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For ODE special issue “New Animal Phylogeny”:New Animal Phylogeny: Future challenges for animal phylogeny in the age of phylogenomicsGonzalo GiribetMuseum of Comparative Zoology & Department of Organismic and Evolutionary Biology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USAEmail: ggiribet@g.harvard.eduAbstractThe science of phylogenetics, and specially the subfield of molecular systematics, has grown exponentially not only in the amount of publications and general interest, but especially in the amount of genetic data available. Modern phylogenomic analyses use large genomic and transcriptomic resources, yet a comprehensive molecular phylogeny of animals, including the newest types of data for all phyla, remains elusive. Future challenges need to address important issues with taxon sampling—especially for rare and small animals—, orthology assignment, algorithmic developments, data storage, and to figure out better ways to integrate information from genomes and morphology in order to place fossils more precisely in the animal tree of life. Such precise placement will also aid in providing more accurate dates to major evolutionary events during the evolution of our closest kingdom.Keywords Genomics – Transcriptomics – Metazoan phylogeny – New animal phylogeny – Fossils – Tip dating – Total evidence datingIntroductionAlmost two decades have passed since the publication of what has become to be known as “The New Animal Phylogeny” PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5BZG91dHRlPC9BdXRob3I+PFllYXI+MjAwMDwvWWVhcj48
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ADDIN EN.CITE.DATA (Adoutte et al. 2000; Halanych 2004)—a new set of phylum-level relationships largely driven by molecular data and mostly derived from two of the most influential papers on animal molecular phylogenetics PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5IYWxhbnljaDwvQXV0aG9yPjxZZWFyPjE5OTU8L1llYXI+
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ADDIN EN.CITE.DATA (Laumer et al. 2015a)—and Ecdysozoa ADDIN EN.CITE <EndNote><Cite><Author>Aguinaldo</Author><Year>1997</Year><RecNum>4044</RecNum><DisplayText>(Aguinaldo et al. 1997)</DisplayText><record><rec-number>4044</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907191">4044</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Aguinaldo, A. M. A.</author><author>Turbeville, J. M.</author><author>Lindford, L. S.</author><author>Rivera, M. C.</author><author>Garey, J. R.</author><author>Raff, R. A.</author><author>Lake, J. A.</author></authors></contributors><titles><title>Evidence for a clade of nematodes, arthropods and other moulting animals</title><secondary-title>Nature</secondary-title></titles><periodical><full-title>Nature</full-title></periodical><pages>489-493</pages><volume>387</volume><keywords><keyword>18S rDNA</keyword><keyword>Animal</keyword><keyword>Animals</keyword><keyword>arthropod</keyword><keyword>Arthropoda</keyword><keyword>Arthropods</keyword><keyword>clades</keyword><keyword>Ecdysozoa</keyword><keyword>Kinorhyncha</keyword><keyword>Lophotrochozoa</keyword><keyword>Metazoa</keyword><keyword>molecular phylogeny</keyword><keyword>molting animals</keyword><keyword>moulting</keyword><keyword>Nematoda</keyword><keyword>nematode</keyword><keyword>nematomorpha</keyword><keyword>Onychophora</keyword><keyword>phylogeny</keyword><keyword>Priapulida</keyword><keyword>Tardigrada</keyword><keyword>POY book</keyword></keywords><dates><year>1997</year><pub-dates><date>1997</date></pub-dates></dates><label>1599</label><urls></urls></record></Cite></EndNote>(Aguinaldo et al. 1997). These studies were based on analyses of 18S rRNA sequence data and were subsequently corroborated by a series of likewise influential papers using larger 18S rRNA data sets, additional markers, and sometimes morphology PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5acnphdsO9PC9BdXRob3I+PFllYXI+MTk5ODwvWWVhcj48
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ADDIN EN.CITE.DATA (e.g., Giribet et al. 1996; Zrzav? et al. 1998; Giribet et al. 2000; Peterson and Eernisse 2001). Few other major changes in animal phylogeny are comparable to the ones introduced by PEVuZE5vdGU+PENpdGUgQXV0aG9yWWVhcj0iMSI+PEF1dGhvcj5IYWxhbnljaDwvQXV0aG9yPjxZ
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ADDIN EN.CITE.DATA Halanych et al. (1995) and ADDIN EN.CITE <EndNote><Cite AuthorYear="1"><Author>Aguinaldo</Author><Year>1997</Year><RecNum>4044</RecNum><DisplayText>Aguinaldo et al. (1997)</DisplayText><record><rec-number>4044</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907191">4044</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Aguinaldo, A. M. A.</author><author>Turbeville, J. M.</author><author>Lindford, L. S.</author><author>Rivera, M. C.</author><author>Garey, J. R.</author><author>Raff, R. A.</author><author>Lake, J. A.</author></authors></contributors><titles><title>Evidence for a clade of nematodes, arthropods and other moulting animals</title><secondary-title>Nature</secondary-title></titles><periodical><full-title>Nature</full-title></periodical><pages>489-493</pages><volume>387</volume><keywords><keyword>18S rDNA</keyword><keyword>Animal</keyword><keyword>Animals</keyword><keyword>arthropod</keyword><keyword>Arthropoda</keyword><keyword>Arthropods</keyword><keyword>clades</keyword><keyword>Ecdysozoa</keyword><keyword>Kinorhyncha</keyword><keyword>Lophotrochozoa</keyword><keyword>Metazoa</keyword><keyword>molecular phylogeny</keyword><keyword>molting animals</keyword><keyword>moulting</keyword><keyword>Nematoda</keyword><keyword>nematode</keyword><keyword>nematomorpha</keyword><keyword>Onychophora</keyword><keyword>phylogeny</keyword><keyword>Priapulida</keyword><keyword>Tardigrada</keyword><keyword>POY book</keyword></keywords><dates><year>1997</year><pub-dates><date>1997</date></pub-dates></dates><label>1599</label><urls></urls></record></Cite></EndNote>Aguinaldo et al. (1997), perhaps followed closely only by another major re-arrangement related to Platyhelminthes, proposing their non-monophyly and a special position for Acoela and Nemertodermatida, as sister groups to the remaining Bilateria—i.e., Nephrozoa PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5DYXJyYW56YTwvQXV0aG9yPjxZZWFyPjE5OTc8L1llYXI+
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ADDIN EN.CITE.DATA (Carranza et al. 1997; Ruiz-Trillo et al. 1999; Jondelius et al. 2002). Much more recently, and based on first-generation phylogenomic analyses, ADDIN EN.CITE <EndNote><Cite AuthorYear="1"><Author>Dunn</Author><Year>2008</Year><RecNum>7465</RecNum><DisplayText>Dunn et al. (2008)</DisplayText><record><rec-number>7465</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907275">7465</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Dunn, C. W.</author><author>Hejnol, A.</author><author>Matus, D. Q.</author><author>Pang, K.</author><author>Browne, W. E.</author><author>Smith, S. A.</author><author>Seaver, E. C.</author><author>Rouse, G. W.</author><author>Obst, M.</author><author>Edgecombe, G. D.</author><author>S?rensen, M. V.</author><author>Haddock, S. H. D.</author><author>Schmidt-Rhaesa, A.</author><author>Okusu, A.</author><author>Kristensen, R. M.</author><author>Wheeler, W. C.</author><author>Martindale, M. Q.</author><author>Giribet, G.</author></authors></contributors><auth-address>Kewalo Marine Laboratory, PBRC, University of Hawaii, 41 Ahui Street, Honolulu, Hawaii 96813, USA. casey_dunn@brown.edu</auth-address><titles><title>Broad phylogenomic sampling improves resolution of the animal tree of life</title><secondary-title>Nature</secondary-title><alt-title>Nature</alt-title></titles><periodical><full-title>Nature</full-title></periodical><alt-periodical><full-title>Nature</full-title></alt-periodical><pages>745-749</pages><volume>452</volume><number>7188</number><keywords><keyword>AToL results</keyword><keyword>Xenoturbella</keyword></keywords><dates><year>2008</year><pub-dates><date>Submitted, September 2007</date></pub-dates></dates><isbn>1476-4687 (Electronic)
