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.

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Color morphs of the temperate, nonsymbiotic corallimorpharian Corynactis californica show variation in pigment pattern and coloring. We collected seven distinct color morphs of C. californica from subtidal locations in Monterey Bay, California, and found that tissue- and color-morph-specific expression of at least six different genes is responsible for this variation. Each morph contains at least three to four distinct genetic loci that code for these colors, and one morph contains at least five loci. These genes encode a subfamily of new GFP-like proteins, which fluoresce across the visible spectrum from green to red, while sharing between 75% to 89% pairwise amino-acid identity. Biophysical characterization reveals interesting spectral properties, including a bright yellow protein, an orange protein, and a red protein exhibiting a "fluorescent timer" phenotype. Phylogenetic analysis indicates that the FP genes from this species evolved together but that diversification of anthozoan fluorescent proteins has taken place outside of phylogenetic constraints, especially within the Corallimorpharia. The discovery of more examples of fluorescent proteins in a non-bioluminescent, nonsymbiotic anthozoan highlights possibilities of adaptive ecological significance unrelated to light regulation for algal symbionts. The patterns and colors of fluorescent proteins in C. californica and similar species may hold meaning for organisms that possess the visual pigments to distinguish them.

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