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Ebihara, A, 2011. Rbcl Phylogeny of Japanese Pteridophyte Flora And

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Ebihara, A, 2011. Rbcl Phylogeny of Japanese Pteridophyte Flora And Bull. Natl. Mus. Nat. Sci., Ser. B, 37(2), pp. 63–74, May 22, 2011 RbcL Phylogeny of Japanese Pteridophyte Flora and Implications on Infrafamilial Systematics Atsushi Ebihara Department of Botany, National Museum of Nature and Science, Amakubo 4–1–1, Tsukuba, 305–0005 Japan E-mail: [email protected] (Received 15 February 2011; accepted 23 March 2011) Abstract A molecular phylogenetic analysis of the Japanese pteridophyte flora was performed using chloroplast rbcL sequences of 93% of Japanese taxa. The obtained tree is provided here and noteworthy or novel results on the infrafamilal taxonomy of the pteridophytes are documented by family. Key words : infrafamilial taxonomy, lycophyte, monilophyte, pteridophyte, rbcL. The pteridophyte flora of Japan comprising genetic analysis was performed by MrBayes 733 taxa has been almost covered by a sequence 3.1.2 (Ronquist and Huelsenbeck, 2003); each of data-set for DNA barcoding (Ebihara et al., four Markov chain Monte Carlo (MCMC) chains 2010). The barcoding project used two chloro- for two independent runs, 20 million generations, plast DNA regions, rbcL and trnH-psbA, and the sampled every 1000 generations using the former alone is not enough for solving deep phy- GTRϩIϩG substitution model, using the “con- logeny at family or higher levels, while the latter straint” options for the families defined by Smith seems unsuitable for phylogenetic analysis due to et al. (2006) except for Woodsiaceae and Dry- its frequent indels. Even if the resolution is limit- opteridaceae whose monophyly is not supported ed, it is worth while to visualize general phyloge- in the analysis by Schuettplez and Pryer (2007). netic relationships in rich floristic sampling. In The first half (10000) trees were discarded as a monilophytes, family-level phylogeny has almost burn-in period based on the diagnosis of the been solved using multiple DNA markers result by Tracer 1.5 (Rambaut and Drummond, (Schuettpelz and Pryer, 2007) and is reflected in 2007). The obtained consensus tree is rooted recent classifications (Smith et al., 2006). In- with lycophytes (Lycopodiaceae, Selaginellaceae frafamiliar relationships, especially those that and Isoetaceae). have not yet appeared in previous studies, are discussed in this paper. Results and Discussion The tree reconstructed in this paper (a tree di- Materials and Methods vided into ten parts; Figs. 1–10) show the present Chloroplast rbcL sequences of 1205 bp in results of Bayesian inference of infrafamiliar length used for the present analysis are exactly relationships and the family or higher-level rela- the same as those used in Ebihara et al. (2010), tionships by Schuettpeltz and Pryer (2007). Since but five of 689 sequences examined by Ebihara et two of the Smith et al. (2006)’s families, Dry- al. (2010) were excluded from the present analy- opteridaceae and Woodsiaceae, are posssibly sis because of their incomplete length. A phylo- non-monophyletic (Schuettpeltz and Pryer, 64 Atsushi Ebihara 2007), infrafamiliar relationships by Schuettpeltz Ophioglossaceae (Fig. 2) and Pryer (2007) are partially adopted for the two The paraphyletic placement of Ophioglossum families. in the present outcome does not match the result Twenty-nine of the 37 families recognized by by Hauk et al. (2003) which suggested its mono- Smith et al. (2006) plus three lycophyte families phyly. The most diversified group of Ophioglos- are distributed in Japan. The following discus- saceae in Japan is Botrychium sect. Sceptridium sion is in sequence of the families adopted by comprising more than 10 taxa. In the sect. Scep- Smith et al. (2006), but families without any tridium all the three polyploid species [tetraploid novel knowledge are omitted. The infrafamilial B. formosanum Tagawa (Sahashi, 1981), and classifications generally follow Iwatsuki (1995), hexaploid B. atrovirens (Sahashi) M. Kato (Sa- but have been modified adopting the results of hashi, 1979) and B. japonicum (Prantl) Underw. recent study. (Sahashi, 1984)] form a distinct clade. Lycopodiaceae (Fig. 1) Cyatheaceae (Fig. 3) Both the Huperzia group and the Lycopodiella Cyathea lepifera (J. Sm. ex Hook.) Copel. and group are supported as monophyletic, but the Ly- C. mertensiana (Hance) C. Chr. et Tardieu be- copodium s.s. group is paraphyletic in the present long to the Sphaeropteris group, one of the three tree. The latter result does not match the result of major groups of the genus (Conant et al., 1996; Wikström and Kenrick (2000) using rbcL and Korall et al., 2007), and the remaining six Japan- trnL intron sequences. ese species belong to the Cyathea s.s. group. Cyathea tuyamae H. Ohba, an endemic species Fig. 1. A part of