About the Authors

Dorothée Diogo

dorothee.diogo@merck.com

Current address: Merck Research Laboratories, Boston, Massachusetts, United States of America

Affiliations Division of Rheumatology, Immunology, and Allergy, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America, Division of Genetics, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America, Program in Medical and Population Genetics, Broad Institute, Cambridge, Massachusetts, United States of America, Partners HealthCare Center for Personalized Genetic Medicine, Boston, Massachusetts, United States of America

Lisa Bastarache

Affiliation Department of Biomedical Informatics, Vanderbilt University, Nashville, Tennessee, United States of America

Katherine P. Liao

Affiliation Division of Rheumatology, Immunology, and Allergy, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America

Robert R. Graham

Affiliation ITGR Human Genetics Group, Genentech Inc, San Francisco, California, United States of America

Robert S. Fulton

Affiliation The Genome Institute, Washington University School of Medicine, St. Louis, Missouri, United States of America

Jeffrey D. Greenberg

Affiliation New York University Hospital for Joint Diseases, New York, New York, United States of America

Steve Eyre

Affiliation Arthritis Research UK Epidemiology Unit, University of Manchester, Manchester Academic Health Sciences Centre, Manchester, United Kingdom

John Bowes

Affiliation Arthritis Research UK Epidemiology Unit, University of Manchester, Manchester Academic Health Sciences Centre, Manchester, United Kingdom

Jing Cui

Affiliation Division of Rheumatology, Immunology, and Allergy, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America

Annette Lee

Affiliation The Feinstein Institute for Medical Research, North Shore-Long Island Jewish Health System, Manhasset, New York, United States of America

Dimitrios A. Pappas

Affiliation Columbia University, College of Physicians and Surgeons, New York, New York, United States of America

Joel M. Kremer

Affiliation The Albany Medical College and The Center for Rheumatology, Albany, New York, United States of America

Anne Barton

Affiliation Arthritis Research UK Epidemiology Unit, University of Manchester, Manchester Academic Health Sciences Centre, Manchester, United Kingdom

Marieke J. H. Coenen

Affiliation Radboud university medical center, Radboud Institute for Health Sciences, Department of Human Genetics, Nijmegen, The Netherlands

Barbara Franke

Affiliation Radboud University Medical Center, Donders Centre for Neurosciences, Department of Psychiatry and Human Genetics, Nijmegen, The Netherlands

Lambertus A. Kiemeney

Affiliation Radboud University Medical Center, Radboud Institute for Health Sciences, Nijmegen, The Netherlands

Xavier Mariette

Affiliations Université Paris-Sud, Orsay, France, APHP–Hôpital Bicêtre, INSERM U1012, Le Kremlin Bicêtre, Paris, France

Corrine Richard-Miceli

Affiliations Université Paris-Sud, Orsay, France, APHP–Hôpital Bicêtre, INSERM U1012, Le Kremlin Bicêtre, Paris, France

Helena Canhão

Affiliations Rheumatology Research Unit, Instituto de Medicina Molecular, Faculdade de Medicina da Universidade de Lisboa, Lisbon, Portugal, Rheumatology Department, Santa Maria Hospital–CHLN, Lisbon, Portugal

João E. Fonseca

Affiliations Rheumatology Research Unit, Instituto de Medicina Molecular, Faculdade de Medicina da Universidade de Lisboa, Lisbon, Portugal, Rheumatology Department, Santa Maria Hospital–CHLN, Lisbon, Portugal

Niek de Vries

Affiliation Amsterdam Rheumatology and Immunology Center, Department of Clinical Immunology & Rheumatology, Academic Medical Center /University of Amsterdam, Amsterdam, The Netherlands

Paul P. Tak

Current address: University of Cambridge, Cambridge, United Kingdom, and GlaxoSmithkline, Stevenage, United Kingdom

Affiliation Amsterdam Rheumatology and Immunology Center, Department of Clinical Immunology & Rheumatology, Academic Medical Center /University of Amsterdam, Amsterdam, The Netherlands

J. Bart A. Crusius

Affiliation Laboratory of Immunogenetics, Department of Medical Microbiology and Infection Control, VU University Medical Center, Amsterdam, The Netherlands

Michael T. Nurmohamed

Affiliation Amsterdam Rheumatology and Immunology Center, Department of Rheumatology, Reade, Amsterdam, The Netherlands

Fina Kurreeman

Affiliation Department of Rheumatology, Leiden University Medical Centre, Leiden, The Netherlands

Ted R. Mikuls

Affiliation Division of Rheumatology and Immunology, Omaha VA and University of Nebraska Medical Center, Omaha, Nebraska, United States of America

Yukinori Okada

Current address: Department of Human Genetics and Disease Diversity, Graduate School of Medical and Dental Sciences, Tokyo Medical and Dental University, and Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan

Affiliations Division of Rheumatology, Immunology, and Allergy, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America, Division of Genetics, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America, Program in Medical and Population Genetics, Broad Institute, Cambridge, Massachusetts, United States of America

Eli A. Stahl

Current address: Division of Psychiatric Genomics, Mt. Sinai School of Medicine, New York, New York, United States of America

Affiliations Division of Rheumatology, Immunology, and Allergy, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America, Division of Genetics, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America, Program in Medical and Population Genetics, Broad Institute, Cambridge, Massachusetts, United States of America

