The paradox of the first tier: an agenda for paleobiology

Abstract

Nature's discontinuities occur both in the hierarchical structuring of genealogical individuals and in the distinct processes operating at different scales of time, here called tiers. Conventional evolutionary theory denies this structuring and attempts to render the larger scales as simple extrapolation from (or reduction to) the familiar and immediate—the struggle among organisms at ecological moments (conventional individuals at the first tier). I propose that we consider distinct processes at three separable tiers of time: ecological moments, normal geological time (trends during millions of years), and periodic mass extinctions.I designate as “the paradox of the first tier” our failure to find progress in life's history, when conventional theory (first tier processes acting on organisms) expects it as a consequence of competition under Darwin's metaphor of the wedge. I suggest a resolution of the paradox: whatever accumulates at the first tier is sufficiently reversed, undone, or overriden by processes of the higher tiers. In particular, punctuated equilibrium at the second tier produces trends for suites of reasons unrelated to the adaptive benefits of organisms (conventional progress). Mass extinction at the third tier, a recurring process now recognized as more frequent, more rapid, more intense, and more different than we had imagined, works by different rules and may undo whatever the lower tiers had accumulated.