0028-0836 (Linking)</isbn><accession-num>18322464</accession-num><urls><related-urls><url> et al. (2008) shook the animal tree once more by placing Ctenophora as the sister group to all other animals, contradicting prior ideas about metazoan evolution and increasing complexity at the base of the tree. However, this result remains contentious PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5SeWFuPC9BdXRob3I+PFllYXI+MjAxMzwvWWVhcj48UmVj
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ADDIN EN.CITE.DATA (e.g., Nosenko et al. 2013; Ryan et al. 2013; Moroz et al. 2014; Halanych 2015; Whelan et al. 2015), unlike the abovementioned clades: Lophotrochozoa (and the more inclusive Spiralia), Ecdysozoa and Nephrozoa, which are found almost universally in recent phylogenomic analyses PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5IZWpub2w8L0F1dGhvcj48WWVhcj4yMDA5PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Hejnol et al. 2009; Nesnidal et al. 2010; Nesnidal et al. 2013; Struck et al. 2014; Laumer et al. 2015a). One notable exception to this consensus is a rather controversial study that claims to place Xenoturbellida and Acoelomorpha within Deuterostomia ADDIN EN.CITE <EndNote><Cite><Author>Philippe</Author><Year>2011</Year><RecNum>14578</RecNum><DisplayText>(Philippe et al. 2011)</DisplayText><record><rec-number>14578</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907337">14578</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Philippe, Hervé</author><author>Brinkmann, Henner</author><author>Copley, Richard R.</author><author>Moroz, Leonid L.</author><author>Nakano, Hiroaki</author><author>Poustka, Albert J.</author><author>Wallberg, Andreas</author><author>Peterson, Kevin J.</author><author>Telford, Maximilian J.</author></authors></contributors><titles><title><style face="normal" font="default" size="100%">Acoelomorph flatworms are deuterostomes related to </style><style face="italic" font="default" size="100%">Xenoturbella</style></title><secondary-title>Nature</secondary-title></titles><periodical><full-title>Nature</full-title></periodical><pages>255-258</pages><volume>470</volume><number>7333</number><dates><year>2011</year></dates><isbn>0028-0836
1476-4687</isbn><urls></urls><electronic-resource-num>10.1038/nature09676</electronic-resource-num></record></Cite></EndNote>(Philippe et al. 2011).While a consensus is emerging with respect to the relationships and composition of these major clades (Halanych 2004; Giribet 2008; Edgecombe et al. 2011; Dunn et al. 2014, a few aspects remain uncertain (see Fig. 1). These include mostly the internal relationships of Ecdysozoa, Spiralia, and Lophotrochozoa, and the base of the animal tree PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5FZGdlY29tYmU8L0F1dGhvcj48WWVhcj4yMDExPC9ZZWFy
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ADDIN EN.CITE.DATA )—specifically, the relative position of Porifera or Ctenophora as the sister group to all remaining Metazoa, a topic that remains contentious due to data and model dependence PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5XaGVsYW48L0F1dGhvcj48WWVhcj4yMDE1PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (e.g., Pick et al. 2010; Nosenko et al. 2013; Whelan et al. 2015). In this review the progress and future challenges to reconstruct the animal tree of life are discussed.Animal phylogenomics—or the use of large data sets to infer animal phylogeniesThe term phylogenomics, which originally had a different meaning ADDIN EN.CITE <EndNote><Cite><Author>Eisen</Author><Year>2003</Year><RecNum>16680</RecNum><DisplayText>(Eisen and Fraser 2003)</DisplayText><record><rec-number>16680</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907365">16680</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Eisen, Jonathan A.</author><author>Fraser, Claire M.</author></authors></contributors><auth-address>Institute for Genomic Research, Rockville, MD 20850, USA. jeisen@</auth-address><titles><title>Phylogenomics: Intersection of evolution and genomics</title><secondary-title>Science</secondary-title><alt-title>Science</alt-title></titles><periodical><full-title>Science</full-title></periodical><alt-periodical><full-title>Science</full-title></alt-periodical><pages>1706-1707</pages><volume>300</volume><number>5626</number><keywords><keyword>Animals</keyword><keyword>Bacteria/genetics</keyword><keyword>*Biological Evolution</keyword><keyword>Computational Biology</keyword><keyword>*Evolution, Molecular</keyword><keyword>Gene Transfer, Horizontal</keyword><keyword>Genome</keyword><keyword>Genome, Human</keyword><keyword>*Genomics</keyword><keyword>Humans</keyword><keyword>*Phylogeny</keyword><keyword>Sequence Analysis, DNA</keyword></keywords><dates><year>2003</year><pub-dates><date>Jun 13</date></pub-dates></dates><isbn>1095-9203 (Electronic)
0036-8075 (Linking)</isbn><accession-num>12805538</accession-num><urls><related-urls><url>;(Eisen and Fraser 2003), has become widespread and now refers to phylogenetic analyses using large amounts of genetic data. It is often associated with the use of data derived from methods that sequence blindly, as opposed to the more traditional PCR-based target sequencing (often using Sanger sequencing). The field has exploded with the generalized use of massive parallel sequencing methods (often called “next generation sequencing” or “NGS”). However, different authors give different meanings to the word phylogenomics, with some using it to refer exclusively to the use of whole-genome data to infer phylogenies ADDIN EN.CITE <EndNote><Cite><Author>Dopazo</Author><Year>2004</Year><RecNum>7312</RecNum><DisplayText>(Dopazo et al. 2004)</DisplayText><record><rec-number>7312</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907274">7312</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Dopazo, H.</author><author>Santoyo, J.</author><author>Dopazo, J.</author></authors></contributors><auth-address>Bioinformatics Unit. Biotechnology Programme, Centro Nacional de Investigaciones Oncologicas (CNIO), Melchor Fernandez Almagro, 3, 28029, Madrid, Spain.</auth-address><titles><title>Phylogenomics and the number of characters required for obtaining an accurate phylogeny of eukaryote model species</title><secondary-title>Bioinformatics</secondary-title></titles><periodical><full-title>Bioinformatics</full-title></periodical><pages>I116-I121</pages><volume>20 Suppl 1</volume><dates><year>2004</year><pub-dates><date>Aug 4</date></pub-dates></dates><accession-num>15262789</accession-num><urls><related-urls><url><style face="underline" font="default" size="100%">;(Dopazo et al. 2004), while now it is mostly used in the sense described above, yet some target gene approaches rightfully qualify as “phylogenomic” ADDIN EN.CITE <EndNote><Cite><Author>Regier</Author><Year>2010</Year><RecNum>15303</RecNum><Prefix>e.g.