Incumbent replacement: Evidence for long-term evolutionary progress

Evolutionary progress is a trend that relaxes trade-off rules. It begins with the evolution of a key adaptation. It continues with the spread of the key adaptation as the clade that contains it replaces some older clade that lacks it. Key adaptations are those that allow for improvement in at least one organismal function at a reduced fitness cost in other functions. Replacement almost certainly involves more than pure chance. It may not often involve competitive extinction. Instead, species from the new clade produce new species to replace already extinct species from the old clade. The key adaptation gives them a higher competitive speciation rate than old-clade sources of replacement. The process, termed incumbent replacement, proceeds at a rate limited by extinction rate. Thus, replacement often seems linked to mass extinction events. The incumbent-replacement hypothesis explains what we know about the replacement of straight-neck turtles (Amphichelydia) by those that can flex their necks and protect their heads in their shells. This replacement occurred four or five times in different biotic provinces. It happened as long ago as the Cretaceous in Eurasia, and as recently as the Pleistocene in mainland Australia. It was accomplished in Gondwanaland by turtles flexing their necks sideways (Pleurodira), and in the north by those flexing their necks into an S-curve (Cryptodira). As is typical of replacements, amphichelydian replacement took millions of years to accomplish wherever it occurred, and much of it in North America took place in a burst associated with and immediately subsequent to a mass extinction.