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Article (Published Version) Article Revised Small Subunit rRNA Analysis Provides Further Evidence that Foraminifera Are Related to Cercozoa BERNEY, Cédric, PAWLOWSKI, Jan Wojciech Abstract There is accumulating evidence that the general shape of the ribosomal DNA-based phylogeny of Eukaryotes is strongly biased by the long-branch attraction phenomenon, leading to an artifactual basal clustering of groups that are probably highly derived. Among these groups, Foraminifera are of particular interest, because their deep phylogenetic position in ribosomal trees contrasts with their Cambrian appearance in the fossil record. A recent actin-based phylogeny of Eukaryotes has proposed that Foraminifera might be closely related to Cercozoa and, thus, branch among the so-called crown of Eukaryotes. Here, we reanalyze the small-subunit ribosomal RNA gene (SSU rDNA) phylogeny by removing all long-branching lineages that could artifactually attract foraminiferan sequences to the base of the tree. Our analyses reveal that Foraminifera branch together with the marine testate filosean Gromia oviformis as a sister group to Cercozoa, in agreement with actin phylogeny. Our study confirms the utility of SSU rDNA as a phylogenetic marker of megaevolutionary history, provided that the artifacts due to the heterogeneity of [...] Reference BERNEY, Cédric, PAWLOWSKI, Jan Wojciech. Revised Small Subunit rRNA Analysis Provides Further Evidence that Foraminifera Are Related to Cercozoa. Journal of Molecular Evolution, 2003, vol. 57, p. S120-S127 DOI : 10.1007/s00239-003-0015-2 Available at: http://archive-ouverte.unige.ch/unige:113627 Disclaimer: layout of this document may differ from the published version. 1 / 1 JMol Evol (2003) 57:S120–S127 DOI: 10.1007/s00239-003-0015-2 Revised Small Subunit rRNA Analysis Provides Further Evidence that Foraminifera Are Related to Cercozoa Ce´ dric Berney, Jan Pawlowski Department of Zoology and Animal Biology, University of Geneva, Geneva, Switzerland Received: 31 July 2002 / Accepted: 11 December 2002 Abstract. There is accumulating evidence that logeny — Long-branch attraction — Small-subunit the general shape of the ribosomal DNA-based ribosomal DNA phylogeny of Eukaryotes is strongly biased by the long-branch attraction phenomenon, leading to an artifactual basal clustering of groups that are prob- Introduction ably highly derived. Among these groups, Fora- minifera are of particular interest, because their deep Phylogenetic studies based on the small-subunit ri- phylogenetic position in ribosomal trees contrasts bosomal RNA gene (SSU rDNA) have yielded a with their Cambrian appearance in the fossil record. ‘‘classical’’ view of eukaryotic evolution, in which A recent actin-based phylogeny of Eukaryotes has several assumed primitive, isolated lineages diverged proposed that Foraminifera might be closely related very early, while other groups, supposedly more re- to Cercozoa and, thus, branch among the so-called cent and apparently simultaneously diverging, cluster crown of Eukaryotes. Here, we reanalyze the small- in the so-called eukaryotic crown (Sogin 1991; Knoll subunit ribosomal RNA gene (SSU rDNA) phylog- 1992). However, the primitive status of ‘‘early’’ eny by removing all long-branching lineages that Eukaryotes, as defined by SSU rDNA phylogenies, could artifactually attract foraminiferan sequences to has been a subject of long debate (see, e.g., Sogin the base of the tree. Our analyses reveal that Fora- 1997). Protein-based phylogenies are now providing minifera branch together with the marine testate accumulating evidences that the distinction between filosean Gromia oviformis as a sister group to ‘‘early’’ and ‘‘crown’’ Eukaryotes is no longer accu- Cercozoa, in agreement with actin phylogeny. Our rate, because many—if not all—‘‘early’’ Eukaryotes study confirms the utility of SSU rDNA as a phylo- are probably highly derived groups, which are arti- genetic marker of megaevolutionary history, provid- factually attracted to the base of SSU rDNA trees by ed that the artifacts due to the heterogeneity of the long-branch attraction (LBA) phenomenon substitution rates in ribosomal genes are circum- (Embley and Hirt 1998; Philippe and Adoutte 1998; vented. Philippe et al. 2000). Most striking is the example of the Microsporidia, which have been clearly shown to Key words: Foraminifera — Cercozoa — Euk- be highly derived, parasitic members of the Fungi aryotes — Gromia oviformis — Molecular phy- (Keeling and Doolittle 1996; Hirt et al. 1999; Keeling et al. 2000; see Van de Peer et al. [2000b] for a re- view). Correspondence to: Ce´ dric Berney, Universite´ de Gene` ve, Station Because protein data are still lacking for a large de Zoologie, 154 route de Malagnou, 1224 Cheˆ ne-Bougeries, part of the known eukaryotic