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Phylogeny of Oedogoniales (Chlorophyceae, Chlorophyta) Inferred from 18S Rdna Sequences with Emphasis on the Relationships in Th

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Phylogeny of Oedogoniales (Chlorophyceae, Chlorophyta) Inferred from 18S Rdna Sequences with Emphasis on the Relationships in Th Pl. Syst. Evol. 265: 179–191 (2007) Plant Systematics DOI 10.1007/s00606-007-0523-4 and Evolution Printed in The Netherlands Phylogeny of Oedogoniales (Chlorophyceae, Chlorophyta) inferred from 18S rDNA sequences with emphasis on the relationships in the genus Oedogonium based on ITS-2 sequences H. Mei1,2, W. Luo3,4, G. X. Liu1, and Z. Y. Hu1 1State Key Laboratory of Freshwater Ecology and Biotechnology, Institute of Hydrobiology, the Chinese Academy of Sciences, Wuhan, China 2Graduated School, the Chinese Academy of Sciences, Beijing, China 3Leibniz Institute of Freshwater Ecology and Inland Fisheries, Department of Limnology of Stratified Lakes, Stechlin - Neuglobsow, Germany 4SOA Key Laboratory for Polar Science, Polar Research Institute of China, Shanghai, China Received March 6, 2006; accepted January 22, 2007 Published online: May 14, 2007 Ó Springer-Verlag 2007 Abstract. The phylogeny of Oedogoniales was Introduction investigated by using nuclear 18S rDNA sequences. Results showed that the genus Oedocladium,asa The Oedogoniales, comprised of three genera separated clade, was clustered within the clade of (Oedogonium, Oedocladium and Bulbochaete), Oedogonium; whereas the genus Bulbochaete was in can be found in freshwater all over the world a comparatively divergent position to the other two (Hirn 1900, Tiffany 1937, Gemeinhardt 1939, genera. The relationship among the species of Gauthie´ r-Lie` vre 1963, Islam and Sarma 1963, Oedogonium was discussed, focusing on ITS-2 Jao 1979, Mrozin´ ska 1985). Their fascinating phylogeny analyzed combining with some morpho- and distinctive features that are well known to logical characteristics. Our results showed that all phycologists set them apart into a very unusual the dioecious nannandrous taxa involved in this order of green algae. While they have no study were resolved into one clade, while all the obvious ancestors, several features common to monocious taxa were clustered into another clade as a sister group to the former. The report also other green algae (e.g. possession of the suggests that the dioecious macrandrous taxa form phycoplast) place them in the Chlorophyta a paraphyly and could be more basally situated (sensu Mattox and Stewart 1984). But the than the dioecious nannandrous and the monoe- systematic position of this group has changed cious taxa by means of molecular phylogeny and over time according to the criteria used by morphotype investigations. different authors, who considered that mor- phological characters and certain aspects of Key words: Oedogoniales, Oedogonium, Oedo- the life cycle were the principal diacritic cladium, Bulbochaete, 18S rDNA, ITS-2, phylo- features (Alberghina et al. 2006). geny, systematics. 180 H. Mei et al.: Phylogeny of Oedogoniales Oedogonium, Bulbochaete and Oedocladium phyly of Oedogoniales, and the Oedogonium are separated based on differences in their group did not appear to be monophyletic. Since filaments and the presence or absence of hairs, the traditional taxonomy of Oedogoniales did however few characters in morphology and not define natural groups and the evolutionary structure could be used to discuss the evolu- position remained uncertain, it was necessary to tionary course of the three genera (Jao 1979). clarify the systematic problem and confirm Tiffany (1930) arranged them in Bulbochaete, many evolutionary hypotheses in a deeper Oedocladium and Oedogonium; Gauthie´ r- way. Except for 18S rDNA, internal transcribed Lie` vre (1963) chose the order Oedocladium, spacer (ITS) sequences of nuclear ribosomal Bulbochaete and Oedogonium; Jao (1979) also DNA have been shown to provide good phylo- agreed with Hirn’s conclusion (1900) as Oe- genetic resolution in recently diverging lineages dogonium, Bulbochaete and Oedocladium; (Kooistra et al. 1992; Van Oppen et al. 1993, However, recent illustration of Mrozin´ ska 1995; Leclerc et al. 1998). In this paper, more (1985) sorted them in the order Oedogonium, 18S rDNA sequences with an emphasis on ITS- Oedocladium and Bulbochaete. The relation- 2ofOedogonium species were involved, which ships of the three genera are still uncertain. obtained a better understanding of the relation- During the past few years various phycolo- ship among the three genera, and the phylogeny gists used different characteristics as criterion among Oedogonium species with a magnitude of for dividing in the three genera of Oedogoniales. sexual differentiation types furtherly. We lim- Jao’s monographic work (1979) based on sam- ited the spacer analysis to ITS-2 because this ple collections from China, presented types of part of sequence contained