the results of Bayesian inference of infrafamiliar phylogeny including Lycopodiaceae, Selaginellaceae and Isoetaceae, based on chloroplast rbcL sequences of 684 Japanese pteridophytes. Posterior probabilities Ն0.90 are shown. Family delimitations, shown by bars on the right side, follow Schuettpelz and Pryer (2008). RbcL Phylogeny of Japanese Pteridophyte 65 Fig. 2. A part of the results of Bayesian inference of infrafamiliar phylogeny including Ophioglossaceae, Psilotaceae, Equisetaceae and Marattiaceae, based on chloroplast rbcL sequences of 684 Japanese pterido- phytes, rooted with lycophytes. Posterior probabilities Ն0.90 are shown. Family delimitations, shown by bars on the right side, follow Smith et al. (2006). Fig. 3. A part of the results of Bayesian inference of infrafamiliar phylogeny including Osmundaceae, Hymeno- phyllaceae, Gleicheniaceae, Dipteridaceae, Lygodiaceae, Schizaeaceae, Marsileaceae and Salviniaceae, based on chloroplast rbcL sequences of 684 Japanese pteridophytes, rooted with lycophytes. Posterior probabilities Ն0.90 are shown. Schematic relationships between families follow Smith et al. (2006). 66 Atsushi Ebihara of Minami-iojima Isl. (Volcano Isls.) character- (Kramer, 1971). Further careful study is neces- ized by branching trunks, is closely related to C. sary especially for bipinnate Lindsaea species. spinulosa Wall. ex Hook. having non-branching trunks. Dennstaedtiaceae (Fig. 4) The current generic classifications are support- Lindsaeaceae (Fig. 4) ed by the present phylogeny. Five of the eight The genus Lindsaea is apparently polyphylet- Microlepia taxa share exactly the same rbcL ic; L. odorata Roxb. (sect. Osmolindsaea) is sis- sequence — this suggests some Japanese Mi- ter to Tapeinidium. Among the other Lindsaea crolepia species are hybrid-origin polyploids, and species, L. orbiculata (Lam.) Mett. ex Kuhn, L. share a maternal ancestor. simulans Ching, L. cambodgensis H. Christ and L. kawabatae Sa. Kurata form an unsolved com- Pteridaceae (Fig. 5) plex. Lindsaea heterophylla Dryand. is also in- All of the five robust clades recognized by cluded in the complex, and this result does not Schuettpelz et al. (2007) within Pteridaceae are conflict with the hypothesis of its hybrid origin distributed in Japan. The PT clade including Fig. 4. A part of the results of Bayesian inference of infrafamiliar phylogeny including Cyatheales (Plagiogyri- aceae, Cibotiaceae and Cyatheaceae), Lindsaeaceae and Dennstaedtiaceae, based on chloroplast rbcL sequences of 684 Japanese pteridophytes, rooted with lycophytes. Posterior probabilities Ն0.90 are shown. Schematic relationships between families follow Smith et al. (2006). RbcL Phylogeny of Japanese Pteridophyte 67 Fig. 5. A part of the results of Bayesian inference of infrafamiliar phylogeny including Pteridaceae based on chloroplast rbcL sequences of 684 Japanese pteridophytes, rooted with lycophytes. Posterior probabilities Ն0.90 are shown. Names of the clades are based on Schuettpelz et al. (2007). Schematic relationships between families follow Smith et al. (2006). Pteris, Onychium and Pityrogramma (intro- P. oshimensis Hieron., P. setulosocotulata Hayata duced), has the largest number of members. The and P. yakuinsularis Sa. Kurata) form an un- other four clades are: AD (Adiantum, Hap- solved complex. Considering the fact that they lopteris [Vittaria] and Antrophyum), CR (Co- are mostly known as apogamously reproducing niogramme and Cryptogramma), CH (Cheilan- taxa (Takamiya, 1996), detailed study on their thes) and CE (Ceratopteris and Acrostichum). In origins and relationships is necessary. the PT clade, the position of Pteris vittata L. is unsolved, but the tree clearly suggests at least the Aspleniaceae (Fig. 6) species occupies the most basal position among There is little discordance with previous phy- Japanese Pteris species. Pteris fauriei Hieron. logenetic analysis (Schneider et al., 2004a). Of and its related species (P. boninensis H. Ohba, P. nine clades recognized by Schneider et al. kawabatae Sa. Kurata, P. kiuschiuensis Hieron., (2004a), members of the seven clades are distrib- P. laurisilvicola Sa. Kurata, P. natiensis Tagawa, uted in Japan. All the Hymenasplenium species 68 Atsushi Ebihara Fig. 6. A part of the results of Bayesian inference of infrafamiliar phylogeny including Aspleniaceae based on chloroplast rbcL sequences of 684 Japanese pteridophytes, rooted with lycophytes. Posterior probabilities Ն0.90 are shown. Names of the clades are based on Schneider et al. (2004a) except for Ab, which is an unnamed clade including Asplenium bullatum. Schematic relationships between families follow Smith et al. (2006). are included in Clade I [the names of the clades Baker, A. pekinense Hance, A. ruprechtii Sa. follow those in Schneider et al. (2004a)]. Asple- Kurata, A. sarelii Hook., A. shimurae (H. Itô) nium ruta-muraria L.
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