`, </Prefix><DisplayText>(e.g., Regier et al. 2010)</DisplayText><record><rec-number>15303</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907348">15303</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Regier, J. C.</author><author>Shultz, J. W.</author><author>Zwick, A.</author><author>Hussey, A.</author><author>Ball, B.</author><author>Wetzer, R.</author><author>Martin, J. W.</author><author>Cunningham, C. W.</author></authors></contributors><auth-address>Center for Biosystems Research, University of Maryland Biotechnology Institute.</auth-address><titles><title>Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences</title><secondary-title>Nature</secondary-title></titles><periodical><full-title>Nature</full-title></periodical><pages>1079-1083</pages><volume>463</volume><edition>2010/02/12</edition><dates><year>2010</year><pub-dates><date>Feb 10</date></pub-dates></dates><isbn>1476-4687 (Electronic)
0028-0836 (Linking)</isbn><accession-num>20147900</accession-num><urls><related-urls><url>;(e.g., Regier et al. 2010). The term phylotranscriptomics has also been applied ADDIN EN.CITE <EndNote><Cite><Author>Oakley</Author><Year>2013</Year><RecNum>13995</RecNum><DisplayText>(Oakley et al. 2013)</DisplayText><record><rec-number>13995</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907329">13995</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Oakley, T. H.</author><author>Wolfe, J. M.</author><author>Lindgren, A. R.</author><author>Zaharoff, A. K.</author></authors></contributors><auth-address>Department of Ecology Evolution and Marine Biology, University of California, Santa Barbara.</auth-address><titles><title>Phylotranscriptomics to bring the understudied into the fold: monophyletic Ostracoda, fossil placement, and pancrustacean phylogeny</title><secondary-title>Molecular Biology and Evolution</secondary-title><alt-title>Molecular biology and evolution</alt-title></titles><periodical><full-title>Molecular Biology and Evolution</full-title></periodical><alt-periodical><full-title>Molecular Biology and Evolution</full-title></alt-periodical><pages>215-233</pages><volume>30</volume><number>1</number><edition>2012/09/15</edition><dates><year>2013</year><pub-dates><date>Oct 13</date></pub-dates></dates><isbn>1537-1719 (Electronic)
0737-4038 (Linking)</isbn><accession-num>22977117</accession-num><urls><related-urls><url>;(Oakley et al. 2013) to refer to phylogenetic analyses using transcriptomic data as the major source of protein-encoding genes.For animal phylogeny, early phylogenomic approaches made use of the genomes of a few model organisms complemented with EST (Expressed Sequence Tags) sequencing using Sanger-based methods PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5NYXJsw6l0YXo8L0F1dGhvcj48WWVhcj4yMDA2PC9ZZWFy
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ADDIN EN.CITE.DATA (e.g., Philippe et al. 2005; Delsuc et al. 2006; Marlétaz et al. 2006; Hausdorf et al. 2007; Philippe et al. 2007; Dunn et al. 2008; Helmkampf et al. 2008; Hejnol et al. 2009; Philippe et al. 2009). These early analyses were followed by a series of papers that incorporated 454 sequence data to the previous data sets, but concentrated into animal subclades. In fact, the only attempts to evaluate animal phylogeny as a whole (including most animal phyla) are still based mostly on ESTs PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5EdW5uPC9BdXRob3I+PFllYXI+MjAwODwvWWVhcj48UmVj
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ADDIN EN.CITE.DATA (Laumer et al. 2015a). It is now clear that Platyhelminthes do not group with the original Lophotrochozoa, but neither do they constitute the proposed clade Platyzoa PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5HaXJpYmV0PC9BdXRob3I+PFllYXI+MjAwMDwvWWVhcj48
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ADDIN EN.CITE.DATA (Cavalier-Smith 1998; Giribet et al. 2000). “Platyzoa” is now understood as a grade of two clades, Gnathifera and Rouphozoa, the latter constituting the sister group to Lophotrochozoa PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5MYXVtZXI8L0F1dGhvcj48WWVhcj4yMDE1PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Struck et al. 2014; Laumer et al. 2015a). The monophyly of the classical clade Lophophorata is likewise emerging in recent analyses PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5OZXNuaWRhbDwvQXV0aG9yPjxZZWFyPjIwMTM8L1llYXI+
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ADDIN EN.CITE.DATA (Nesnidal et al. 2013; Laumer et al. 2015a). However, a few aspects of spiralian phylogeny remain unresolved, such as the position of Cycliophora PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5MYXVtZXI8L0F1dGhvcj48WWVhcj4yMDE1PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Laumer et al. 2015a), a group difficult to assess molecularly as well as morphologically PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5OZXZlczwvQXV0aG9yPjxZZWFyPjIwMDk8L1llYXI+PFJl
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ADDIN EN.CITE.DATA (e.g., Neves et al. 2009).Major progress has also been made in the internal phylogeny of traditionally difficult phyla, like Annelida PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5MYXVtZXI8L0F1dGhvcj48WWVhcj4yMDE1PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Struck et al. 2011; Weigert et al. 2014; Andrade et al. 2015; Laumer et al. 2015a; Lemer et al. 2015), Mollusca PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5TbWl0aDwvQXV0aG9yPjxZZWFyPjIwMTE8L1llYXI+PFJl
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ADDIN EN.CITE.DATA (Kocot et al. 2011; Smith et al. 2011; Kocot et al. 2013; Zapata et al. 2014; González et al. 2015) and Platyhelminthes PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5MYXVtZXI8L0F1dGhvcj48WWVhcj4yMDE1PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Egger et al. 2015; Laumer et al. 2015b). Phylogenomics has provided the datasets that contain enough information to find resolution and support for some of the most recalcitrant clades in animal evolution. For instance, it is now well accepted that Annelida includes many taxa formerly considered different phyla or with supposed affiliations with other animal groups, such as Sipuncula, Echiura, Pogonophora and Vestimentifera, Myzostomida, or Diurodrilida PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5BbmRyYWRlPC9BdXRob3I+PFllYXI+MjAxNTwvWWVhcj48
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ADDIN EN.CITE.DATA (Struck et al. 2011; Kvist and Siddall 2013; Weigert et al. 2014; Andrade et al. 2015; Laumer et al. 2015a). As the costs of generating molecular sequence data decrease and the techniques for obtaining RNA and DNA become more sensitive, progress is now being made in many new directions, from the inclusion of previously impracticable small samples PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5BbmRyYWRlPC9BdXRob3I+PFllYXI+MjAxNTwvWWVhcj48
