About the Authors

Noriko Tonomura

Affiliations Broad Institute of MIT and Harvard, Cambridge, Massachusetts, United States of America, Department of Clinical Sciences, Cummings School of Veterinary Medicine at Tufts University, North Grafton, Massachusetts, United States of America

Ingegerd Elvers

Affiliations Broad Institute of MIT and Harvard, Cambridge, Massachusetts, United States of America, Science for Life Laboratory, Dept. of Medical Biochemistry and Microbiology, Uppsala University, Uppsala, Sweden

Rachael Thomas

Affiliation Department of Molecular Biomedical Sciences, College of Veterinary Medicine, & Center for Comparative Medicine and Translational Research, North Carolina State University, Raleigh, North Carolina, United States of America

Kate Megquier

Affiliations Broad Institute of MIT and Harvard, Cambridge, Massachusetts, United States of America, Science for Life Laboratory, Dept. of Medical Biochemistry and Microbiology, Uppsala University, Uppsala, Sweden

Jason Turner-Maier

Affiliation Broad Institute of MIT and Harvard, Cambridge, Massachusetts, United States of America

Cedric Howald

Affiliation Broad Institute of MIT and Harvard, Cambridge, Massachusetts, United States of America

Aaron L. Sarver

Affiliation Department of Veterinary Clinical Sciences, College of Veterinary Medicine, University of Minnesota, Saint Paul, Minnesota, United States of America

Ross Swofford

Affiliation Broad Institute of MIT and Harvard, Cambridge, Massachusetts, United States of America

Aric M. Frantz

Affiliations Department of Veterinary Clinical Sciences, College of Veterinary Medicine, University of Minnesota, Saint Paul, Minnesota, United States of America, Masonic Cancer Center, University of Minnesota, Minneapolis, Minnesota, United States of America

Daisuke Ito

Affiliations Department of Veterinary Clinical Sciences, College of Veterinary Medicine, University of Minnesota, Saint Paul, Minnesota, United States of America, Masonic Cancer Center, University of Minnesota, Minneapolis, Minnesota, United States of America

Evan Mauceli

Affiliations Broad Institute of MIT and Harvard, Cambridge, Massachusetts, United States of America, Department of Laboratory Medicine, Boston Children’s Hospital, Boston, Massachusetts, United States of America

Maja Arendt

Affiliation Science for Life Laboratory, Dept. of Medical Biochemistry and Microbiology, Uppsala University, Uppsala, Sweden

Hyun Ji Noh

Affiliation Broad Institute of MIT and Harvard, Cambridge, Massachusetts, United States of America

Michele Koltookian

Affiliation Broad Institute of MIT and Harvard, Cambridge, Massachusetts, United States of America

Tara Biagi

Affiliation Broad Institute of MIT and Harvard, Cambridge, Massachusetts, United States of America

Sarah Fryc

Affiliation Broad Institute of MIT and Harvard, Cambridge, Massachusetts, United States of America

Christina Williams

Affiliation Department of Molecular Biomedical Sciences, College of Veterinary Medicine, & Center for Comparative Medicine and Translational Research, North Carolina State University, Raleigh, North Carolina, United States of America

Anne C. Avery

Affiliations Department of Microbiology, Immunology, and Pathology, Colorado State University College of Veterinary Medicine and Biomedical Sciences, Fort Collins, Colorado, United States of America, Animal Cancer Center, Colorado State University College of Veterinary Medicine and Biomedical Sciences, Fort Collins, Colorado, United States of America

Jong-Hyuk Kim

Affiliations Department of Veterinary Clinical Sciences, College of Veterinary Medicine, University of Minnesota, Saint Paul, Minnesota, United States of America, Masonic Cancer Center, University of Minnesota, Minneapolis, Minnesota, United States of America

Lisa Barber

Affiliation Department of Clinical Sciences, Cummings School of Veterinary Medicine at Tufts University, North Grafton, Massachusetts, United States of America

Kristine Burgess

Affiliation Department of Clinical Sciences, Cummings School of Veterinary Medicine at Tufts University, North Grafton, Massachusetts, United States of America

Eric S. Lander

Affiliation Broad Institute of MIT and Harvard, Cambridge, Massachusetts, United States of America

Elinor K. Karlsson

Affiliations Broad Institute of MIT and Harvard, Cambridge, Massachusetts, United States of America, FAS Center for Systems Biology, Harvard University, Cambridge, Massachusetts, United States of America

Chieko Azuma

Current address: University of Massachusetts Medical School, Worcester, Massachusetts, United States of America

Affiliation Department of Clinical Sciences, Cummings School of Veterinary Medicine at Tufts University, North Grafton, Massachusetts, United States of America

Jaime F. Modiano

Contributed equally to this work with: Jaime F. Modiano, Matthew Breen, Kerstin Lindblad-Toh

Affiliations Department of Veterinary Clinical Sciences, College of Veterinary Medicine, University of Minnesota, Saint Paul, Minnesota, United States of America, Masonic Cancer Center, University of Minnesota, Minneapolis, Minnesota, United States of America

Matthew Breen

Contributed equally to this work with: Jaime F. Modiano, Matthew Breen, Kerstin Lindblad-Toh

Affiliations Department of Molecular Biomedical Sciences, College of Veterinary Medicine, & Center for Comparative Medicine and Translational Research, North Carolina State University, Raleigh, North Carolina, United States of America, Cancer Genetics Program, University of North Carolina Lineberger Comprehensive Cancer Center, Raleigh, North Carolina, United States of America

Kerstin Lindblad-Toh

Contributed equally to this work with: Jaime F. Modiano, Matthew Breen, Kerstin Lindblad-Toh

kersli@broadinstitute.org

Affiliations Broad Institute of MIT and Harvard, Cambridge, Massachusetts, United States of America, Science for Life Laboratory, Dept. of Medical Biochemistry and Microbiology, Uppsala University, Uppsala, Sweden

Competing Interests

I have read the journal’s policy and the authors of this manuscript have the following competing interests: A patent application has been filed related to this work.

Author Contributions

Conceived and designed the experiments: KLT MB CA NT JFM. Analyzed the data: NT EKK IE RS JTM CH ALS JFM HJN KM EM MA KLT. Wrote the paper: NT KLT IE KM. Coordinated, collected, made/confirmed diagnoses and characterized samples for the study: CA MB LB KB RT BD JFM KM TB SF NT MK. Prepared samples for genotyping and managed data generation: MK RS TB SF KM NT. Collected, coordinated, and prepared samples for whole genome sequencing: RS MK RT NT. Prepared the samples for expression studies: AMF ALS DI JHK JFM. Performed immunophenotyping: ACA. Took part in hypotheses and conclusions discussions: ESL CW.