. 2002a. Molecular phylogeny of hypnobryalean mosses as inferred from a large-scale dataset of chloroplast rbeL, with special reference to the Hypnaceae and possibly related families. Hikobia 13: 645-665.

223 Bryology in the New Millennium

APPENDIX A 12085 (TNS); Ectropothecium in cubans Taxon sampling and DDBJ/EMBL/ (Reinw. & Hornsch.) A. Jaeger, GenBank accession numbers. AB332294, Taiwan, N. Nishimura 12087 The sequence of information is: species, (TNS); Ectropothecium moritzii A. Jaeger, accession number, locality, and AB332293, Taiwan, N. Nishimura specimen number (herbarium). 12075 (TNS); Ectropothecium obtusulum haldanianum (Grev.) H.A. (Cardot) Z. Iwats., AB332302, Japan: Crum, AB332268, Japan: Nagano, M. Ehime, N. Nishimura 12055 (TNS); Higuchi 42196 (TNS); Calliergonella Ectropothecium ptychofolium N.Nishim, lindbergii (Mitt.) Hedenas, AB332264, AB332255, Malaysia, Borneo, M. Japan: Iwate, N. Nishimura 11954 (TNS); Higuchi 45055 (TNS); Ectropothecium chrysophyllus (Brid.) sodale (Sull.) Mitt., AB332240,Vanuatu, R.S. Chopra, AB332275, Japan: Nagano, M. Higuchi 39817 (TNS); Ectropothecium M. Higuchi 37073 (TNS); zollingeri (Mull. Hal. ) A. Jaeger, halleri (Sw. ex Hedw.) M. Fleisch., AB332295, Taiwan, N. Nishimura 12108 AB332270, Japan: Shizuoka, M. Higuchi (TNS); Ectropothecium zollinger; (Mull. 39601 (TNS); Ctenidium andoi N. Hal.) A. Jaeger, AB332305, Japan: Nishim, AB332297, Taiwan, N. Ogasawara IsIs., Y. Tateishi 20556 Nishimura 12099 (TNS); Ctenidium (TNS); ssp. calyginosus has tile (Mitt.) Lindb., AB334102, Japan: (Mitt.) Mizush., AB332277, Japan: Kochi, N. Nishimura 11145 (OKAY, TNS, Kochi, K. Kawai 2844 (TNS); HIRO); Ctenidium homalophyllum Broth. Glossadelphus ogatae Broth. & Yasuda, & Yasuda ex Iisiba, AB332298, Japan: AB332280, Japan: Miyazaki, M. Higuchi Kochi, K. Kawai 2913 (TNS); Ctenidium 47056 (TNS); Gollania japonica (Cardot) luzonense Broth., AB332310, Japan: Ando & Higuchi, AB332248, Taiwan, Kagoshima, Y. Tateishi 22288 (TNS); M. Higuchi 42036 (TNS); Gollania Ctenidium malacobolum (Mull.Hal.) philippinensis (Broth.) Nog., AB332241, Broth., AB332251, Taiwan, M. Higuchi Taiwan, M. Higuchi 40681 (TNS); 43795 (TNS); Ctenidium malacobolum Gollania sinensis Broth. & Paris, (Mull.Hal.) Broth., AB332284, Japan: AB332259, China, Sichuan, M. Higuchi Miyazaki, N. Nishimura 12039 (TNS); 29776 (TNS); Gollania taxiphylloides (Hedw.) Mitt., Ando & Higuchi, AB332242, Japan: AB334103, Japan: Kochi, N. Nishimura Tokyo, M. Higuchi 47030 (TNS); Gollania 11139 (OKAY, TNS, HIRO); Ctenidium varians (Mitt.) Broth., AB332249, Japan: precrassum Sakurai, AB332299, Japan: Kagoshima, M. Higuchi 44796 (TNS); Okayama, N. Nishimura 12109 (TNS); Gollania sp., AB332247, Taiwan, M. Ctenidium pinnatum (Broth. & Paris) Higuchi 43851 (TNS); Broth., AB332252, Taiwan, M. Higuchi capillaceum Broth. ex Iisiba, AB334106, 43853 (TNS); Ctenidium polychaetum Japan: Hiroshima, H. Tsubota 4977 (Bosch & Sande Lac.) Broth., AB332254, (HIRO); Homomallium japonico-adnatum Malaysia, Borneo, M. Higuchi 45075 (Broth.) Broth., AB332243, Japan: (TNS); Ctenidium serralifolium Cardot Tokyo, M. Higuchi 47039 (TNS); (Broth.), AB332253, Malaysia, Borneo, Hondaclla caperata (Mitt.) B.C.Tan & M. Higuchi 44960 (TNS); Dacryophyllum Z.Iwats., AB332279, Japan: Tokyo, M. falcifolium Ireland, AB332238, USA: Higuchi 47036 (TNS); California, K. M. Kellman 4674 (TNS); himalaya11lnn (Mitt.) Broth., AB332274, japonicum Cardot, AB332278, Japan: Nagano, M. Higuchi 43016 Japan: Iwate, N. Nishimura 11971 (TNS); (TNS); Hypnum calcicolum Ando, Ectropothecium andoi N. Nishim, AB332244, Japan: Tokyo, M. Higuchi AB332309, Japan: Kagoshima, Y. 47019 (TNS); Hypnum callichroum ssp. Tateishi 22286 (TNS); Ectropothecium japonicum Ando, AB332260, Japan: dealbatum (Reinw. & Hornsch.) A. Nagano, M. Higuchi 43042 (TNS); Jaeger, AB332296, Taiwan, N. Nishimura Hedw., AB332287, Japan: Fukui, N. Nishimura 12008 (TNS);

