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A Molecular Contribution for Understanding the Lembophyllaceae (Bryopsida) Based on Noncoding Chloroplast Regions (Cpdna) and It

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A Molecular Contribution for Understanding the Lembophyllaceae (Bryopsida) Based on Noncoding Chloroplast Regions (Cpdna) and It J. Hattori Bot. Lab. No. 89: 71- 92 (Dec. 2000) A MOLECULAR CONTRIBUTION FOR UNDERSTANDING THE LEMBOPHYLLACEAE (BRYOPSIDA) BASED ON NONCODING CHLOROPLAST REGIONS (CPDNA) AND ITS2 (NRDNA) SEQUENCE DATA STUDIES IN AUSTRAL TEMPERATE RAIN FOREST BRYOPHYTES 8 12 3 2 DIETMAR QUANDT ' , RAYMOND S. TANGNEY , JAN-PETER FRAHM 1 and WOLFGANG FREY ABSTRACT. So far morphological approaches have produced little consensus in formulating clear definitions of the mostly southern hemispheric Lembophyllaceae and Meteoriaceae, two pleurocar­ pous and mainly epiphytic moss families. The generic limits proposed by various authors have result­ ed in an overlap of family concepts, and the systematic position of genera such as Weymouthia and Pilotrichella has been unsettled since the description of these families by Brotherus ( 1907) and Kind­ berg ( 1897). A molecular approach based on sequence data of two non coding regions of the chloro­ plast DNA (cpDNA), trnLuAA intron and trnLuAA 3'exon- trnFGAA intergenic spacer (trn data) as well as sequence data of the internal transcribed spacer 2 (ITS2) of the nuclear ribosomal DNA (nrDNA) provides new evidence supporting the transfer of the genus Weymouthia into the Lembophyllaceae. Although the inter- and intrageneric sequence divergence is extremely low, the highly congruent maximally parsimonious trees from trn and ITS2 data sets suggest a close relationship of the Lembo­ phyllaceae taxa sensu Tangney ( 1997 b ). The Lembophyllaceae as treated here contain five genera Camptochaete, Lembophyllum, Fifea, Fallaciella and Weymouthia . Within the Meteoriaceae Meteori­ um and Papillaria are retained as separate genera, while Weymouthia is excluded from the Meteori­ aceae and transferred to the Lembophyllaceae. The molecular data does not align Pilotrichella with either the Meteoriaceae or the Lembophyllaceae sampled. A transfer of Pilotrichella into the Lembo­ phyllaceae is not supported. Pulchrinodus in.flatus is neither part of the genus Weymouthia nor of the Meteoriaceae or Lembophyllaceae. KEY WORDS : Lembophyllaceae, Meteoriaceae, Bryopsida, noncoding chloroplast DNA, nuclear ribo­ somal DNA, molecular systematics, molecular evolution INTRODUCTION One of the most striking features of the humid temperate rain forests occurring in New Zealand and Tasmania certainly are the epiphytic bryophytes, especially members of the Lembophyllaceae and Meteoriaceae. These two pleurocarpous moss families, almost limited to the Southern Hemisphere, are partly characterised by their weft-forming or pen- 1 Institut fiir Biologie-Systematische Botanik und Pfianzengeographie- , Freie Universitiit Berlin, Altensteinstraf3e 6, D-14195 Berlin, Germany. 2 Botanisches Institut und Botanischer Garten, Rheinische Friedrich-Wilhelms-Universitiit Bonn, Meckenheimer Allee 170, D-53115 Bonn, Germany. 3 Department of Botany, University of Otago, P. 0. Box 56, Dunedin, New Zealand. Present ad­ dress: Department of Biodiversity and Systematic Biology, National Museum and Gallery Cardiff, Cathays Park, Cardiff CF JO 3NP, United Kingdom. 72 J. Hattori Bot. Lab. No. 89 2 0 0 0 dant life-form and a commonly epiphytic habitat. Since their first description by Kindberg (1897, Meteoriaceae, cited in Buck 1994) and Brotherus (1907, Lembophyllaceae, and Me­ teoriaceae worldwide), there has been little agreement over family concepts. This has been reflected in recent years by substantial changes concerning the genera and the status of dif­ ferent members earlier included within these families (Buck & Vitt 1986, Crum 1991, Buck 1994, Tangney 1997 b ). Initially, Brotherus ( 1907) included four genera within the Lembophyllaceae: Lembo­ phyllum, Camptochaete, Dolichomitra, and lsothecium. By adding Porotrichopsis, Rigodi­ um, and Acrocladium, as well as his new genus Plasteurhynchium Fleischer (1915- 23) drastically enlarged the family, after he had already added Tripterocladium and his newly described genus Porotrichodendron (Fleischer 1906-08). Subsequently, Brotherus ( 1925), largely followed Fleischer's ( 1906-08, 1915- 23) treatment of the family and accepted the inclusions. However, he (Brotherus 1925) added Dolichomitriopsis and his new genus Elmeriobryum, while Isotheciopsis was included later on (Brotherus 1929). Horikawa & Ando (1964) expanded the family to 14 genera by adding Dixonia. As circumscribed by Fleischer (I 906--08, I 9 I 5- 23) and Brotherus (1907, I 925), this family seemed to be no more than an assortment of genera of little similarity and doubtful relationship (Andrews 1952, Crum 1991). Therefore, some recent authors drastically re­ duced the family to the