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ADDIN EN.CITE.DATA (e.g., Andrade et al. 2015; Laumer et al. 2015b) to applying phylogenomic approaches to much more recent divergences, including species problems, relationships within families, or among families, etc. Future challengesResolving outstanding issues in animal phylogeny will require a concerted effort to gather genomic data from key taxa PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Mb3BlejwvQXV0aG9yPjxZZWFyPjIwMTQ8L1llYXI+PFJl
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ADDIN EN.CITE.DATA (Zhang et al. 2014), to provide two examples. In addition, computational development of software and hardware will become more limiting than generating the genomic data per se. Storing the vast amounts of raw and processed genomic data is also emerging as a critical issue. If our goal is to provide a well-resolved animal phylogeny, several areas must thus move forward:1. Taxon samplingAlthough the first phylogenomic analyses made use of the available genomes, those studies generated little data ADDIN EN.CITE <EndNote><Cite><Author>Philippe</Author><Year>2005</Year><RecNum>14583</RecNum><Prefix>e.g.`, </Prefix><DisplayText>(e.g., Philippe et al. 2005)</DisplayText><record><rec-number>14583</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907337">14583</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Philippe, H.</author><author>Lartillot, N.</author><author>Brinkmann, H.</author></authors></contributors><auth-address>Canadian Institute for Advanced Research. Departement de Biochimie, Universite de Montreal, Succursale Centre-Ville, Montreal, Quebec H3C3J7, Canada.</auth-address><titles><title>Multigene analyses of bilaterian animals corroborate the monophyly of Ecdysozoa, Lophotrochozoa and Protostomia</title><secondary-title>Molecular Biology and Evolution</secondary-title></titles><periodical><full-title>Molecular Biology and Evolution</full-title></periodical><pages>1246-1253</pages><volume>22</volume><number>5</number><dates><year>2005</year><pub-dates><date>Feb 9</date></pub-dates></dates><accession-num>15703236</accession-num><urls><related-urls><url><style face="underline" font="default" size="100%">;(e.g., Philippe et al. 2005). Taxon sampling was later emphasized in subsequent EST analyses PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5EdW5uPC9BdXRob3I+PFllYXI+MjAwODwvWWVhcj48UmVj
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ADDIN EN.CITE.DATA (e.g., Kocot et al. 2011; Smith et al. 2011; Struck et al. 2011; Kvist and Siddall 2013; Andrade et al. 2014; Cannon et al. 2014; Fernández et al. 2014; Sharma and Wheeler 2014; Telford et al. 2014; von Reumont and Wa?gele 2014; Weigert et al. 2014; Zapata et al. 2014; Andrade et al. 2015; Egger et al. 2015; Laumer et al. 2015b). A series of studies has focused on particular nodes of the animal tree of life PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5MYXVtZXI8L0F1dGhvcj48WWVhcj4yMDE1PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Nesnidal et al. 2013; Nosenko et al. 2013; Ryan et al. 2013; Moroz et al. 2014; Srivastava et al. 2014; Struck et al. 2014; Laumer et al. 2015a). However, no metazoan-broad studies using NGS data have included data from all or most animal phyla; and only through thorough taxon sampling will we be able to resolve the position of the last truly enigmatic taxa, such as cycliophorans, dicyemids and rhombozoans PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5MYXVtZXI8L0F1dGhvcj48WWVhcj4yMDE1PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (Laumer et al. 2015a; Laumer et al. 2015b), emphasis will have to shift towards optimizing fieldwork for “rare” animals. This will require taxonomic expertise as perhaps one of the most crucial components for the future of phylogenomics.2. Data matricesData matrix assembly is a fundamental step in phylogenomic analyses, and different methods have been employed, from “manually” curating matrices with a few ill-defined, pre-selected genes ADDIN EN.CITE <EndNote><Cite><Author>Delsuc</Author><Year>2006</Year><RecNum>7025</RecNum><Prefix>e.g.`, </Prefix><DisplayText>(e.g., Delsuc et al. 2006)</DisplayText><record><rec-number>7025</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907271">7025</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Delsuc, F.</author><author>Brinkmann, H.</author><author>Chourrout, D.</author><author>Philippe, H.</author></authors></contributors><auth-address>Departement de Biochimie, Centre Robert-Cedergren, Universite de Montreal, Succursale Centre-Ville, Montreal, Quebec H3C3J7, Canada.</auth-address><titles><title>Tunicates and not cephalochordates are the closest living relatives of vertebrates</title><secondary-title>Nature</secondary-title></titles><periodical><full-title>Nature</full-title></periodical><pages>965-968</pages><volume>439</volume><number>7079</number><keywords><keyword>Animals</keyword><keyword>Evolution, Molecular</keyword><keyword>Likelihood Functions</keyword><keyword>*Phylogeny</keyword><keyword>Research Support, Non-U.S. Gov't</keyword><keyword>Urochordata/*classification/genetics/physiology</keyword><keyword>Vertebrates/*classification/genetics</keyword></keywords><dates><year>2006</year><pub-dates><date>Feb 23</date></pub-dates></dates><accession-num>16495997</accession-num><urls><related-urls><url><style face="underline" font="default" size="100%"> </style></url></related-urls></urls></record></Cite></EndNote>(e.g., Delsuc et al. 2006), to automated methods of orthology detection and gene selection PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Lb2NvdDwvQXV0aG9yPjxZZWFyPjIwMTE8L1llYXI+PFJl
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ADDIN EN.CITE.DATA (e.g., Dunn et al. 2008; Kocot et al. 2011). Automated methods for generating a data matrix—manual methods are mostly ignored in modern evolutionary biology—can be divided into three steps, each crucial in its own way: (1) gene prediction, (2) orthology assignment, and (3) data set assembly. Gene prediction methods (“assemblers”) are numerous and often must deal with common issues such as paralogy and alternative splicing, and cannot be dealt with here due to space limitations. Numerous assemblers, their computation efficiency and limitations are discussed and compared elsewhere PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5FYXJsPC9BdXRob3I+PFllYXI+MjAxMTwvWWVhcj48UmVj
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ADDIN EN.CITE.DATA (e.g., Earl et al. 2011; Bradnam et al. 2013).Orthology assignment can rely on several methods designed to perform some sort of all-by-all comparisons. These can range from pairwise comparisons of distances to graphic methods. Some common orthology assignment methods used in phylogenomics are HaMStR ADDIN EN.CITE <EndNote><Cite><Author>Ebersberger</Author><Year>2009</Year><RecNum>7599</RecNum><DisplayText>(Ebersberger et al. 2009)</DisplayText><record><rec-number>7599</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907277">7599</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Ebersberger, I.</author><author>Strauss, S.</author><author>von Haeseler, A.</author></authors></contributors><auth-address>Center for Integrative Bioinformatics Vienna, Max F, Perutz Laboratories, Vienna, Austria. ingo.ebersberger@univie.ac.at</auth-address><titles><title>HaMStR: Profile hidden markov model based search for orthologs in ESTs</title><secondary-title>BMC Evolutionary Biology</secondary-title><alt-title>BMC evolutionary biology</alt-title></titles><periodical><full-title>BMC Evolutionary Biology</full-title></periodical><alt-periodical><full-title>BMC Evolutionary Biology</full-title></alt-periodical><pages>157</pages><volume>9</volume><edition>2009/07/10</edition><keywords><keyword>Algorithms</keyword><keyword>Animals</keyword><keyword>Cluster Analysis</keyword><keyword>*Expressed Sequence Tags</keyword><keyword>Fungi/genetics</keyword><keyword>Genome, Fungal</keyword><keyword>Genomics/*methods</keyword><keyword>Humans</keyword><keyword>Markov Chains</keyword><keyword>*Models, Genetic</keyword><keyword>*Phylogeny</keyword><keyword>Sequence Analysis, DNA</keyword><keyword>Software</keyword></keywords><dates><year>2009</year></dates><isbn>1471-2148 (Electronic)
