Immune cells are armed with Toll-like receptors (TLRs) for sensing and responding to pathogens and other danger cues1. The role of extracellular signal-regulated kinases 1/2 (Erk1/2) in TLR signalling remains enigmatic, with both pro- and anti-inflammatory functions described. We reveal here that the immune-specific transmembrane adaptor SCIMP is a direct scaffold for Erk1/2 in TLR pathways, with high-resolution live cell imaging revealing that SCIMP guides the spatial and temporal recruitment of Erk2 to membrane ruffles and macropinosomes for pro-inflammatory TLR4 signalling. SCIMP- deficient mice display defects in Erk1/2 recruitment to TLR4, c-Fos activation and pro- inflammatory cytokine production, with these effects being phenocopied by Erk1/2 signalling inhibition. Our findings thus delineate a selective role for SCIMP as a key scaffold2 3 for the membrane recruitment of Erk1/2 kinase to initiate TLR-mediated pro- inflammatory responses in macrophages.