key genera Camptochaete and Lembophyllum (Buck 1980, Buck & Vitt 1986, Tangney 1997 b ), while discussing an inclusion of the genus Weymouthia (Streimann I 99 I b, Buck 1994, Tangney 1997 b ). Crum (1991) even excluded Camp­ tochaete, leaving the Lembophyllaceae only with Fifea (formerly part of Camptochaete) and Lembophyllum. Whereas Walther (1983) only excluded the four genera Acrocladium, Tripterocladium, lsothecium, and Elmeriobryum from the former 14 in his synopsis. While Buck ( 1980) as well as Buck & Vitt ( 1986) considered only Camptochaete and Lembophyl­ lum within the Lembophyllaceae, Buck ( L994) transferred Weymouthia, Squamidium, Pi­ lotrichella and Pseudopilotrichum, formerly Meteoriaceae, subfamily Pilotrichelloideae (Brotherus I 925), into the Lembophyllaceae. The most recent generic revision of the Lem­ bophyllaceae (Tangney 1997 b) proposed a solution comparable to Buck & Vitt (I 986). Tangney (I 997 b) only included Lembophyllum and Camptochaete as well as the two monotypic genera Fifea and Fallaciella (both formerly Camptochaete), and Weymouthia within the Lembophyllaceae. But it has to be mentioned that a proposed relationship of the genera Squamidium, Pilotrichella, and Pseudopilotrichum (Buck I 994) was not part in the study of Tangney (1997 b ). One of the most interesting genera is the palaeoaustral genus Weymouthia with its dis­ cussed relationship either to Papillaria and Pilotrichella (both Meteoriaceae) or Lembo­ phyllum (Lembophyllaceae). Assuming a close relationship to the genera Papillaria and Pi­ /otrichella, Weymouthia was originally placed within the Meteoriaceae (Brotherus 1907, 1925). On the other hand, Lembophyllum (Weymouthia) cochlearifolium (Schwaegr.) Lindb. was initially treated by Brotherus (1907, I 925) within the monotypic section Pseudo-Wey­ mouthia as part of the Lembophyllaceae. Later, Dixon (I 927) recognized that Weymouthia billardierei is conspecific to L. cochlearifolium and synonymised both under Weymouthia coch/earifolia. While discussing a close relationship between the genera Weymouthia and D. Q UANDT et al. : A molecular contribution for understanding the Lembophyllaceae 73 trnl uAA trnluAA trnFoAA 5'-Exon 3'-Exon 265 bp lntron Spacer c- E- Fig. 1. The trnLuAA- trnF0 AA region of the chloroplast genome (cpDNA). Positions and directions of primers (C, D, E, and F) used to amplify the two analysed noncoding re­ gions (trnLuAA intron and trnLuAA 3 'exon- trnF GAA intergenic spacer) are indicated below and correspond to Taberlet et al. ( 1991 ). Sizes with respect to Weymouthia cochlearifolia (Schwaegr.) Dix. lembophyllum, Dixon ( 1927) retained the systematic position of Weymouthia within the Meteoriaceae. Subsequently, this hypothesized close relationship was pointed out by sever­ al authors (Streimann 1991 b, Buck 1994, Tangney 1997 b ). Due to the interconnected taxonomic history, it is necessary to examine the relation­ ship between the Lembophyllaceae and Meteoriaceae, and to investigate the systematic po­ sition of different species or genera discussed in both families. As morphological ap­ proaches have not been able to provide agreement on definitions of the Meteoriaceae and Lembophyllaceae, a molecular approach seems to be of considerable value for gathering further evidence to resolve the relationships between the different taxa of the Lembophyl­ laceae and Meteoriaceae. Sequence data provides powerful evidence for resolving the relationship among close­ ly related species. During the past decade, several molecular markers from the chloroplast DNA (cpDNA) or the nuclear genome have become available for inferring plant phyloge­ nies. Initially, cpDNA analysis of angiosperms mainly focussed on coding regions like rbcL (e.g. Chase et al. 1993, Price & Palmer 1993), matK (e.g. Hilu & Liang 1997, Hilu et al. 2000) or atpB (e.g. Hoot et al. 1995). More recently noncoding chloroplast regions be­ came popular, mainly to address questions on lower taxonomic level or concerning biogeo­ graphic questions (e.g. Gielly et al. 1996, Small et al. 1998, Frey et al. 1999, Bayer et al. 2000). Few studies have been carried out to resolve the relationships on lower taxonomic level within bryophytes so far. Most of these are intrafamiliar studies based on cpDNA and focus on rbcL (Orthotrichaceae, Goffinet et al. 1998), rps4 (Hyvonen et al. 1999) or the trnT-trnL-trnF region (Dicranaceae, Stech 1999 a,b ). The trnLuAA intron (Fig. 1) as well as the trnLuAA 3 'exon- trnF GAA intergenic spacer (Fig. 1), we used in our study was first intro­ duced by Taberlet et al. ( 1991) as a tool for molecular systematics and seems to be very useful at the inter- and intrageneric level (Bohle et al. 1996, Gielly & Taberlet 1996 a,b, Frey et al. 1999, Stech 1999 a,b,c,d). 74 J. Hattori Bot. Lab. No. 89 2 0 0 0 Repeat unit .. Transcription unit .. 5'-ETS 5.8S \ 18S nrDNA nrDNA 26S nrDNA IGS (NTS) 18S nrDNA 4 ~~--_!---- !/ ~ 1 2 _118S nrDNA
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