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Molecular and Genomic Data Identify the Closest Living Relative of Primates Science Jan E. Janecka, etal. Sc/ence 318, 792(2007); DOI: 10.1126/science.1147555 The following resources related to this article are available online at www.sciencemag.org (this Information is current as of November 2, 2007): Updated information and services, including iiigli-resolution figures, can be found in tine online version of this article at: http://www.sciencennag.org/cgi/content/full/318/5851/792 Supporting Online IVIaterial can be found at: o http://www.sciencemag.org/cgi/content/full/318/5851/792/DC1 o CM cvT This article cites 23 articles, 9 of which can be accessed for free: 0) http://www.sciencennag.Org/cgi/content/full/318/5851/792#otherarticles .Q E 0)> This article appears in the following subject collections: o Evolution z http://www.sciencennag.org/cgi/collection/evolution o O) Infornnation about obtaining reprints of this article or about obtaining permission to reproduce o d) this article in whole or in part can be found at: ça http://www.sciencennag.org/about/permissions.dtl E 0) ü o ü E p •o 0) •o oça o Q Science (print ISSN 0036-8075; online ISSN 1095-9203) is published weekly, except the last week in December, by the American Association for the Advancement of Science, 1200 New York Avenue NW, Washington, DC 20005. Copyright 2007 by the American Association for the Advancement of Science; all rights reserved. The title Science is a registered trademark of AAAS. I REPORTS 14. M. S. Grinolds, V. A. Lobastov, ]. Weissenrieder, 30. ]. B. Goodenough, Phys. Rev. 120, 67 (1960). 38. From the reflectivity of 0.28 and the absorption depth A. H. Zewail, Proc. Notl. Acod. Sei. U.S.A. 103, 18427 31. T. C. Koethe et al.. Phys. Rev. Lett. 97, 116402 of 100 nm at 800 nm (20), the threshold fluence corresponds (2006). (2006). to 450 m]/mm^ at the surface. With the unit cell volume 15. V. A. Lobastov, ]. Weissenrieder, ]. Tang, A. H. Zewail, 32. Given the x-ray wavelength (1.54 A) and diffraction angle of 118 A^ {21), which contains four vanadium atoms, Nono Lett. 7, 2552 (2007). (13.9°) reported (22), the Bragg peak {22) should be this energy density gives -0.05 photon per vanadium. 16. H. S. Park, ]. S. Baskin, O.-H. Kwon, A. H. Zewail, Nano indexed as (Oil) of the monochnic phase. It was assigned 39. G. 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Rossman for the crystal-cutting equipment, 75, 011004 (2006), and references therein. -100 ps, as calculated from the electron mobility of 1 to and L M. Henling for help with the x-ray measurements. 26. H.-T. Kim et oL. Phys. Rev. Lett. 97, 266401 (2006). 10 cm^/(V-s) for metallic vanadium dioxide (20). The This work was supported by the NSF, by the Air Force 27. P. Baum, A. H. Zewail, Proc. Noti Acad. Sei. U.S.A. 103, diffusion of such electrons into the deeper regions may Office of Scientific Research, and by the Gordon and 16105 (2006). contribute to generation of shear. Betty Moore Center for Physical Biology at Caltech. 28. D.-S. Yang, N. Gedik, A. H. Zewail, ]. Phys. Chem. C 111, 37. Shear motion leads to a change in principal axes {34). P.B. was partially supported by the Alexander von 4889 (2007). Because not all Bragg spots are equally well in Humboldt Foundation. 29. V. A. Lobastov et al., in Ultrofost Optics IV, F. Krausz, phase with the Ewald sphere at the same time {28), G. Korn, P. Corkum, \. A. Walmsley, Eds., voL 95 of shear motion may enhance or suppress the Bragg Springer Series in Opticol Sciences (Springer, New York, intensities to values above or below the initial intensity, 12 July 2007; accepted 19 September 2007 2004), pp. 419-435. as observed. 