diversity, the phylo- Gene` ve, Switzerland; email: [email protected] genetic position of many groups of protists remains S121 unclear. Among these groups, the Foraminifera are any support in SSU rDNA sequences for a cercozoan of particular interest, because their evolutionary his- affiliation of Foraminifera. tory can be inferred from their exceptionally well preserved fossil record. Foraminifera are a large and morphologically diverse group of mainly marine, Materials and Methods testate protists with typical filamentous, granuloret- iculose pseudopodia (Lee et al. 2000). Molecular data The complete SSU rDNA sequences of 5 Foraminifera and 49 from both large- and small-subunit rDNAs have other Eukaryotes were manually aligned using the Genetic Data Environment software (Larsen et al. 1993), following the secondary suggested a very ancient origin for the group (Paw- structure models proposed by Neefs et al. (1993) and Wuyts et al. lowski et al. 1994, 1996, 1999). But as discussed be- (2000). Sequences were chosen so that representatives of all well- fore (Pawlowski et al. 1996; Sogin 1997), rRNA genes defined domains of the ‘‘crown’’ were included in the data set, using from the Foraminifera are highly divergent. There- the studies by Van de Peer and De Wachter (1997) and Van de Peer fore, a very early branching of the group must be et al. (2000a) as references. Eukaryotic lineages that were excluded from our data set comprise all groups (Diplomonadida, Paraba- taken suspiciously. salia, Microsporidia, Euglenozoa, Heterolobosea, Radiolaria, A recent actin-based phylogeny of Eukaryotes Mycetozoa, Entamoebidae, and Pelobiontida) that are thought to showed a close relationship among the Foraminifera, have undergone a process of accelerated rate of evolution and, Cercomonas, and Chlorarachnion (Keeling 2001), thus, may act as long branches in phylogenetic analyses (see, e.g., suggesting a possible inclusion of Foraminifera Stiller and Hall 1999; Morin 2000; Philippe and Germot 2000; Bolivar et al. 2001; Lo´ pez-Garcia et al. 2002). Relative rate tests within Cercozoa. This group of ‘‘crown’’ Eukaryotes performed with RRTree (Robinson-Rechavi and Huchon 2000) was recently described on the basis of molecular data confirmed at both 5 and 1% levels that the representatives of these (Bhattacharya et al. 1995). The Cercozoa consist of a groups display significantly higher rates of substitution compared diverse assemblage of organisms, such as Cercomon- to crown species. The names, phylogenetic position, and GenBank as-like amoeboflagellates, the euglyphid testate filose accession numbers of the sequences used in this study are indicated in Table 1. amoebae, the plasmodiophorid plant pathogens, and Evolutionary trees were inferred using the neighbor-joining the chlorarachniophyte green amoebae (Cavalier- (NJ) method (Saitou and Nei 1987), the maximum-parsimony Smith 1998; see also Cavalier-Smith and Chao 1997; (MP) method, and the maximum-likelihood (ML) method (Fel- Atkins et al. 2000; Ku¨ hn et al. 2000; Bulman et al. senstein 1981). The reliability of internal branches was assessed 2001; Vickerman et al. 2002). Clear morphological using the bootstrap method (Felsenstein 1985), with 1000 replicates for NJanalyses, 500 replicates for MP analyses, and 100 replicates evidence is yet lacking to support the Cercozoa, but a for ML analyses. The Phylo_Win program (Galtier et al. 1996) was close relationship between Cercomonas and Chlor- employed for distance computations and NJbuilding and boot- arachnion was also recovered on the basis of both strapping, using the HKY85 model of substitution (Hasegawa et al. alpha- and beta-tubulin phylogenies (Keeling et al. 1985). MP and ML analyses were performed with PAUP* (Swof- 1998, 1999). ford 1998). The most parsimonious trees for each MP bootstrap replicate were determined using a heuristic search procedure with A possibility to avoid LBA artifacts, and to find 10 random-addition-sequence replicates and tree bisection–recon- the true phylogenetic position of an assumed fast- nection branch-swapping. The transversions cost was set to twice evolving, derived group of Eukaryotes, is to suppress the transitions cost. For ML analyses, the TrN model of substi- all other possibly fast-evolving taxa from the analyses tution was used (Tamura and Nei 1993), taking into account a and, as suggested by Swofford et al. (1996), to re- proportion of invariable sites, and a gamma-shaped distribution of rates of substitution among sites, with eight categories. All neces- construct a tree including only ingroup taxa. These sary parameters were estimated from the data set using Modeltest two conditions are necessary in order to avoid the (Posada and Crandall 1998). Starting trees of ML searches were attraction of
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