Review of the major post–Middle Ordovician lithotectonic elements of the Appalachian orogen indicates that the middle to late Paleozoic geologic evolution of the Appalachian margin was less uniform than that of the early Paleozoic. Evolutionary divergence between the northern and southern segments of the orogen started in the Late Ordovician to Silurian with staggered accretion of the first peri-Gondwanan elements to reach the Laurentia margin, Carolinia in the south and Ganderia in the north. Divergence was amplified during the Silurian, specifically with respect to the nature of the Laurentian margin and the history of accretion. During this time frame, the northern margin was convergent, whereas the amagmatic southern margin may well have been a transform boundary. In terms of accretion, the Late Silurian–Early Devonian docking of Avalonia was restricted to the northern segment, whereas the southern Appalachians appear to have been largely quiescent during this interval. The evolutionary paths of the two segments of the margin converge on a common history in the Late Devonian during the Famennian event; we suggest that this tectonism was related to the initial marginwide interaction of Laurentia with the peri-Gondwanan blocks of Meguma and Suwanee, providing a uniform tectonic template for margin evolution. The Laurentian-Gondwanan collision is marked by second-order divergences in history. Specifically, during the Carboniferous, the southern segment records a larger component of shortening than the northern Appalachians.