Two kinds of factors set the tempo and direction of organic and cultural evolution, those external to biotic evolutionary process, such as changes in the earth’s physical and chemical environments, and those internal to it, such as the time required for chance factors to lead lineages across adaptive valleys to a new niche space (Valentine 1985). The relative importance of these two sorts of processes is widely debated. Valentine (1973) argued that marine invertebrate diversity patterns responded to seafloor spreading as this process generated more or less niche space. He suggested that natural selection is a powerful force and that earth’s biota are in near equilibrium with the niches available on the geological time scale. Walker and Valentine (1984) modeled the evolution of species assuming a logistic speciation rate limited by internal factors and a diversity-independent death rate caused by ongoing environmental change. Fitting this model to the observed evolution of shelled marine invertebrates suggests that the lag between extinctions and the evolution of new species leaves perhaps 30% of ecological niches unfilled. In this model, the biota lag environmental change by perhaps a few million years. However, as Valentine (1985) notes, if adaptive landscapes have whole suites of niches protected by deep maladaptive valleys, the waiting time for some pioneering species to cross the divide may be very long, generating the rare events that set new body plans and generate major adaptive radiations. Eldredge and Gould (1972) and Gould (2002) championed the idea that internal processes such as genetic and developmental constraints, coupled with the complexity of the adaptive landscape, resulted in a highly historically contingent evolutionary process. On Gould’s account, most of the history of life had to do not with a relatively close tracking of a changing environment but with the halting evolutionary exploration a deeply fissured niche space, mostly by rapid bursts of evolution as a fissure was crossed, followed by long periods of stasis..