Paleobiogeography, high-resolution stratigraphy, and the future of Paleozoic biostratigraphy: Fine-scale diachroneity of the Wenlock (Silurian) conodont Kockelella walliseri

The Wenlock Epoch of the Silurian Period has become one of the chronostratigraphically best-constrained intervals of the Paleozoic. The integration of multiple chronostratigraphic tools, such as conodont and graptolite biostratigraphy, sequence stratigraphy, and δ13Ccarb chemostratigraphy, has greatly improved global chronostratigraphic correlation and portions of the Wenlock can now be correlated with precision better than ± 100 kyr. Additionally, such detailed and integrated chronostratigraphy provides an opportunity to evaluate the fidelity of individual chronostratigraphic tools. Here, we use conodont biostratigraphy, sequence stratigraphy and carbon isotope (δ13Ccarb) chemostratigraphy to demonstrate that the conodont Kockelella walliseri, an important guide fossil for middle and upper Sheinwoodian strata (lower stage of the Wenlock Series), first appears at least one full stratigraphic sequence lower in Laurentia than in Baltica. Rather than serving as a demonstration of the unreliability of conodont biostratigraphy, this example serves to demonstrate the promise of high-resolution Paleozoic stratigraphy. The temporal difference between the two first occurrences was likely less than 1 million years, and although it is conceptually understood that speciation and colonization must have been non-instantaneous events, Paleozoic paleobiogeographic variability on such short timescales (tens to hundreds of kyr) traditionally has been ignored or considered to be of little practical importance. The expansion of high-resolution Paleozoic stratigraphy in the future will require robust biostratigraphic zonations that embrace the integration of multiple chronostratigraphic tools as well as the paleobiogeographic variability in ranges that they will inevitably demonstrate. In addition, a better understanding of the paleobiogeographic migration histories of marine organisms will provide a unique tool for future Paleozoic paleoceanography and paleobiology research.