Niche conservatism and ecological change during the Late Devonian mass extinction.

Studies of the fossil record can inform our understanding of not only the causes of mass extinctions, but also their effects on biodiversity, ecology and evolution. Here, we examine regional-scale ecological changes resulting from a Late Devonian mass extinction event using brachiopod fossil assemblages from the Appalachian Basin. About half of the species went extinct, but were largely replaced by new immigrant taxa. Both before and after the extinction, the primary gradient in faunal composition was correlated with onshore-offshore position, with a second gradient attributed to frequency of disturbance. Survivors of the extinction displayed a strong degree of niche conservatism along these gradients. Despite these indicators of ecological stability, the pre- and post-extinction faunas were quite distinct at the order level, with atrypids and strophomenids largely replaced by productids, whose spiny shells may have provided greater resistance to disturbance and/or predation. Thus, extinction survivors persisted in similar ecological niches despite environmental perturbations and considerable change in the taxonomic and ecological composition of the regional species pool.

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.

Extinction intensity, selectivity and their combined macroevolutionary influence in the fossil record.

The macroevolutionary effects of extinction derive from both intensity of taxonomic losses and selectivity of losses with respect to ecology, physiology and/or higher taxonomy. Increasingly, palaeontologists are using logistic regression to quantify extinction selectivity because the selectivity metric is independent of extinction intensity and multiple predictor variables can be assessed simultaneously. We illustrate the use of logistic regression with an analysis of physiological buffering capacity and extinction risk in the Phanerozoic marine fossil record. We propose the geometric mean of extinction intensity and selectivity as a metric for the influence of extinction events. The end-Permian mass extinction had the largest influence on the physiological composition of the fauna owing to its combination of high intensity and strong selectivity. In addition to providing a quantitative measure of influence to compare among past events, this approach provides an avenue for quantifying the risk posed by the emerging biodiversity crisis that goes beyond a simple projection of taxonomic losses.

Modelling the ecological-functional diversification of marine Metazoa on geological time scales.

The ecological traits and functional capabilities of marine animals have changed significantly since their origin in the late Precambrian. These changes can be analysed quantitatively using multi-dimensional parameter spaces in which the ecological lifestyles of species are represented by particular combinations of parameter values. Here, we present models that describe the filling of this multi-dimensional 'ecospace' by ecological lifestyles during metazoan diversification. These models reflect varying assumptions about the processes that drove ecological diversification; they contrast diffusive expansion with driven expansion and niche conservatism with niche partitioning. Some models highlight the importance of interactions among organisms (ecosystem engineering and predator-prey escalation) in promoting new lifestyles or eliminating existing ones. These models reflect processes that were not mutually exclusive; rigorous analyses will continue to reveal their applicability to episodes in metazoan history.

Accelerated mass extinction in an isolated biota during Late Devonian climate changes.

The fossil record can illuminate factors that contribute to extinction risk during times of global environmental disturbance; for example, inferred thermal tolerance was an important predictor of extinction during several mass extinctions that corresponded with climate change. Additionally, members of geographically isolated biotas may face higher risk because they have less opportunity to migrate to suitable climate refugia during environmental disturbances. Here, we investigate how different types of risk intersect in the well-preserved brachiopod fauna of the Appalachian Foreland Basin during the two pulses of the Frasnian-Famennian mass extinction (Late Devonian, ~ 372 Ma). The selectivity of extinction is consistent with climate change (cooling) as a primary kill mechanism in this fauna. Overall, the extinction was mild relative to other regions, despite the many endemic species. However, vulnerable taxa went extinct more rapidly, during the first extinction pulse, such that the second pulse was insignificant. These results suggest that vulnerable taxa in geographically isolated biotas face heightened extinction risk at the initiation of environmental stress, but that taxa in other regions may eventually see elevated extinction risk if environmental stress repeats or intensifies.

Ecologically diverse clades dominate the oceans via extinction resistance.

Ecological differentiation is correlated with taxonomic diversity in many clades, and ecological divergence is often assumed to be a cause and/or consequence of high speciation rate. However, an analysis of 30,074 genera of living marine animals and 19,992 genera of fossil marine animals indicates that greater ecological differentiation in the modern oceans is actually associated with lower rates of origination over evolutionary time. Ecologically differentiated clades became taxonomically diverse over time because they were better buffered against extinction, particularly during mass extinctions, which primarily affected genus-rich, ecologically homogeneous clades. The relationship between ecological differentiation and taxonomic richness was weak early in the evolution of animals but has strengthened over geological time as successive extinction events reshaped the marine fauna.

Ecological selectivity of the emerging mass extinction in the oceans.

To better predict the ecological and evolutionary effects of the emerging biodiversity crisis in the modern oceans, we compared the association between extinction threat and ecological traits in modern marine animals to associations observed during past extinction events using a database of 2497 marine vertebrate and mollusc genera. We find that extinction threat in the modern oceans is strongly associated with large body size, whereas past extinction events were either nonselective or preferentially removed smaller-bodied taxa. Pelagic animals were victimized more than benthic animals during previous mass extinctions but are not preferentially threatened in the modern ocean. The differential importance of large-bodied animals to ecosystem function portends greater future ecological disruption than that caused by similar levels of taxonomic loss in past mass extinction events.