About the Authors

Jian Zhao

Affiliation Division of Rheumatology, Department of Medicine, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America

Hui Wu

Affiliation Division of Rheumatology, Department of Medicine, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America

Melanie Khosravi

Affiliation Division of Rheumatology, Department of Medicine, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America

Huijuan Cui

Affiliation Joint Molecular Rheumatology Laboratory of Institute of Health Sciences and Shanghai Renji Hospital, Shanghai Jiao Tong University School of Medicine, Shanghai Institutes for Biological Sciences, and Chinese Academy of Sciences, Shanghai, China

Xiaoxia Qian

Affiliation Joint Molecular Rheumatology Laboratory of Institute of Health Sciences and Shanghai Renji Hospital, Shanghai Jiao Tong University School of Medicine, Shanghai Institutes for Biological Sciences, and Chinese Academy of Sciences, Shanghai, China

Jennifer A. Kelly

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

Kenneth M. Kaufman

Affiliations Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America, United States Department of Veterans Affairs Medical Center, Oklahoma City, Oklahoma, 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

Adrienne H. Williams

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

Mary E. Comeau

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

Julie T. Ziegler

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

Miranda C. Marion

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

Adam Adler

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

Marta E. Alarcón-Riquelme

Affiliations Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America, Center for Genomics and Oncological Research, Pfizer-University of Granada-Junta de Andalucia, Granada, Spain

BIOLUPUS Network

Memberships of the consortia are available in the Acknowledgments.

GENLES Network

Memberships of the consortia are available in the Acknowledgments.

Bernardo A. Pons-Estel

Affiliation Sanatorio Parque, Rosario, Argentina

John B. Harley

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

Sang-Cheol Bae

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

So-Young Bang

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

Soo-Kyung Cho

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

Chaim O. Jacob

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

Timothy J. Vyse

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

Timothy B. Niewold

Affiliation Section of Rheumatology and Gwen Knapp Center for Lupus and Immunology Research, University of Chicago, Chicago, Illinois, 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

Kathy L. Moser

Affiliation Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, 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

Jeffrey C. Edberg

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

Elizabeth E. Brown

Affiliations Department of Medicine, University of Alabama at Birmingham, Birmingham, Alabama, United States of America, Department of Epidemiology, University of Alabama at Birmingham, Birmingham, Alabama, United States of America

Graciela S. Alarcon

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

Michelle A. Petri

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

Rosalind Ramsey-Goldman

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

Luis M. Vilá

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

John D. Reveille

Affiliation Rheumatology and Clinical Immunogenetics, University of Texas Health Science Center at Houston, Houston, Texas, 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

Gary S. Gilkeson

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

Diane L. Kamen

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

Barry I. Freedman

Affiliation Department of Internal Medicine, Wake Forest University Health Sciences, Winston-Salem, North Carolina, United States of America

Juan-Manuel Anaya

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

Joan T. Merrill

Affiliation Clinical Pharmacology, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, 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

R. Hal Scofield

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

Anne M. Stevens

Affiliations Division of Rheumatology, Department of Pediatrics, University of Washington, Seattle, Washington, United States of America, Center for Immunity and Immunotherapies, Seattle Children's Research Institute, Seattle, Washington, 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

Deh-Ming Chang

Affiliation National Defense Medical Center, Taipei, Taiwan

Yeong Wook Song

Affiliation Division of Rheumatology, Seoul National University, Seoul, Korea

Ji Ah Park

Affiliation Division of Rheumatology, Seoul National University, Seoul, Korea

Eun Young Lee

Affiliation Division of Rheumatology, Seoul National University, Seoul, Korea

Susan A. Boackle

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

Jennifer M. Grossman

Affiliation Division of Rheumatology, Department of Medicine, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America

Bevra H. Hahn

Affiliation Division of Rheumatology, Department of Medicine, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America

Timothy H. J. Goodship

Affiliation Institute of Human Genetics, Newcastle University, Newcastle upon Tyne, United Kingdom

Rita M. Cantor

Affiliation Department of Human Genetics, University of California Los Angeles, Los Angeles, California, United States of America

Chack-Yung Yu

Affiliation Department of Pediatrics, The Ohio State University, Columbus, Ohio, United States of America

Nan Shen

Affiliation Joint Molecular Rheumatology Laboratory of Institute of Health Sciences and Shanghai Renji Hospital, Shanghai Jiao Tong University School of Medicine, Shanghai Institutes for Biological Sciences, and Chinese Academy of Sciences, Shanghai, China

Betty P. Tsao

btsao@mednet.ucla.edu

Affiliation Division of Rheumatology, Department of Medicine, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: BP Tsao, J Zhao, H Wu, C-Y Yu, N Shen. Performed the experiments: J Zhao, H Wu, M Khosravi, H Cui, X Qian, A Adler. Analyzed the data: J Zhao, H Wu, M Khosravi, H Cui, X Qian, JA Kelly, KM Kaufman, CD Langefeld, AH Williams, ME Comeau, JT Ziegler, MC Marion, SB Glenn. Contributed reagents/materials/analysis tools: BP Tsao, N Shen, C-Y Yu, ME Alarcón-Riquelme, BA Pons-Estel, JB Harley, S-C Bae, S-Y Bang, S-K Cho, CO Jacob, TJ Vyse, TB Niewold, PM Gaffney, KL Moser, RP Kimberly, JC Edberg, EE Brown, GS Alarcon, MA Petri, R Ramsey-Goldman, LM Vilá, JD Reveille, JA James, GS Gilkeson, DL Kamen, BI Freedman, J-M Anaya, JT Merrill, LA Criswell, RH Scofield, AM Stevens, JM Guthridge, D-M Chang, YW Song, JA Park EY Lee, SA Boackle, JM Grossman BH Hahn, THJ Goodship, RM Cantor. Wrote the paper: J Zhao. Revised the manuscript: BP Tsao, SA Boackle, BH Hahn, JM Grossman, C-Y Yu, EE Brown, JD Reveille, JA James, RM Cantor.