A.P.C. was supported by a Wellcome Trust Clinical Career Development Fellowship no. WT104551MA; A.F. was supported by Arthritis Research UK Clinician Scientist Fellowship no. 18547; F.B. was supported by an Arthritis Research UK Senior Fellowship; H.M.M. was supported by an Arthritis Research UK Career Development Fellowship (19899); C.W. was supported by a Deutsche Forschungsgemeinschaft (DFG) Fellowship (ref. 319464273); A.J.N was supported by a Versus Arthritis Career Development Fellowship no. 21743; K.W. was supported by Rheumatology Research Foundation Scientist Development Award; K.J. was supported by a Wellcome Trust PhD studentship; S.N.S. and M.A. are supported by the Kennedy Trust for Rheumatology Research. K.D. was supported by a Deutsche Forschungsgemeinschaft (DFG) award CRC1181. This work was supported by the Arthritis Research UK Rheumatoid Arthritis Pathogenesis Centre of Excellence no. 20298 (RACE); The National Institutes of Health Accelerating Medicines Partnership in RA/SLE and Arthritis Research UK programme grant no. 19791 (to C.D.B.). This paper presents independent research supported by the NIHR Birmingham Biomedical Research Centre at the University Hospitals Birmingham NHS Foundation Trust and the University of Birmingham. The views expressed are those of the author(s) and not necessarily those of the NHS, the NIHR, our funding bodies or the Department of Health.

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Nature thanks Jason Cyster, Thomas A. Wynn and the other anonymous reviewer(s) for their contribution to the peer review of this work.