Sex and the shifting biodiversity dynamics of marine animals in deep time.

The fossil record of marine animals suggests that diversity-dependent processes exerted strong control on biodiversification: after the Ordovician Radiation, genus richness did not trend for hundreds of millions of years. However, diversity subsequently rose dramatically in the Cretaceous and Cenozoic (145 million years ago-present), indicating that limits on diversification can be overcome by ecological or evolutionary change. Here, we show that the Cretaceous-Cenozoic radiation was driven by increased diversification in animals that transfer sperm between adults during fertilization, whereas animals that broadcast sperm into the water column have not changed significantly in richness since the Late Ordovician (∼450 million years ago). We argue that the former group radiated in part because directed sperm transfer permits smaller population sizes and additional modes of prezygotic isolation, as has been argued previously for the coincident radiation of angiosperms. Directed sperm transfer tends to co-occur with many ecological traits, such as a predatory lifestyle. Ecological specialization likely operated synergistically with mode of fertilization in driving the diversification that began during the Mesozoic marine revolution. Plausibly, the ultimate driver of diversification was an increase in food availability, but its effects on the fauna were regulated by fundamental reproductive and ecological traits.

Secondary evolutionary escalation between brachiopods and enemies of other prey.

The fossil record of predation indicates that attacks on Paleozoic brachiopods were very rare, especially compared to those on post-Paleozoic mollusks, yet stratigraphically and geographically widespread. Drilling frequencies were very low in the early Paleozoic (<<1%) and went up slightly in the mid-to-late Paleozoic. Present-day brachiopods revealed frequencies only slightly higher. The persistent rarity of drilling suggests that brachiopods were the secondary casualties of mistaken or opportunistic attacks by the enemies of other taxa. Such sporadic attacks became slightly more frequent as trophic systems escalated and predators diversified. Some evolutionarily persistent biotic interactions may be incidental rather than coevolutionary or escalatory in nature.

Anatomical and ecological constraints on Phanerozoic animal diversity in the marine realm.

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.