1471-2148 (Linking)</isbn><accession-num>19586527</accession-num><urls><related-urls><url>;(Ebersberger et al. 2009), OMA ADDIN EN.CITE <EndNote><Cite><Author>Altenhoff</Author><Year>2011</Year><RecNum>4219</RecNum><DisplayText>(Altenhoff et al. 2011)</DisplayText><record><rec-number>4219</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907193">4219</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Altenhoff, A. M.</author><author>Schneider, A.</author><author>Gonnet, G. H.</author><author>Dessimoz, C.</author></authors></contributors><auth-address>Dessimoz, C
ETH, Univ Str 6, CH-8092 Zurich, Switzerland
ETH, Univ Str 6, CH-8092 Zurich, Switzerland
ETH, CH-8092 Zurich, Switzerland
Swiss Inst Bioinformat, CH-8092 Zurich, Switzerland
Univ Edinburgh, Inst Evolutionary Biol, Edinburgh EH9 3JT, Midlothian, Scotland</auth-address><titles><title>OMA 2011: orthology inference among 1000 complete genomes</title><secondary-title>Nucleic Acids Research</secondary-title><alt-title>Nucleic Acids Res</alt-title></titles><periodical><full-title>Nucleic Acids Research</full-title></periodical><alt-periodical><full-title>Nucleic Acids Res</full-title><abbr-1>Nucleic acids research</abbr-1></alt-periodical><pages>D289-D294</pages><volume>39</volume><keywords><keyword>biotechnology-information</keyword><keyword>database resources</keyword><keyword>national-center</keyword><keyword>identification</keyword><keyword>browser</keyword></keywords><dates><year>2011</year><pub-dates><date>Jan</date></pub-dates></dates><isbn>0305-1048</isbn><accession-num>WOS:000285831700049</accession-num><urls><related-urls><url><Go to ISI>://WOS:000285831700049</url><url>;(Altenhoff et al. 2011) or the All-By-All BLASTP search, followed by a phylogenetic approach to identify orthologous sequences ADDIN EN.CITE <EndNote><Cite><Author>Smith</Author><Year>2011</Year><RecNum>3211</RecNum><DisplayText>(Smith et al. 2011)</DisplayText><record><rec-number>3211</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907184">3211</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Smith, S.</author><author>Wilson, N. G.</author><author>Goetz, F.</author><author>Feehery, C.</author><author>Andrade, S. C. S.</author><author>Rouse, G. W.</author><author>Giribet, G.</author><author>Dunn, C. W.</author></authors></contributors><titles><title>Resolving the evolutionary relationships of molluscs with phylogenomic tools</title><secondary-title>Nature</secondary-title></titles><periodical><full-title>Nature</full-title></periodical><pages>364-367</pages><volume>480</volume><keywords><keyword>NSF #0844881</keyword></keywords><dates><year>2011</year><pub-dates><date>submitted, 7 April 2011
Re-submitted, 10 June 2011
re-submitted, 20 July 2011</date></pub-dates></dates><orig-pub>Published online 26 October 2011</orig-pub><urls></urls><electronic-resource-num>10.1038/nature10526</electronic-resource-num></record></Cite></EndNote>(Smith et al. 2011), as implemented in Agalma ADDIN EN.CITE <EndNote><Cite><Author>Dunn</Author><Year>2013</Year><RecNum>7466</RecNum><DisplayText>(Dunn et al. 2013)</DisplayText><record><rec-number>7466</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907275">7466</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Dunn, Casey W.</author><author>Howison, Mark</author><author>Zapata, Felipe</author></authors></contributors><auth-address>Department of Ecology and Evolutionary Biology, Brown University, Providence, Rhode Island, USA. casey_dunn@brown.edu.</auth-address><titles><title>Agalma: an automated phylogenomics workflow</title><secondary-title>BMC Bioinformatics</secondary-title><alt-title>BMC bioinformatics</alt-title></titles><periodical><full-title>BMC Bioinformatics</full-title></periodical><alt-periodical><full-title>BMC Bioinformatics</full-title></alt-periodical><pages>330</pages><volume>14</volume><keywords><keyword>Gene Expression Profiling/*methods</keyword><keyword>Genome</keyword><keyword>Genomics/*methods</keyword><keyword>*Phylogeny</keyword><keyword>Sequence Analysis, DNA/methods</keyword><keyword>*Software</keyword></keywords><dates><year>2013</year></dates><isbn>1471-2105 (Electronic)
1471-2105 (Linking)</isbn><accession-num>24252138</accession-num><urls><related-urls><url>;(Dunn et al. 2013). Few phylogenomic studies have studied the impact of alternative methods for orthology assignment on phylogenetic estimation, but the few that have show that results often converge when enough information becomes available ADDIN EN.CITE <EndNote><Cite><Author>Zapata</Author><Year>2014</Year><RecNum>21</RecNum><Prefix>e.g.`, </Prefix><DisplayText>(e.g., Zapata et al. 2014)</DisplayText><record><rec-number>21</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907156">21</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author> Zapata, Felipe</author><author>Wilson, Nerida G.</author><author>Howison, Mark</author><author>Andrade, Sónia C. S.</author><author>J?rger, Katharina M.</author><author>Schr?dl, M.</author><author>Goetz, Freya E.</author><author>Giribet, Gonzalo</author><author>Dunn, Casey W.</author></authors></contributors><titles><title>Phylogenomic analyses of deep gastropod relationships reject Orthogastropoda</title><secondary-title>Proceedings of the Royal Society B: Biological Sciences</secondary-title></titles><periodical><full-title>Proceedings of the Royal Society B: Biological Sciences</full-title></periodical><pages>20141739</pages><volume>281</volume><dates><year>2014</year><pub-dates><date>submitted, 13 July 2014</date></pub-dates></dates><urls><related-urls><url>;(e.g., Zapata et al. 2014).Once each gene prediction (often called orthogroups) has been assigned into orthology groups (often referred to as orthoclusters), a decision must be made about which of these orthoclusters are selected. The process of gene selection can be done by choosing each orthocluster found above a certain threshold (i.e., in ≥ 50% of taxa). Such methods, however, can be affected by key—but poorly sampled—taxa, and some have used methods for optimizing the representation of genes in poor libraries ADDIN EN.CITE <EndNote><Cite><Author>Laumer</Author><Year>2015</Year><RecNum>11646</RecNum><DisplayText>(Laumer et al. 2015b)</DisplayText><record><rec-number>11646</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907310">11646</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Laumer, Christopher E.</author><author>Hejnol, Andreas</author><author>Giribet, Gonzalo</author></authors></contributors><auth-address>Museum of Comparative Zoology, Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, United States.