224 Phylogenetic Analysis of Hypnaceae Based on rbcL

Hypnum cupressiforme var. filiforme Brid., 40848 (TNS); Neodolichomitra AB332261, Russia, Saint Petersburg, T. yunnanensis (Besch.) T.J. Kop., Yoshida s. n. (TNS); Hypnum dieckii AB332258, China, Sichuan, M. Higuchi Renauld & Cardot, AB332263, Japan: 29749 (TNS); Oedicladium serricuspe Iwate, N. Nishimura 11987 (TNS); (Broth.) Nog. & Z. Iwats., AB332304, Hypnum fauriei Cardot, AB332262, Japan: Kagoshima, N. Nishimura 11929 Japan: Tokyo, M. Higuchi 47011 (TNS); (TNS); Pseudotaxiphyllum maebarae Hypnum fujiyamae (Broth.) Paris, (Sakurai) Z. Iwats., AB334105, Japan: AB332300, Japan: Iwate, K. Kawai 3541 Hiroshima, H. Tsubota 4956 (HIRO); (TNS); Hypnum oldhamii (Mitt.) A.Jaeger Pseudotaxiphyllum pohliaecarpum (Sull. & Sauerb., AB332281, Japan: Miyazaki, & Lesq.) Z. Iwats., AB332271, Japan: M. Higuchi 47065 (TNS); Hypnum Ibaraki, M. Higuchi 47050 (TNS); plicatulum (Lindb.) A. Jaeger & Sauerb., crista-castrensis (Hedw.) De Not., AB332266, Japan: Nagano, M. Higuchi AB332269, Japan: Nagano, M. Higuchi 43097 (TNS); Hypnum plumaeforme 41606 (TNS); Racopilum aristatum Mitt., Wilson, AB332288, Japan: Miyazaki, N. AB332291, Japan: Miyazaki, N. Nishimura 12029 (TNS); Hypnum Nishimura 12025 (TNS); plumaeforme var. minus Broth. ex Ando, rugosum (Ehrh. ex Hedw.) Kindb., AB332289, Japan: Miyazaki, N. AB332273, Japan: Nagano, M. Higuchi Nishimura 12026 (TNS); Hypnum 43103 (TNS); uncinata (Hedw.) sakuraii (Sakurai) Ando, AB332290, Loeske, AB332276, Japan: Nagano, M. Japan: Miyazaki, N. Nishimura 12030 Higuchi 42944 (TNS); Stereodontopsis (TNS); Hypnum tristo-viride (Broth.) pseudorevoluta (Reimers) Ando, Paris, AB332267, Japan: Nagano, M. AB332245, Japan: Kochi, K. Kawai 2810 Higuchi 43159 (TNS); Isopterygiopsis (TNS); Taxiphyllopsis iwatsukii Higuchi pulchella (Hedw.) Z.Iwats., AB332272, & Deguchi, AB332282, Japan: Japan: Nagano, M. Higuchi 41549 Okayama, H. Kiguchi s. n. (TNS); (TNS); IsoptenJgiopsis pulchella (Hedw.) Taxiphyllum altemans (Cardot) Z.Iwats., Z. Iwats. AB332307, New Zealand, AB332301, Japan: Okayama, M. South Isl., M. Higuchi 46764 (TNS); Chishiki 4900 (TNS); Taxiphyllum Isopterygium minutirameum (Mull. Hal.) cuspidifolium (Cardot) Z. Iwats., A. Jaeger, AB332306, Taiwan, N. AB332246, Japan: Tokyo, M. Higuchi Nishimura 12095 (TNS); Isopterygium 46994 (TNS); Vesicularia ferriei (Cardot propaguliferum Toyama, AB332286, & Ther.) Broth., AB332303, Japan: Japan: Miyazaki, N. Nishimura 12037 Ehime, N. Nishimura 12045 (TNS); (TNS); Isopterygium sp., AB332308, New Vesicula ria montagnei (Schimp.) Broth., Zealand, South lsI., M. Higuchi 46988 AB332292, Taiwan, N. Nishimura 12064 (TNS); Leptocladiella psilura (Mitt.) M. (TNS); sp., AB334104, Australia: Fleisch., AB332250, Taiwan, M. Higuchi Tasmania, H. Deguchi 36184 (HIRO).

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