About the Authors

Bahram Namjou

Contributed equally to this work with: Bahram Namjou, Xana Kim-Howard, Celi Sun

Affiliations Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America, Cincinnati Children's Hospital Medical Center, Cincinnati, Ohio, United States of America

Xana Kim-Howard

Contributed equally to this work with: Bahram Namjou, Xana Kim-Howard, Celi Sun

Affiliation Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America

Celi Sun

Contributed equally to this work with: Bahram Namjou, Xana Kim-Howard, Celi Sun

Affiliation Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America

Adam Adler

Affiliation Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America

Sharon A. Chung

Affiliation Rosalind Russell Medical Research Center for Arthritis, Department of Medicine, University of California San Francisco, San Francisco, California, United States of America

Kenneth M. Kaufman

Affiliations Cincinnati Children's Hospital Medical Center, Cincinnati, Ohio, United States of America, US Department of Veterans Affairs Medical Center, Cincinnati, Ohio, United States of America

Jennifer A. Kelly

Affiliation Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America

Stuart B. Glenn

Affiliation Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America

Joel M. Guthridge

Affiliation Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America

Robert H. Scofield

Affiliations Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America, Department of Medicine, University of Oklahoma Health Sciences Center, Oklahoma City, Oklahoma, United States of America

Robert P. Kimberly

Affiliation Department of Medicine, University of Alabama at Birmingham, Birmingham, Alabama, United States of America

Elizabeth E. Brown

Affiliation Department of Medicine, University of Alabama at Birmingham, Birmingham, Alabama, United States of America

Graciela S. Alarcón

Affiliation Department of Medicine, University of Alabama at Birmingham, Birmingham, Alabama, United States of America

Jeffrey C. Edberg

Affiliation Department of Medicine, University of Alabama at Birmingham, Birmingham, Alabama, United States of America

Jae-Hoon Kim

Affiliation Department of Rheumatology, Hanyang University Hospital for Rheumatic Diseases, Seoul, Korea

Jiyoung Choi

Affiliation Department of Rheumatology, Hanyang University Hospital for Rheumatic Diseases, Seoul, Korea

Rosalind Ramsey-Goldman

Affiliation Division of Rheumatology, Northwestern University Feinberg School of Medicine, Chicago, Illinois, United States of America

Michelle A. Petri

Affiliation Department of Medicine, Johns Hopkins University School of Medicine, Baltimore, Maryland, United States of America

John D. Reveille

Affiliation Department of Internal Medicine, University of Texas Health Science Center at Houston, Houston, Texas, United States of America

Luis M. Vilá

Affiliation Department of Medicine, University of Puerto Rico Medical Sciences Campus, San Juan, Puerto Rico

Susan A. Boackle

Affiliation Division of Rheumatology, University of Colorado School of Medicine, Aurora, Colorado, United States of America

Barry I. Freedman

Affiliation Center for Public Health Genomics and Department of Biostatistical Sciences, Wake Forest University Health Sciences, Wake Forest, North Carolina, United States of America

Betty P. Tsao

Affiliation Division of Rheumatology, University of California Los Angeles, Los Angeles, California, United States of America

Carl D. Langefeld

Affiliation Department of Biostatistical Sciences, Wake Forest University Health Sciences, Wake Forest, North Carolina, United States of America

Timothy J. Vyse

Affiliation Divisions of Genetics and Molecular Medicine and Immunology, King's College London, London, United Kingdom

Chaim O. Jacob

Affiliation Department of Medicine, Keck School of Medicine, University of Southern California, Los Angeles, California, United States of America

Bernardo Pons-Estel

Affiliation Department of Medicine, Sanatorio Parque, Rosario, Argentina

on behalf of the Argentine Collaborative Group

Membership of the Argentine Collaborative Group and the BIOLUPUS and GENLES networks is provided in the Acknowledgments.

Timothy B. Niewold

Affiliation Division of Rheumatology and Department of Immunology, Mayo Clinic, Rochester, Minnesota, United States of America

Kathy L. Moser Sivils

Affiliation Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America

Joan T. Merrill

Affiliation Clinical Pharmacology, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America

Juan-Manuel Anaya

Affiliation Center for Autoimmune Diseases Research, Universidad del Rosario, Bogota, Colombia

Gary S. Gilkeson

Affiliation Department of Medicine, Division of Rheumatology, Medical University of South Carolina, Charleston, South Carolina, United States of America

Patrick M. Gaffney

Affiliation Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America

Sang-Cheol Bae

Affiliation Department of Rheumatology, Hanyang University Hospital for Rheumatic Diseases, Seoul, Korea

Marta E. Alarcón-Riquelme

Affiliations Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America, Centro de Genómica e Investigación Oncológica (GENYO) Pfizer-Universidad de Granada-Junta de Andalucía, Granada, Spain

on behalf of the BIOLUPUS and GENLES networks

Membership of the Argentine Collaborative Group and the BIOLUPUS and GENLES networks is provided in the Acknowledgments.

John B. Harley

Affiliations Cincinnati Children's Hospital Medical Center, Cincinnati, Ohio, United States of America, US Department of Veterans Affairs Medical Center, Cincinnati, Ohio, United States of America

Lindsey A. Criswell

Affiliation Rosalind Russell Medical Research Center for Arthritis, Department of Medicine, University of California San Francisco, San Francisco, California, United States of America

Judith A. James

Affiliations Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America, Department of Medicine, University of Oklahoma Health Sciences Center, Oklahoma City, Oklahoma, United States of America

Swapan K. Nath

Swapan-nath@omrf.org

Affiliation Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: SKN. Performed the experiments: KMK AA. Analyzed the data: BN XKH CS. Contributed reagents/materials/analysis tools: SAC JAK SBG JMG RHS RPK EEB GSA JCE JHK JC RR MAP JDR LMV SAB BIF BPT CDL TJV COJ BP TBN KLM JTM JMA GSG PMG SCB MA JBH LAC JAJ. Wrote the paper: BN XKH CS SAC JAJ SKN.