Sars International Centre for Marine Molecular Biology, University of Bergen, Bergen, Norway.</auth-address><titles><title>Nuclear genomic signals of the "microturbellarian" roots of platyhelminth evolutionary innovation</title><secondary-title>eLife</secondary-title><alt-title>eLife</alt-title></titles><periodical><full-title>eLife</full-title><abbr-1>eLife</abbr-1></periodical><alt-periodical><full-title>eLife</full-title><abbr-1>eLife</abbr-1></alt-periodical><pages>e05503</pages><volume>4</volume><dates><year>2015</year><pub-dates><date>Submitted, 5 November 2014</date></pub-dates></dates><isbn>2050-084X (Electronic)
2050-084X (Linking)</isbn><accession-num>25764302</accession-num><urls><related-urls><url>;(Laumer et al. 2015b). In addition to these criteria based on percentage of occupancy allowed into the final matrix ADDIN EN.CITE <EndNote><Cite><Author>Hejnol</Author><Year>2009</Year><RecNum>9799</RecNum><DisplayText>(Hejnol et al. 2009)</DisplayText><record><rec-number>9799</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907296">9799</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Hejnol, A.</author><author>Obst, M.</author><author>Stamatakis, A.</author><author>Ott. M.</author><author>Rouse, G. W.</author><author>Edgecombe, G. D.</author><author>Martinez, P.</author><author>Bagu?à, J.</author><author>Bailly, X.</author><author>Jondelius, U.</author><author>Wiens, M.</author><author>Müller, W. E. G.</author><author>Seaver, E.</author><author>Wheeler, W. C.</author><author>Martindale, M. Q.</author><author>Giribet, G.</author><author>Dunn, C. W.</author></authors></contributors><titles><title>Assessing the root of bilaterian animals with scalable phylogenomic methods</title><secondary-title>Proceedings of the Royal Society B: Biological Sciences</secondary-title></titles><periodical><full-title>Proceedings of the Royal Society B: Biological Sciences</full-title></periodical><pages>4261-4270</pages><volume>276</volume><keywords><keyword>AToL results</keyword><keyword>Cycliophora</keyword></keywords><dates><year>2009</year><pub-dates><date>Submitted, 30 March 2009
Resubmitted, April 2009
Resubmitted, May 2009</date></pub-dates></dates><urls><related-urls><url>;(Hejnol et al. 2009), criteria based on data properties have been used for designing the final data set. One such approach is BaCoCa (BAse COmposition CAlculator), which combines multiple statistical approaches to identify biases in aligned sequence data ADDIN EN.CITE <EndNote><Cite><Author>Kück</Author><Year>2014</Year><RecNum>11402</RecNum><DisplayText>(Kück and Struck 2014)</DisplayText><record><rec-number>11402</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907308">11402</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Kück, P.</author><author>Struck, T. H.</author></authors></contributors><auth-address>Zoological Research Museum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany. Electronic address: patrick_kueck@web.de.</auth-address><titles><title>BaCoCa – A heuristic software tool for the parallel assessment of sequence biases in hundreds of gene and taxon partitions</title><secondary-title>Molecular Phylogenetics and Evolution</secondary-title><alt-title>Molecular phylogenetics and evolution</alt-title></titles><periodical><full-title>Molecular Phylogenetics and Evolution</full-title><abbr-1>Mol. Phylogenet. Evol.</abbr-1></periodical><alt-periodical><full-title>Molecular Phylogenetics and Evolution</full-title><abbr-1>Mol. Phylogenet. Evol.</abbr-1></alt-periodical><pages>94-98</pages><volume>70</volume><dates><year>2014</year><pub-dates><date>Jan</date></pub-dates></dates><isbn>1095-9513 (Electronic)
1055-7903 (Linking)</isbn><accession-num>24076250</accession-num><urls><related-urls><url>;(Kück and Struck 2014). Others have used methods to select sets of genes with different evolutionary rates ADDIN EN.CITE <EndNote><Cite><Author>Fernández</Author><Year>2014</Year><RecNum>8109</RecNum><DisplayText>(Fernández et al. 2014)</DisplayText><record><rec-number>8109</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907281">8109</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Fernández, Rosa</author><author>Hormiga, Gustavo</author><author>Giribet, Gonzalo</author></authors></contributors><auth-address>Museum of Comparative Zoology and Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA 02138, USA. Electronic address: rfernandezgarcia@g.harvard.edu.
Department of Biological Sciences, The George Washington University, Washington, D.C. 20052, USA.
Museum of Comparative Zoology and Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA 02138, USA.</auth-address><titles><title>Phylogenomic analysis of spiders reveals nonmonophyly of orb weavers</title><secondary-title>Current Biology</secondary-title><alt-title>Current biology : CB</alt-title></titles><periodical><full-title>Current Biology</full-title></periodical><alt-periodical><full-title>Curr Biol</full-title><abbr-1>Current biology : CB</abbr-1></alt-periodical><pages>1772-1777</pages><volume>24</volume><number>15</number><dates><year>2014</year><pub-dates><date>Submitted, 1 May 2014</date></pub-dates></dates><isbn>1879-0445 (Electronic)
0960-9822 (Linking)</isbn><accession-num>25042584</accession-num><urls><related-urls><url>;(Fernández et al. 2014), added bins of genes with increasing evolutionary rates ADDIN EN.CITE <EndNote><Cite><Author>Sharma</Author><Year>2014</Year><RecNum>2788</RecNum><DisplayText>(Sharma and Wheeler 2014)</DisplayText><record><rec-number>2788</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907181">2788</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Sharma, Prashant P.</author><author>Wheeler, Ward C.</author></authors></contributors><titles><title>Cross-bracing uncalibrated nodes in molecular dating improves congruence of fossil and molecular age estimates</title><secondary-title>Frontiers in Zoology</secondary-title></titles><periodical><full-title>Frontiers in Zoology</full-title></periodical><pages>57</pages><volume>11</volume><number>1</number><dates><year>2014</year></dates><isbn>1742-9994</isbn><urls><related-urls><url>;(Sharma and Wheeler 2014), or used measures such as phylogenetic informativeness to select sets of genes PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Mw7NwZXotR2lyw6FsZGV6PC9BdXRob3I+PFllYXI+MjAx
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ADDIN EN.CITE.DATA (López-Giráldez et al. 2013). Each of these strategies may drive results according to the properties of the selected sets of genes.3. Algorithmic developmentsAlgorithmic developments are obviously important in phylogenomic analyses, and these may apply to any step of the phylogenetic analysis. An interesting study actually quantified the computational effort invested in the different steps leading to a phylogenomic analysis, from transcriptome assembly to phylogenetic tree inference ADDIN EN.CITE <EndNote><Cite><Author>Dunn</Author><Year>2013</Year><RecNum>7466</RecNum><DisplayText>(Dunn et al. 2013)</DisplayText><record><rec-number>7466</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907275">7466</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Dunn, Casey W.</author><author>Howison, Mark</author><author>Zapata, Felipe</author></authors></contributors><auth-address>Department of Ecology and Evolutionary Biology, Brown University, Providence, Rhode Island, USA. casey_dunn@brown.edu.</auth-address><titles><title>Agalma: an automated phylogenomics workflow</title><secondary-title>BMC Bioinformatics</secondary-title><alt-title>BMC bioinformatics</alt-title></titles><periodical><full-title>BMC Bioinformatics</full-title></periodical><alt-periodical><full-title>BMC Bioinformatics</full-title></alt-periodical><pages>330</pages><volume>14</volume><keywords><keyword>Gene Expression Profiling/*methods</keyword><keyword>Genome</keyword><keyword>Genomics/*methods</keyword><keyword>*Phylogeny</keyword><keyword>Sequence Analysis, DNA/methods</keyword><keyword>*Software</keyword></keywords><dates><year>2013</year></dates><isbn>1471-2105 (Electronic)
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ADDIN EN.CITE.DATA (Lartillot et al. 2013), often does not converge on large data sets ADDIN EN.CITE <EndNote><Cite><Author>Fernández</Author><Year>2014</Year><RecNum>8109</RecNum><Prefix>e.g.`, </Prefix><DisplayText>(e.g., Fernández et al. 2014)</DisplayText><record><rec-number>8109</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907281">8109</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Fernández, Rosa</author><author>Hormiga, Gustavo</author><author>Giribet, Gonzalo</author></authors></contributors><auth-address>Museum of Comparative Zoology and Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA 02138, USA. Electronic address: rfernandezgarcia@g.harvard.edu.
