About the Authors

Julio E. Molineros

Contributed equally to this work with: Julio E. Molineros, Amit K. Maiti, Celi Sun

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

Amit K. Maiti

Contributed equally to this work with: Julio E. Molineros, Amit K. Maiti, 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: Julio E. Molineros, Amit K. Maiti, Celi Sun

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

Loren L. Looger

Affiliation Howard Hughes Medical Institute, Janelia Farm Research Campus, Ashburn, Virginia, United States of America

Shizhong Han

Affiliations Arthritis and Clinical Immunology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, United States of America, Department of Psychiatry, Yale School of Medicine, New Haven, Connecticut, United States of America

Xana Kim-Howard

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

Stuart Glenn

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

Jennifer A. Kelly

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

Timothy B. Niewold

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

Gary S. Gilkeson

Affiliation Division of Rheumatology, Medical University of South Carolina, Charleston, South Carolina, 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

Michelle Petri

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

Rosalind Ramsey-Goldman

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

John D. Reveille

Affiliation Department of Rheumatology and Clinical Immunogenetics, University of Texas Health Science Center at Houston, Houston, Texas, United States of America

Luis M. Vilá

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

Barry I. Freedman

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

Betty P. Tsao

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

Lindsey A. Criswell

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

Chaim O. Jacob

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

Jason H. Moore

Affiliation Department of Genetics, Dartmouth Medical School, Lebanon, New Hampshire, United States of America

Timothy J. Vyse

Affiliations Division of Genetics and Molecular Medicine, King's College London, London, United Kingdom, Division of Immunology, Infection and Inflammatory Diseases, Kings College London, London, United Kingdom

Carl L. Langefeld

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

Patrick M. Gaffney

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

Kathy L. Moser

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

R. Hal Scofield

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, Centro de Genómica e Investigación Oncológica (GENyO)–Pfizer/Universidad de Granada/Junta de Andalucía, Granada, Spain

on behalf of the BIOLUPUS Network

Membership of the BIOLUPUS Network is available in Text S1.

Scott M. Williams

Affiliation Department of Genetics, Geisel School of Medicine, Dartmouth College, Hanover, New Hampshire, United States of America

Joan T. Merrill

Affiliation Clinical Pharmacology Research Program, Oklahoma Medical Research Foundation, Oklahoma City, Oklahoma, 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, College of Medicine, University of Oklahoma Health Sciences Center, Oklahoma City, Oklahoma, United States of America

Kenneth M. Kaufman

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

John B. Harley

Affiliation Cincinnati Children's Hospital Medical Center and the U.S. Department of Veterans Affairs Medical Center, Cincinnati, Ohio, 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: AKM JEM. Analyzed the data: JEM CS SH SG XK-H LLL. Contributed reagents/materials/analysis tools: JBH KMK JAK AA SG CS CLL PMG JAJ MEA-R TBN EEB RR-G GSA MP JDR LMV BPT BIF LAC COJ KLM JCE JMG TJV GSG JTM JHM SMW RHS RPK. Wrote the paper: JEM AKM LLL SKN.