Person:

Bambach, Richard

Issues of directionality in the history of life can be framed in terms of six major evolutionary steps, or megatrajectories (cf. Maynard Smith and Szathmary 1995): (1) evolution from the origin of life to the last common ancestor of extant organisms, (2) the metabolic diversification of bacteria and archaea, (3) evolution of eukaryotic cells, (4) multicellularity, (5) the invasion of the land and (6) technological intelligence. Within each megatrajectory, overall diversification conforms to a pattern of increasing variance bounded by a right wall as well as one on the left. However, the expanding envelope of forms and physiologies also reflects-at least in part-directional evolution within clades. Each megatrajectory has introduced fundamentally new evolutionary entities that garner resources in new ways, resulting in an unambiguously directional pattern of increasing ecological complexity marked by expanding ecospace utilization. The sequential addition of megatrajectories adheres to logical rules of ecosystem function, providing a blueprint for evolution that may have been followed to varying degrees wherever life has arisen.

The repeated association during the late Neoproterozoic Era of large carbon-isotopic excursions, continental glaciation, and stratigraphically anomalous carbonate precipitation provides a framework for interpreting the reprise of these conditions on the Late Permian Earth. A paleoceanographic model that was developed to explain these stratigraphically linked phenomena suggests that the overturn of anoxic deep oceans during the Late Permian introduced high concentrations of carbon dioxide into surficial environments. The predicted physiological and climatic consequences for marine and terrestrial organisms are in good accord with the observed timing and selectivity of Late Permian mass extinction.

We grouped the fossil records of marine animal genera into suites defined by function and physiology, The stratigraphic coherence of the resulting diversity history indicates the importance of ecological structure in constraining taxonomic richness through time. The proportional representation of major functional groups was stably maintained for intervals as long as 200 million years, despite evolutionary turnover and changes in total diversity. Early Paleozoic radiations established stable ecosystem relationships, and thereafter only the great era-bounding mass extinctions were able to break patterns of incumbency, permitting the emergence of new community structures with distinct proportional diversity relationships.