higher base conser- sexual differentiation as groundwork for classi- vation and it was proven to be most useful at fication below the rank of genus Oedogonium species and genus level (Van Nues et al. 1995; and genus Bulbochaete. Since living samples of Hershkovitz and Lewis 1996; Coleman and Mai genus Oedocladium had not been found, until 1997; Stiger et al. 2000, 2003; Coleman 2003; Liu (1993) did report this new record in China. Hegewald and Wolf 2003). In addition, the ITS- Mrozin´ ska (1991, 1993) proposed to divide the 2 region is informative and easy to be aligned genera Oedogonium and Bulbochaete into sec- guided by secondary structure (Mai and Cole- tions based on the number of spermatozoids man 1997). Morphological characters were also produced in the antheridial cell respectively, combined with the ITS-2 study to assess the which characteristics should be chosen as the relationships among Oedogonium taxa. criterion in taxonomical classification below the rank of genera. Yet the evolutionary relation- ships between the genera remained problematic. Materials and methods To date, only a limited number of molecular Sampling and cultivation. Twelve Oedogonium taxa phylogenetic studies have been based on and one Oedocladium taxon were collected respec- sequences of the Oedogoniales. Correlative tively in Hubei Province 110.6°)115°E, studies (Booton et al. 1998, Buchheim et al. 30.4°)31.7°N (PRC) and Zhangjiajie 110.4°E, 2001, Shoup and Lewis 2003, Krienitz et al. 29.1°N (Hunan Province, PRC). Samples were 2003) have taken not more than four species of isolated from field by the authors and cultivated this order into analyses, and have only drawn at 20°Cto25°C at a 16:8-h light:dark cycle under 25 lmol photons m)2Æs)1, in modified Bold’s Basal preliminary conclusions: The Oedogoniales was medium (BBM, Nichols and Bold 1965) supple- Bulbochaete )1 monophyletic; and could be more mented with soilwater 70 mlÆl . basally situated phylogenetically than the other Taxon selection. A total of 28 Oedogoniales two genera. The latest research of Alberghina taxa were selected in 18S rDNA or ITS-2 sequence (2006) reported a further phylogenetic analysis analyses. All these organisms, their origins, strain using 18S rDNA of 10 Oedogonium species in number or reference and the GenBank accession Argentina, which also demonstrated the mono- numbers of each sequence are listed in Table 1. For Table 1. Oedogoniales taxa used for 18S rDNA or ITS-2 sequence analyses, strain source, strain No. or reference, Genbank accession numbers H. Mei et al.: Phylogeny of Oedogoniales 181 and sexual reproduction type and sperm No. in each antheridium of the Oedogonium species used in ITS-2 analysis. Sequences marked with an asterisk were determind by this study. FACHB: Freshwater Algae Culture Collection, Institute of Hydrobiology, Chinese Academy of Science Taxon, strain source 18S rDNA ITS-2 sequence Type of No. of sexual sperm per differentiation antheridium Strain No. or Strain No. or reference Acc.No. reference Acc.No. Bulbochaete rectangularis var. Booton U83132 UTEX LB954. AY962677 hiloensis (Nordst) Tiffany et al. 1998 Oedocladium carolinianum Beaney Booton U83135 & Hoffman et al. 1998 Oedocladium prescottii Islam * (Sandy FACHB 993 DQ078298 soil, Zhangjiajie, China) Oedogonium acrosporum De Bary Alberghina DQ115892 et al. 2006 Oedogonium angustistomum Hoffmann Booton U83134 UTEX LB1557 AY962676 Dioecious 2 et al. 1998 macrandrous Oedogonium borisianum UTEX LB2239 AY962670 Dioecious 1 (Le Clerc) Wittr., nannandrous Oedogonium brevicingulatum Jao FACHB 999 DQ413052 FACHB 999 DQ413058 Monoecious 2 * (Pond at Institute of Hydrobiology, CAS, Wuhan, China) Oedogonium cardiacum Wittr. Booton U83133 UTEX LB40 AY962675 Dioecious 2 et al. 1998 macrandrous Oedogonium calliandrum Hoffm. UTEX LB1554 AY962672 Dioecious 2 macrandrous Oedogonium crispum (Hass.) Wittr. ACOI 1287 AY962680 Monoecious 2 Oedogonium cylindrosporum Jao * (Pond Present DQ078297 Present study DQ078300 Dioecious 2 at Zhifang, Wuhan, China) study macrandrous Oedogonium eminens (Hirn) Tiff. * (Pond Present study DQ078302 Dioecious 1 at Zhifang, Wuhan, China) nannandrous Oedogonium fragile Wittr. SCCAP K0093 AY962679 Monoecious 2 Oedogonium globosum Nordst. * (Pond FACHB 992 DQ413051 FACHB 992 DQ413057 Monoecious 2 at Zhifang, Wuhan, China) Table 1. (Continued) 182 H. Mei et al.: Phylogeny of Oedogoniales Taxon, strain source 18S rDNA ITS-2 sequence Type of No. of sexual sperm per differentiation antheridium Strain No. or Strain No. or reference Acc.No. reference Acc.No. Oedogonium nodulosum Wittr. * (Pond FACHB 996 DQ018735 FACHB 996 DQ078301 Monoecious 2 at Macheng city, Hubei Province, China) Oedogonium oblongum Wittr., ACOI 1118 AY962681 Monoecious 2 Oedogonium pakistanense Islam & Sarma FACHB 995 DQ076244 FACHB 995 DQ413060 Monoecious 2 * (Sandy soil, Xiantao County, Hubei Province, China) Oedogonium pseudohirnii Jao * (Soil, FACHB 994
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