Illite crystallinity data from the Silurian slate belts of England and Wales indicate anchizone to low epizone metamorphism during the Acadian deformation in late Early Devonian time. This metamorphic grade implies a substantial overburden, now eroded, of Lower Devonian non-marine sediments of the Old Red Sandstone (ORS) magnafacies. A minimum 3.5 km pre-tectonic thickness of ‘lost’ ORS is estimated in the southern Lake District and comparable thicknesses in North Wales and East Anglia. Tectonically driven subsidence of the underlying Avalonian crust is required to accommodate such thicknesses of non-marine sediment. One proposed mechanism is flexure of the Avalonian footwall during convergence that continued from Iapetus closure in the Silurian until Acadian cleavage formation in the Emsian. The evidence for this model in the critical area of northwest England is reviewed and found to be unconvincing. An alternative model is developed following a recent suggestion that the Early Devonian was a period not of continued convergence but of orogen-wide sinistral transtension. Transtensional accommodation of the lost ORS is evidenced by Early Devonian extensional faults, by synchronous lamprophyric magmatism, and by compatibility with previously diagnosed sediment provenance patterns. A summary of Siluro-Devonian tectonostratigraphy for Britain south of the Highland Border emphasizes that, unlike the Scottish Highlands, this area was not affected by the Scandian Orogeny, but was by the Acadian. An important period of sinistral transtension in the Early Devonian (c.420–400 Ma) was common to both regions. This was a time of high heat flow, lamprophyric and more evolved magmatism, and major southward sediment transport, involving mainly recycled metamorphic detritus from the Highlands and from contemporaneous volcanicity. Old Red Sandstone, deposited in coalescing transtensional basins over much of Britain from the Midland Valley to the Welsh Borders, was largely removed and recycled southward during Acadian inversion.