Department of Biological Sciences, The George Washington University, Washington, D.C. 20052, USA.
Museum of Comparative Zoology and Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA 02138, USA.</auth-address><titles><title>Phylogenomic analysis of spiders reveals nonmonophyly of orb weavers</title><secondary-title>Current Biology</secondary-title><alt-title>Current biology : CB</alt-title></titles><periodical><full-title>Current Biology</full-title></periodical><alt-periodical><full-title>Curr Biol</full-title><abbr-1>Current biology : CB</abbr-1></alt-periodical><pages>1772-1777</pages><volume>24</volume><number>15</number><dates><year>2014</year><pub-dates><date>Submitted, 1 May 2014</date></pub-dates></dates><isbn>1879-0445 (Electronic)
0960-9822 (Linking)</isbn><accession-num>25042584</accession-num><urls><related-urls><url>;(e.g., Fernández et al. 2014). In fact, increased complexity in evolutionary models, often required for dealing with heterogeneous genomic and phylogenomic datasets, comes at a high computational cost. Faster and more efficient algorithms thus need to be developed to cope with the ever-increasing size of datasets ADDIN EN.CITE <EndNote><Cite><Author>Stamatakis</Author><Year>2014</Year><RecNum>3513</RecNum><DisplayText>(Stamatakis 2014a, 2014b)</DisplayText><record><rec-number>3513</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907187">3513</key></foreign-keys><ref-type name="Computer Program">9</ref-type><contributors><authors><author>Stamatakis, A.</author></authors></contributors><titles><title>ExaBayes User's Manual</title></titles><dates><year>2014</year></dates><urls></urls></record></Cite><Cite><Author>Stamatakis</Author><Year>2014</Year><RecNum>3512</RecNum><record><rec-number>3512</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907187">3512</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Stamatakis, A.</author></authors></contributors><auth-address>Scientific Computing Group, Heidelberg Institute for Theoretical Studies, Heidelberg, Germany.</auth-address><titles><title>RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies</title><secondary-title>Bioinformatics</secondary-title><alt-title>Bioinformatics</alt-title></titles><periodical><full-title>Bioinformatics</full-title></periodical><alt-periodical><full-title>Bioinformatics</full-title></alt-periodical><dates><year>2014</year><pub-dates><date>Jan 21</date></pub-dates></dates><isbn>1367-4811 (Electronic)
1367-4803 (Linking)</isbn><accession-num>24451623</accession-num><urls><related-urls><url>;(Stamatakis 2014a, 2014b). For phylogenetic inference, fast and accurate methods, like parsimony, may be able to provide a quick tree hypothesis without the burden of other methods ADDIN EN.CITE <EndNote><Cite><Author>Kvist</Author><Year>2013</Year><RecNum>11505</RecNum><DisplayText>(Kvist and Siddall 2013)</DisplayText><record><rec-number>11505</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907309">11505</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Kvist, Sebastian</author><author>Siddall, Mark E.</author></authors></contributors><titles><title>Phylogenomics of Annelida revisited: a cladistic approach using genome-wide expressed sequence tag data mining and examining the effects of missing data</title><secondary-title>Cladistics</secondary-title></titles><periodical><full-title>Cladistics</full-title></periodical><pages>435-448</pages><volume>29</volume><number>4</number><dates><year>2013</year></dates><isbn>07483007</isbn><urls><related-urls><url>;(Kvist and Siddall 2013), but parsimony software needs to incorporate realistic amino acid transition matrices before they can be generally applied to large phylogenomic analyses.4. Data storageThe amounts of data generated for phylogenetic analyses have grown exponentially in the past few years, to the point of being now considered part of the “Big Data science”, and these are just going to get much bigger PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5TdGVwaGVuczwvQXV0aG9yPjxZZWFyPjIwMTU8L1llYXI+
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ADDIN EN.CITE.DATA (Stephens et al. 2015). Although the genes utilized in each analysis per each terminal may only be in the range of a few hundreds or thousands, the raw data generated and all intermediate steps after sanitation, assembly, translation to amino acids, and orthogroup assignment, as well as tracking all connections between these steps, are orders of magnitude larger and require huge storage requirements. Some of these legacy data are often lost due to lack of storage resources, resulting in duplicated computational efforts through many of these steps, and some authors only publish the few genes utilized in the final analysis, violating the objective of data transparency (e.g., testing alternative assembly programs would not possible). While some public databases store both the raw data (e.g., Sequence Read Archive database [SRA] of NCBI?), assemblies, and data matrices (e.g., Dryad), additional effort is needed to connect the actual data to specimens ADDIN EN.CITE <EndNote><Cite><Author>Fernández</Author><Year>2015</Year><RecNum>16513</RecNum><Prefix>see for example </Prefix><DisplayText>(see for example Fernández and Giribet 2015)</DisplayText><record><rec-number>16513</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907362">16513</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Fernández, Rosa</author><author>Giribet, Gonzalo</author></authors></contributors><titles><title>Unnoticed in the tropics: phylogenomic resolution of the poorly known arachnid order Ricinulei (Arachnida)</title><secondary-title>Royal Society Open Science</secondary-title></titles><periodical><full-title>Royal Society Open Science</full-title></periodical><pages>150065</pages><volume>2</volume><number>6</number><dates><year>2015</year></dates><isbn>2054-5703</isbn><urls></urls><electronic-resource-num>10.1098/rsos.150065</electronic-resource-num></record></Cite></EndNote>(see for example Fernández and Giribet 2015). Although some global efforts attempt to do that PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Mb3BlejwvQXV0aG9yPjxZZWFyPjIwMTQ8L1llYXI+PFJl
