The thoughts and writings of Stephen Jay Gould have had an enormous impact on the shaping of macroevolutionary theory. The notion of punctuated equilibria (Eldredge and Gould 1972) remained prominent throughout his work. It also unleashed a storm of debate in paleontology and evolutionary theory. A second theme that recurs throughout Gould's opus is heterochrony (evolution by changes in the rates and timing of ontogenetic events, sensu Gould 1977a). His analyses of these two subjects have inspired many of us to explore further and add to them. My contribution discusses their expansion to encompass large numbers of lineages through long time, and the relationship of punctuated equilibria and heterochrony to physical environmental change, and to each other.

Much of what is known about the long-term history of biodiversity and rates of taxonomic evolution in the fossil record derives from literature-based compilations of fossil stratigraphic ranges. It has been suggested that taxonomic and stratigraphic errors in these compilations are randomly distributed and, therefore, introduce no significant bias to macroevolutionary patterns. Here we compare a new, comprehensive global database of Ordovician and Early Silurian crinoids to Sepkoski's global genus compendium.

Approximately 44% of the crinoid genera resolved to substage in Sepkoski's compendium are taxonomically inaccurate (i.e., invalid, nomina dubia, or column genera) or have incorrect first and/or last occurrences. Errors in Sepkoski's compendium result from incomplete coverage of existing taxonomic work and incorrect stratigraphic correlations that, in some cases, are propagated throughout the taxonomic literature. Stratigraphic range errors are nonrandomly distributed among substages in Sepkoski's compendium. The result is underestimated richness in the Early Silurian and significantly overestimated rates of extinction in the Late Ordovician. There is no similar bias in Sepkoski's substage origination rates for crinoids.

At the stage-level of temporal resolution, Sepkoski's crinoid data are more accurate. In this case, only 32% of the compendium's crinoid genera contain some stratigraphic or taxonomic inaccuracy. However, errors still result in incorrect macroevolutionary patterns, particularly with respect to rate of origination in the Ashgill, which is significantly underestimated in Sepkoski's compendium. Genera described since the completion of Sepkoski's compendium have had relatively little effect on estimated rates of evolution at both stage and substage resolution.

These results suggest that macroevolutionary patterns among some taxa in Sepkoski's compilation may be significantly influenced by nonrandomly distributed taxonomic inaccuracies and stratigraphic range errors. In the case of the apparent end-Ordovician mass extinction among crinoids, the revised history reveals a dramatically reduced role for extinction at the substage-level of temporal resolution. At the stage level, Sepkoski's original compilation strongly exaggerates the excess of extinction over origination in the Ashgill. Although biases inherent in the stratigraphic record remain unaccounted for, removing taxonomic and stratigraphic errors in Sepkoski's compendium substantially changes our understanding of the nature of large-scale biotic change for an important Paleozoic taxon during the end-Ordovician.

We outline a plausible evolutionary sequence that led from prokaryotes to the origin of the first nucleated cell. The nucleus is postulated to evolve after the archaebacterium and eubacterium merged to form the symbiotic ancestor of amitochondriate protists. Descendants of these amitochondriate cells (archaeprotists) today thrive in organic-rich anoxic habitats where they are amenable to study. Eukaryosis, the origin of nucleated cells, occurred by the middle Proterozoic Eon prior to the deposition in sediments of well-preserved microfossils such as Vandalosphaeridium and the spiny spheres in the Doushantou cherts of China.

Mass extinctions are important to macroevolution not only because they involve a sharp increase in extinction intensity over “background” levels, but also because they bring a change in extinction selectivity, and these quantitative and qualitative shifts set the stage for evolutionary recoveries. The set of extinction intensities for all stratigraphic stages appears to fall into a single right-skewed distribution, but this apparent continuity may derive from failure to factor out the well-known secular trend in background extinction: high early Paleozoic rates fill in the gap between later background extinction and the major mass extinctions. In any case, the failure of many organism-, species-, and clade-level traits to predict survivorship during mass extinctions is a more important challenge to the extrapolationist premise that all macroevolutionary processes are simply smooth extensions of microevolution. Although a variety of factors have been found to correlate with taxon survivorship for particular extinction events, the most pervasive effect involves geographic range at the clade level, an emergent property independent of the range sizes of constituent species. Such differential extinction would impose “nonconstructive selectivity,” in which survivorship is unrelated to many organismic traits but is not strictly random. It also implies that correlations among taxon attributes may obscure causation, and even the focal level of selection, in the survival of a trait or clade, for example when widespread taxa within a major group tend to have particular body sizes, trophic habits, or metabolic rates. Survivorship patterns will also be sensitive to the inexact correlations of taxonomic, morphological, and functional diversity, to phylogenetically nonrandom extinction, and to the topology of evolutionary trees. Evolutionary recoveries may be as important as the extinction events themselves in shaping the long-term trajectories of individual clades and permitting once-marginal groups to diversify, but we know little about sorting processes during recovery intervals. However, both empirical extrapolationism (where outcomes can be predicted from observation of pre- or post-extinction patterns) and theoretical extrapolationism (where mechanisms reside exclusively at the level of organisms within populations) evidently fail during mass extinctions and their evolutionary aftermath. This does not mean that conventional natural selection was inoperative during mass extinctions, but that many features that promoted survivorship during background times were superseded as predictive factors by higher-level attributes. Many intriguing issues remain, including the generality of survivorship rules across extinction events; the potential for gradational changes in selectivity patterns with extinction intensity or the volatility of target clades; the heritability of clade-level traits; the macroevolutionary consequences of the inexact correlations between taxonomic, morphological, and functional diversity; the factors governing the dynamics and outcome of recoveries; and the spatial fabric of extinctions and recoveries. The detection of general survivorship rules—including the disappearance of many patterns evident during background times—demonstrates that studies of mass extinctions and recovery can contribute substantially to evolutionary theory.