Abstract
BackgroundThe class Chlorophyceae (Chlorophyta) includes morphologically and ecologically diverse green algae. Most of the documented species belong to the clade formed by the Chlamydomonadales (also called Volvocales) and Sphaeropleales. Although studies based on the nuclear 18S rRNA gene or a few combined genes have shed light on the diversity and phylogenetic structure of the Chlamydomonadales, the positions of many of the monophyletic groups identified remain uncertain. Here, we used a chloroplast phylogenomic approach to delineate the relationships among these lineages.ResultsTo generate the analyzed amino acid and nucleotide data sets, we sequenced the chloroplast DNAs (cpDNAs) of 24 chlorophycean taxa; these included representatives from 16 of the 21 primary clades previously recognized in the Chlamydomonadales, two taxa from a coccoid lineage (Jenufa) that was suspected to be sister to the Golenkiniaceae, and two sphaeroplealeans. Using Bayesian and/or maximum likelihood inference methods, we analyzed an amino acid data set that was assembled from 69 cpDNA-encoded proteins of 73 core chlorophyte (including 33 chlorophyceans), as well as two nucleotide data sets that were generated from the 69 genes coding for these proteins and 29 RNA-coding genes. The protein and gene phylogenies were congruent and robustly resolved the branching order of most of the investigated lineages. Within the Chlamydomonadales, 22 taxa formed an assemblage of five major clades/lineages. The earliest-diverging clade displayed Hafniomonas laevis and the Crucicarteria, and was followed by the Radicarteria and then by the Chloromonadinia. The latter lineage was sister to two superclades, one consisting of the Oogamochlamydinia and Reinhardtinia and the other of the Caudivolvoxa and Xenovolvoxa. To our surprise, the Jenufa species and the two spine-bearing green algae belonging to the Golenkinia and Treubaria genera were recovered in a highly supported monophyletic group that also included three taxa representing distinct families of the Sphaeropleales (Bracteacoccaceae, Mychonastaceae, and Scenedesmaceae).ConclusionsOur phylogenomic study advances our knowledge regarding the circumscription and internal structure of the Chlamydomonadales, suggesting that a previously unrecognized lineage is sister to the Sphaeropleales. In addition, it offers new insights into the flagellar structures of the founding members of both the Chlamydomonadales and Sphaeropleales.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-015-0544-5) contains supplementary material, which is available to authorized users.
Highlights
The class Chlorophyceae (Chlorophyta) includes morphologically and ecologically diverse green algae
We examined a total of 29 chlamydomonadalean and sphaeroplealean taxa that represent 16 of the 21 primary clades that Nakada et al [16] recovered for the Chlamydomonadales, three of the major lineages recognized for the Sphaeropleales (Scenedesmaceae, Mychonastaceae and Bracteacoccaceae) as well as the Jenufa lineage, which was suspected to be sister to the Golenkinia outgroup used by Nakada et al [16]
This lineage is detected as a strongly supported clade that unites the spine-bearing Golenkinia longispicula with the two Jenufa species (Figs. 1 and 3). This clade may comprise the Treubarinia; the latter multi-genera lineage, which includes Cylindrocapsa, Trochiscia and Elakatothrix in addition to Treubaria [2, 16, 37], is represented in our study by the spine-bearing Treubaria triappendiculata. While the latter alga is sister to the Golenkinia + Jenufa lineages in the Bayesian protein tree (PP = 0.98) and the maximum likelihood (ML) gene tree inferred with the PCG12RNA data set (BS = 85 %), it branches at the base of the lineages formed by the representatives of the Scenedesmaceae, Mychonastaceae and Bracteacoccace in the ML protein tree (BS = 54 %) and the gene tree inferred with the PCG123degenRNA data set (BS = 54 %)
Summary
The class Chlorophyceae (Chlorophyta) includes morphologically and ecologically diverse green algae. The Chlorophyceae occupies the crown of the Chlorophyta, one of the two divisions of the Viridiplantae [1] This monophyletic class of green algae comprises five orders or main clades [2, 3] that form two major lineages [4, 5]: the Chlamydomonadales (or Volvocales) + Sphaeropleales (CS clade) and the Oedogoniales + Chaetophorales + Chaetopeltidales (OCC clade). Members of the Chlorophyceae are found in a wide range of habitats and display diverse cell organizations (unicells, coccoids, colonies, simple flattened thalli, unbranched and branched filaments) [1, 6] Their motile cells exhibit variability at the level of the flagellar apparatus. The members of the Oedogoniales have an unusual flagellar apparatus that is characterized by a stephanokont arrangement of flagella [13]
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