The end of the line: competitive exclusion and the extinction of historical entities.

Identifying competitive exclusion at the macroevolutionary scale has typically relied on demonstrating a reciprocal, contradictory response by two co-occurring, functionally similar clades. Finding definitive examples of such a response in fossil time series has proven challenging, however, as has controlling for the effects of a changing physical environment. We take a novel approach to this issue by quantifying variation in trait values that capture almost the entirety of function for steam locomotives (SL), a known example of competitive exclusion from material culture, with the goal of identifying patterns suitable for assessing clade replacement in the fossil record. Our analyses find evidence of an immediate, directional response to the first appearance of a direct competitor, with subsequent competitors further reducing the realized niche of SLs, until extinction was the inevitable outcome. These results demonstrate when interspecific competition should lead to extinction and suggest that clade replacement may only occur when niche overlap between an incumbent and its competitors is near absolute and where the incumbent is incapable of transitioning to a new adaptive zone. Our findings provide the basis for a new approach to analyse putative examples of competitive exclusion that is largely free of assumptions.