David E. Larson

Affiliation The Genome Institute, Washington University School of Medicine, St. Louis, Missouri, United States of America

Tracie L. Deluca

Affiliation The Genome Institute, Washington University School of Medicine, St. Louis, Missouri, United States of America

Michelle O'Laughlin

Affiliation The Genome Institute, Washington University School of Medicine, St. Louis, Missouri, United States of America

Catrina C. Fronick

Affiliation The Genome Institute, Washington University School of Medicine, St. Louis, Missouri, United States of America

Lucinda L. Fulton

Affiliation The Genome Institute, Washington University School of Medicine, St. Louis, Missouri, United States of America

Roman Kosoy

Affiliation Department of Biochemistry and Molecular Medicine, University of California Davis, Davis, California, United States of America

Michael Ransom

Affiliation Department of Biochemistry and Molecular Medicine, University of California Davis, Davis, California, United States of America

Tushar R. Bhangale

Affiliation ITGR Human Genetics Group, Genentech Inc, San Francisco, California, United States of America

Ward Ortmann

Affiliation ITGR Human Genetics Group, Genentech Inc, San Francisco, California, United States of America

Andrew Cagan

Affiliation Information Systems, Partners Healthcare, Charlestown, Massachusetts, United States of America

Vivian Gainer

Affiliation Information Systems, Partners Healthcare, Charlestown, Massachusetts, United States of America

Elizabeth W. Karlson

Affiliation Division of Rheumatology, Immunology, and Allergy, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America

Isaac Kohane

Affiliation Information Systems, Partners Healthcare, Charlestown, Massachusetts, United States of America

Shawn N. Murphy

Affiliation Information Systems, Partners Healthcare, Charlestown, Massachusetts, United States of America

Javier Martin

Affiliation Instituto de Parasitologia y Biomedicina Lopez-Neyra, CSIC, Granada, 18100, Spain

Alexandra Zhernakova

Affiliations Department of Rheumatology, Leiden University Medical Centre, Leiden, The Netherlands, Genetics Department, University Medical Center and Groningen University, Groningen, The Netherlands

Lars Klareskog

Affiliation Rheumatology Unit, Department of Medicine, Karolinska Institutet and Karolinska University Hospital Solna, Stockholm, Sweden

Leonid Padyukov

Affiliation Rheumatology Unit, Department of Medicine, Karolinska Institutet and Karolinska University Hospital Solna, Stockholm, Sweden

Jane Worthington

Affiliation Arthritis Research UK Epidemiology Unit, University of Manchester, Manchester Academic Health Sciences Centre, Manchester, United Kingdom

Elaine R. Mardis

Affiliation The Genome Institute, Washington University School of Medicine, St. Louis, Missouri, United States of America

Michael F. Seldin

Affiliation Division of Rheumatology and Immunology, Omaha VA and University of Nebraska Medical Center, Omaha, Nebraska, United States of America

Peter K. Gregersen

Affiliation The Feinstein Institute for Medical Research, North Shore-Long Island Jewish Health System, Manhasset, New York, United States of America

Timothy Behrens

Affiliation ITGR Human Genetics Group, Genentech Inc, San Francisco, California, United States of America

Soumya Raychaudhuri

Affiliations Division of Rheumatology, Immunology, and Allergy, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America, Division of Genetics, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America, Program in Medical and Population Genetics, Broad Institute, Cambridge, Massachusetts, United States of America, Partners HealthCare Center for Personalized Genetic Medicine, Boston, Massachusetts, United States of America, Arthritis Research UK Epidemiology Unit, University of Manchester, Manchester Academic Health Sciences Centre, Manchester, United Kingdom

Joshua C. Denny

Affiliations Department of Biomedical Informatics, Vanderbilt University, Nashville, Tennessee, United States of America, Department of Medicine, Vanderbilt University, Nashville, Tennessee, United States of America

Robert M. Plenge

Current address: Merck Research Laboratories, Boston, Massachusetts, United States of America

Affiliations Division of Rheumatology, Immunology, and Allergy, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America, Division of Genetics, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America, Program in Medical and Population Genetics, Broad Institute, Cambridge, Massachusetts, United States of America

Competing Interests

The authors would like to emphasize that Drs T. Behrens, R.R. Graham, T.R. Bhangale and W. Ortmann are employed by Genentech Inc. Dr. J.D. Greenberg is an employee and shareholder in Corrona and has received consulting fees from AstraZeneca, Celgene, Novartis and Pfizer. Dr D.A. Pappas is an employee of Corrona, LLC, and Novartis instructor. J.M. Kremer is a shareholder in Corrona and receives employment compensation. However, this does not alter the authors' adherence to PLOS ONE policies on sharing data and materials.

Author Contributions

Conceived and designed the experiments: DD RMP. Performed the experiments: DD RMP JCD KPL SR. Analyzed the data: DD LB. Contributed reagents/materials/analysis tools: RPG RSF JDG SE JB JC AL DAP JMK AB MJHC BF LAK XM CRM HC JEF NdV PPT JBAC MTN FK TRM YO EAS DEL TLD MO CCF LLF RK MR TRB WO AC VG EWK IK SNM JM AZ LK LP JW ERM MFS PKG TB. Wrote the paper: DD RPM.