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ADDIN EN.CITE.DATA (Giribet 2010; Giribet 2015; Pyron 2015; Wanninger 2015), especially under the phylogenomic paradigm. While it is clear that molecular data derived from hundreds of genes now often take precedence for reconstructing deep evolutionary histories, morphology still remains an important source of phylogenetic information ADDIN EN.CITE <EndNote><Cite><Author>Burleigh</Author><Year>2013</Year><RecNum>5546</RecNum><DisplayText>(Burleigh et al. 2013)</DisplayText><record><rec-number>5546</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907257">5546</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Burleigh, J. G.</author><author>Alphonse, K.</author><author>Alverson, A. J.</author><author>Bik, H. M.</author><author>Blank, C.</author><author>Cirranello, A. L.</author><author>Cui, H.</author><author>Daly, M.</author><author>Dietterich, T. G.</author><author>Gasparich, G.</author><author>Irvine, J.</author><author>Julius, M.</author><author>Kaufman, S.</author><author>Law, E.</author><author>Liu, J.</author><author>Moore, L.</author><author>O'Leary, M. A.</author><author>Passarotti, M.</author><author>Ranade, S.</author><author>Simmons, N. B.</author><author>Stevenson, D. W.</author><author>Thacker, R. W.</author><author>Theriot, E. C.</author><author>Todorovic, S.</author><author>Velazco, P. M.</author><author>Walls, R. L.</author><author>Wolfe, J. M.</author><author>Yu, M.</author></authors></contributors><auth-address>University of Florida.</auth-address><titles><title>Next-generation phenomics for the Tree of Life</title><secondary-title>PLoS Currents Tree of Life</secondary-title><alt-title>PLoS currents</alt-title></titles><periodical><full-title>PLoS Currents Tree of Life</full-title></periodical><alt-periodical><full-title>PLoS Currents</full-title><abbr-1>PLoS currents</abbr-1></alt-periodical><volume>5</volume><dates><year>2013</year></dates><isbn>2157-3999 (Electronic)</isbn><accession-num>23827969</accession-num><urls><related-urls><url>;(Burleigh et al. 2013) and the only means with which to place extinct lineages in a phylogenetic context PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Ob3ZhY2VrPC9BdXRob3I+PFllYXI+MTk5MjwvWWVhcj48
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ADDIN EN.CITE.DATA (Parham et al. 2012). Room for error in placement of fossils is generally much higher when fossils are assigned to nodes, and thus, an alternative using fossils as terminals in a combined analysis of molecules and morphology is now back in fashion PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5QeXJvbjwvQXV0aG9yPjxZZWFyPjIwMTE8L1llYXI+PFJl
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ADDIN EN.CITE.DATA (Murienne et al. 2010; Pyron 2011; Wood et al. 2013; Garwood et al. 2014; Sharma and Giribet 2014; Arcila et al. 2015)—this is often referred to as total evidence dating or tip-dating. (Combining fossils with molecules was more common in earlier total evidence analyses of animal relationships using Sanger data PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5XaGVlbGVyPC9BdXRob3I+PFllYXI+MTk5MzwvWWVhcj48
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ADDIN EN.CITE.DATA (Giribet 2015; Pyron 2015).Final RemarksIt is undisputed that molecules have taken a prominent role in reconstructing animal phylogeny and that a new consensus of animal relationships is emerging (see Fig. 1). The first generation of molecular phylogenetic analyses provided a refreshed framework for the animal tree, proposing key hypotheses such as Ecdysozoa, Spiralia and Lophotrochozoa. The current generation of phylogenomic analyses has explored and tested these hypotheses in great depth, has instructed new ones, especially with respect to the position of Ctenophora or Xenacoelomorpha, and the newest datasets are finally resolving recalcitrant nodes of the animal tree, like the internal relationships of Mollusca ADDIN EN.CITE <EndNote><Cite><Author>Sigwart</Author><Year>2015</Year><RecNum>16762</RecNum><Prefix>but see </Prefix><Suffix> for a discussion on alternative molluscan relationships</Suffix><DisplayText>(but see Sigwart and Lindberg 2015 for a discussion on alternative molluscan relationships)</DisplayText><record><rec-number>16762</rec-number><foreign-keys><key app="EN" db-id="trfzxzswofparuesv5b5rp0g0ar222tat9tz" timestamp="1436907366">16762</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Sigwart, Julia D.</author><author>Lindberg, David R.</author></authors></contributors><auth-address>Marine Laboratory, Queen's University Belfast, BT22 1PF, Northern Ireland, UK; and Department of Integrative Biology, Museum of Paleontology and Center for Computational Biology, University of California, Berkeley, CA, 94720, USA j.sigwart@qub.ac.uk.
Marine Laboratory, Queen's University Belfast, BT22 1PF, Northern Ireland, UK; and Department of Integrative Biology, Museum of Paleontology and Center for Computational Biology, University of California, Berkeley, CA, 94720, USA.</auth-address><titles><title>Consensus and confusion in molluscan trees: Evaluating morphological and molecular phylogenies</title><secondary-title>Systematic Biology</secondary-title><alt-title>Systematic biology</alt-title></titles><periodical><full-title>Systematic Biology</full-title></periodical><alt-periodical><full-title>Systematic Biology</full-title></alt-periodical><pages>384-395</pages><volume>64</volume><number>3</number><dates><year>2015</year><pub-dates><date>May</date></pub-dates></dates><isbn>1076-836X (Electronic)
1063-5157 (Linking)</isbn><accession-num>25472575</accession-num><urls><related-urls><url>;(but see Sigwart and Lindberg 2015 for a discussion on alternative molluscan relationships), Annelida, or the membership of some phyla and supraphyletic clades. Nonetheless, morphology cannot be simply discarded as a resource for inferring relationships, as it is the ultimate test for the proposed phylogenies and the only possible way to place huge amounts of extinct diversity onto the animal tree of life. Fossils are also crucial for dating phylogenies, and tip-dating is developing as a preferred and more accurate method for estimating divergences.Generating transcriptomes or genomes is now possible for non-model organisms and feasible for many researchers. Transcriptomes require live, frozen or RNAlater-preserved tissues, which has forced many of us to re-collect tissues that were not preserved in such ways, making years’ worth of collecting efforts inaccessible to phylogenomics. However, the possibility of cheap sequencing of genomes now allows making use of large numbers of specimens collected in ethanol and preserved in freezers in many museums and university collections. Transcriptome sequencing is now possible for single individuals of the tiniest animals PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5MYXVtZXI8L0F1dGhvcj48WWVhcj4yMDE1PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA (e.g., Laumer et al. 2015a; Laumer et al. 2015b), and genome amplification also allows sequencing of genomes from the smallest animals. Having eliminated these previous technical limitations, emphasis will now shift towards selecting the species of interest and will once again require taxonomic expertise for collecting, vouchering and identifying such organisms. Genomics has largely driven our understanding of animal relationships in recent times, but animal phylogeny remains the arena of zoologists, and not of just molecular biologists who rely on others for specimens and taxonomic identifications, and who may not always understand the importance of keeping vouchers and basic information about the collected specimens.AcknowledgementsAndreas Wanninger solicited this review, based on work developed during the past few years in part supported by the US National Science Foundation (Grants #0334932, #0531757, #0732903, and #1457539). 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