Late Cretaceous ammonoids show that drivers of diversification are regionally heterogeneous.

Palaeontologists have long sought to explain the diversification of individual clades to whole biotas at global scales. Advances in our understanding of the spatial distribution of the fossil record through geological time, however, has demonstrated that global trends in biodiversity were a mosaic of regionally heterogeneous diversification processes. Drivers of diversification must presumably have also displayed regional variation to produce the spatial disparities observed in past taxonomic richness. Here, we analyse the fossil record of ammonoids, pelagic shelled cephalopods, through the Late Cretaceous, characterised by some palaeontologists as an interval of biotic decline prior to their total extinction at the Cretaceous-Paleogene boundary. We regionally subdivide this record to eliminate the impacts of spatial sampling biases and infer regional origination and extinction rates corrected for temporal sampling biases using Bayesian methods. We then model these rates using biotic and abiotic drivers commonly inferred to influence diversification. Ammonoid diversification dynamics and responses to this common set of diversity drivers were regionally heterogeneous, do not support ecological decline, and demonstrate that their global diversification signal is influenced by spatial disparities in sampling effort. These results call into question the feasibility of seeking drivers of diversity at global scales in the fossil record.

Ecological niche conservatism spurs diversification in response to climate change.

Lengthy debate has surrounded the theoretical and empirical science of whether climatic niche evolution is related to increased or decreased rates of biological diversification. Because species can persist for thousands to millions of years, these questions cross broad scales of time and space. Thus, short-term experiments may not provide comprehensive understanding of the system, leading to the emergence of contrasting opinions: niche evolution may increase diversity by allowing species to explore and colonize new geographic areas across which they could speciate; or, niche conservatism might augment biodiversity by supporting isolation of populations that may then undergo allopatric speciation. Here, we use a simulation approach to test how biological diversification responds to different rates and modes of niche evolution. We find that niche conservatism promotes biological diversification, whereas labile niches-whether adapting to the conditions available or changing randomly-generally led to slower diversification rates. These novel results provide a framework for understanding how Earth-life interactions produced such a diverse biota.

Knowledge gaps and missing links in understanding mass extinctions: Can mathematical modeling help?

Extinction of species, and even clades, is a normal part of the macroevolutionary process. However, several times in Earth history the rate of species and clade extinctions increased dramatically compared to the observed "background" extinction rate. Such episodes are global, short-lived, and associated with substantial environmental changes, especially to the carbon cycle. Consequently, these events are dubbed "mass extinctions" (MEs). Investigations surrounding the circumstances causing and/or contributing to mass extinctions are on-going, but consensus has not yet been reached, particularly as to common ME triggers or periodicities. In part this reflects the incomplete nature of the fossil and geologic record, which - although providing significant information about the taxa and paleoenvironmental context of MEs - is spatiotemporally discontinuous and preserved at relatively low resolution. Mathematical models provide an important opportunity to potentially compensate for missing linkages in data availability and resolution. Mathematical models may provide a means to connect ecosystem scale processes (i.e., the extinction of individual organisms) to global scale processes (i.e., extinction of whole species and clades). Such a view would substantially improve our understanding not only of how MEs precipitate, but also how biological and paleobiological sciences may inform each other. Here we provide suggestions for how to integrate mathematical models into ME research, starting with a change of focus from ME triggers to organismal kill mechanisms since these are much more standard across time and spatial scales. We conclude that the advantage of integrating mathematical models with standard geological, geochemical, and ecological methods is great and researchers should work towards better utilization of these methods in ME investigations.

Spatio-temporal climate change contributes to latitudinal diversity gradients.

The latitudinal diversity gradient (LDG), where the number of species increases from the poles to the Equator, ranks among the broadest and most notable biodiversity patterns on Earth. The pattern of species-rich tropics relative to species-poor temperate areas has been recognized for well over a century, but the generative mechanisms are still debated vigorously. We use simulations to test whether spatio-temporal climatic changes could generate large-scale patterns of biodiversity as a function of only three biological processes-speciation, extinction and dispersal-omitting adaptive niche evolution, diversity-dependence and coexistence limits. In our simulations, speciation resulted from range disjunctions, whereas extinction occurred when no suitable sites were accessible to species. Simulations generated clear LDGs that closely match empirical LDGs for three major vertebrate groups. Higher tropical diversity primarily resulted from higher low-latitude speciation, driven by spatio-temporal variation in precipitation rather than in temperature. This suggests that spatio-temporal changes in low-latitude precipitation prompted geographical range disjunctions over Earth's history, leading to high rates of allopatric speciation that contributed to LDGs. Overall, we show that major global biodiversity patterns can derive from interactions of species' niches (fixed a priori in our simulations) with dynamic climate across complex, existing landscapes, without invoking biotic interactions or niche-related adaptations.

Impacts of Niche Breadth and Dispersal Ability on Macroevolutionary Patterns.

We describe a spatially explicit simulation experiment designed to assess relative impacts of macroecological traits on patterns of biological diversification under changing environmental conditions. Using a simulation framework, we assessed impacts of species' niche breadth (i.e., the range of their abiotic tolerances) and dispersal ability on resulting patterns of speciation and extinction and evaluated how these traits, in conjunction with environmental change, shape biological diversification. Simulation results supported both niche breadth and dispersal ability as important drivers of diversification in the face of environmental change, and suggested that the rate of environmental change influences how species interact with the extrinsic environment to generate diversity. Niche breadth had greater effects on speciation and extinction than dispersal ability when climate changed rapidly, whereas dispersal ability effects were elevated when climate changed slowly. Our simulations provide a bottom-up perspective on the generation and maintenance of diversity under climate change, offering a better understanding of potential interactions between species' intrinsic macroecological characteristics and a dynamic extrinsic environment in the process of biological diversification.

Sharks that pass in the night: using Geographical Information Systems to investigate competition in the Cretaceous Western Interior Seaway.

One way the effects of both ecology and environment on species can be observed in the fossil record is as changes in geographical distribution and range size. The prevalence of competitive interactions and species replacements in the fossil record has long been investigated and many evolutionary perspectives, including those of Darwin, have emphasized the importance of competitive interactions that ultimately lead one species to replace another. However, evidence for such phenomena in the fossil record is not always manifest. Here we use new quantitative analytical techniques based on Geographical Information Systems and PaleoGIS tectonic reconstructions to consider this issue in greater detail. The abundant, well-preserved fossil marine vertebrates of the Late Cretaceous Western Interior Seaway of North America provide the component data for this study. Statistical analysis of distributional and range size changes in taxa confirms earlier ideas that the relative frequency of competitive replacement in the fossil record is limited to non-existent. It appears that typically, environmental gradients played the primary role in determining species distributions, with competitive interactions playing a more minor role.