@ARTICLE{TreeBASE2Ref15322, author = {Casey W Dunn and Andreas Hejnol and David Q Matus and Kevin Pang and William E Browne and Stephen A Smith and Elaine Seaver and G. W. Rouse and Matthias Obst and Gregory D. Edgecombe and Martin V S?rensen and Steven H. D. Haddock and Andreas Schmidt-Rhaesa and Akiko Okusu and R. M. Kristensen and Ward C. Wheeler and Mark Q. Martindale and Gonzalo Giribet}, title = {Broad phylogenomic sampling improves resolution of the Animal Tree of Life}, year = {2008}, keywords = {}, doi = {10.1038/nature06614}, url = {}, pmid = {18322464 }, journal = {Nature}, volume = {452}, number = {7188}, pages = {745--749}, abstract = {Long-held ideas regarding the evolutionary relationships among animals have recently been upended by sometimes controversial hypotheses based largely on insights from molecular data1,2. These new hypotheses include a clade of moulting animals (Ecdysozoa)3 and the close relationship of the lophophorates to molluscs and annelids (Lophotrochozoa)4. Many relationships remain disputed, including those that are required to polarize key features of character evolution, and support for deep nodes is often low. Phylogenomic approaches, which utilize data from many genes, have shown promise for resolving deep animal relationships, but are hindered by a lack of data from many important groups. Here we report a total of 39.9 Mb of Expressed Sequence Tags (ESTs) from 29 animals (Supplementary Table 1) belonging to 21 phyla, including 11 phyla previously lacking genomic or EST data. Analyzed in combination with existing sequences (Figs. 1, 2; Supplementary Fig. 1), our data reinforce several previously identified clades that split deeply in the animal tree (including Protostomia, Ecdysozoa, and Lophotrochozoa), unambiguously resolve multiple long-standing issues for which there was strong conflicting support in earlier studies with less data (such as velvet worms rather than tardigrades as the sister group of arthropods5), and provide molecular support for the monophyly of molluscs, a group long recognized by morphologists, for the first time. In addition, we find strong support for several new hypotheses. These include a clade that unites annelids (including sipunculans and echiurans) with nemerteans, phoronids, and brachiopods, molluscs as sister to that assemblage, and the placement of ctenophores as the earliest diverging extant multicellular animals. A single origin of spiral cleavage (with subsequent losses) is inferred from well-supported nodes. Many relationships between a stable subset of taxa find strong support, and a diminishing number of lineages remain recalcitrant to placement on the tree.} }

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