10.1126/science.ll47724 Sundatheria (3, 6, 14), although support for this Molecular and Genomic Data hypothesis was lower than for other mammalian interordinal clades (15). Primatomorpha, proposed Identify the Closest Living on morphological grounds (12), has also been in- dicated by some molecular studies (16, 17). Other studies have failed to reject altemative hypotheses, Relative of Primates and analyses of different character subsets support contradictory topologies (18-20). ]an E. Janecka,^ Webb Miller/ Thomas H. Pringle,' Frank Wiens,'' Annette Zitzmann,^ To improve our understanding of early Kristofer l\A. Helgen,* Mark S. Springer/ William ]. Murphy^* euarchontan evolution and determine the closest living relative of primates, we used two inde- A full understanding of primate morphological and genomic evolution requires the pendent molecular approaches. We first screened identification of their closest living relative. In order to resolve the ancestral relationships among a nonredundant set of 197,522 protein-coding primates and their closest relatives, we searched multispecies genome alignments for exons from the human University of Califor- phylogenetically informative rare genomic changes within the superordinal group Euarchonta, nia Santa Cruz Known Genes track to identify which includes the orders Primates, Dermoptera (colugos), and Scandentia (treeshrews). We also rare genomic changes (exonic indels) that would constructed phylogenetic trees from 14 kilobases of nuclear genes for representatives from most potentially support the three a priori hypotheses. major primate lineages, both extant colugos, and multiple treeshrews, including the pentail We also assembled and analyzed a 14-kb nu- treeshrew, Ptilocercus lowii, the only living member of the family Ptilocercidae. A relaxed clear gene data set of 19 gene fragments in order molecular clock analysis including Ptilocercus suggests that treeshrews arose approximately to estimate a phylogeny and time scale for ex- 63 million years ago. Our data show that colugos are the closest living relatives of primates and tant euarchontans. To mitigate against the pos- indicate that their divergence occurred in the Cretaceous. sible effects of long-branch attraction (LBA), which can incorrecfly place rapidly evolving clades The origins of modem primates and their the monophyly of treeshrews, colugos (flying fossil relatives remain a topic of intense lemurs), and primates in the clade Euarchonta, ^Department of Veterinary Integrative Biosciences, Texas A&M debate (i-i), as there has been an in- with a sister-group relationship to Glires [which University, College Station, TX 77843, USA. ^Center for Com- parative Genomics and Bioinformatics, Pennsylvania State creased focus on identifying adaptive evolution- includes rodents and lagomorphs (3, 6-8)\. In University, University Park, PA 16802, USA. ^Sperling ary changes within primates and the dynamics contrast, the relationships within Euarchonta Foundation, Eugene, OR 97405, USA. "Department of Animal of genome evolution within the primate lineage are not well resolved, most likely because of the Physiology, University of Bayreuth, D-95440 Bayreuth, {4, 5). Resolving higher primate relationships rapid evolution of these groups and inadequate Germany. ^Zoological Institute, Johann Wolfgang Goethe- University, D-60054 Frankfurt/Main, Germany. 'National has been challenging, making it difficult to sampling within Scandentia and Dermoptera. Museum of Natural History, Smithsonian Institution, Wash- identify character transformations in early pri- Three hypotheses have been proposed: (i) a ington, DC 20013, USA. 'Department of Biology, University mate evolution. An essential part of this chal- sister-group relationship between treeshrews and of California, Riverside, CA 92521, USA. lenge is to determine the closest living relative primates (9-11), (ii) a sister-group relationship *To whom correspondence should be addressed at the to primates, which would provide a broader between colugos and primates [Primatomorpha Department of Veterinary Integrative Biosciences, College of context for understanding primate evolution. (12)], and (iii) both colugos and treeshrews as Veterinary Medicine and Biomédical Sciences, Veterinary Medical Administration Building, Room 107, Texas A&M DNA sequence and morphological studies, sister to the primates [Sundatheria (2, 13)]. Mo- University, College Station, TX 77843-4458, USA.