Geerat J. Vermeij | Paleobiology
Historians have debated whether pathways and events from the past to the present are influenced largely by contingency, the dependence of outcomes on particular prior conditions, or whether there is long-term emergent directional change. Previous arguments for predictability in evolutionary history relied on the high frequency of convergence, but the repeated evolution of widely favored adaptations need not imply long-term directionality. Using evidence from the fossil record and arguments concerning the metabolic evolution of organisms, I show here that power (total energy taken up and expended per unit time) has increased stepwise over time at ecosystem-level and global scales thanks to the ratchet-like, cumulative effects of competition and cooperation and to the disproportionate influence of powerful top competitors and opportunistic species on emergent ecosystem properties and processes. The history of life therefore exhibits emergent directionality at large ecosystem-wide scales toward greater power.