Modern evolutionary theory developed with the main objective to explain and predict changes in populations from one generation to the next, and how species evolve. Largely independently, paleontologists have accrued a body of knowledge informed by patterns of organismal diversity in deep time, but with the limitations of the fossil record. How mechanisms causing differentiation between populations ultimately contribute to macroevolution, however, remains a central question in evolutionary biology. I propose to address this question by studying mechanisms of differentiation at various levels of biological organization in evolutionary radiations of freshwater mollusks of the East African Rift System (EARS), an outstanding emerging model system. The EARS is geographically subdivided in a set of quasi-replicate systems to study evolution at various spatial and taxonomic scales; its freshwater mollusks are diverse, with life-history traits that typically correlate well to ecological niches; diversification is currently ongoing so that processes leading to speciation can be reconstructed accurately; and ancestors can be followed over long periods with substantial phylogenetic and time control. We will develop insight into the link between microevolution and macroevolution in two work packages (WPs): in WP1 we will study how processes of population differentiation lead to speciation and in WP2 we will embed our findings of WP1 into their wider taxonomic and geographic context to provide insight into macroevolution. Our approach will be similar in both WPs in that we will integrate data on genetic diversity, morphological disparity and habitat characteristics to examine how diversity originates (WP1) and is maintained (WP2). Methodologically, we will develop a next-generation sequencing (NGS) pipeline with two steps: 1) transcriptome sequencing to construct a reference transcriptome, and 2) the construction of an NGS library from transcriptomic data for gene-capture approaches (which can be used on existing EtOH-preserved collections). Additionally, morphological disparity will be studied with the same geometric morphometric methods across both WPs, and ecological data will be collected from sampling localities. In WP1 we will focus on mechanisms of differentiation in two mollusk clades, i.e. Bellamya [4 nominal species] and Nyassunio [3 nominal species] from the Malawi Basin. We aim to determine which processes (ecological or non-ecological) have caused differentiation and how these processes have interacted in space and time in the multiple speciation events that took place or are ongoing. We will obtain insight into mechanisms by comparing the genetic, morphological and environmental bases of differentiation from as many populations in the basin as possible. Specifically, single-nucleotide polymorphism data will be obtained from NGS, geometric morphometric data from shells, and these data will be subjected to population genomic studies together with habitat information. In WP2 we will establish relations in monophyletic clades of Viviparidae and Unionidae (African Bellamyinae and Coelaturini, to which Bellamya and Nyassunio, respectively, belong) on a rift-wide scale from NGS sequence data. These reconstructions will be used to test hypotheses on biogeographic patterns, parallel and iterative evolution, and trait evolution in relation to diversification. Secondly, we will integrate fossil taxa to study patterns of extinction selectivity and to test whether macroevolutionary conclusions based on the modern fauna hold when fossils are included. Finally, results from WPs 1 & 2 will be integrated to examine to what extent microevolutionary patterns affect macroevolution, and thus how drastic differences in diversity even in closely related clades can be explained. Beyond a profound influence on evolutionary biology, this understanding is vital to understand the implications of environmental changes on modern diversity.

The Wallacea region, lying between the Borneo to the west and Papau New Guinea to the east, is one of the world's biodiversity hotspots, hosting incredibly high levels of biodiversity, much of which is unique to the region. This exceptional level of biodiversity and endemism reflects evolutionary diversification and radiation over millions of years in one of the world's most geologically complex and active regions. The region's exceptional biodiversity, however, is threatened by climate change, direct exploitation and habitat destruction and fragmentation from land use change. Continued habitat loss and fragmentation is expected to precipitate population declines, increase extinction rates, and could also lead to 'reverse speciation' where disturbance pushes recently diverged species together, leading to increased hybridisation, genetic homogenisation, and species' collapse. Already, approximately 1,300 Indonesian species have been listed as at risk of extinction, but the vast majority of the region's biodiversity has not been assessed and we lack basic information on the distribution and diversification of many groups, let alone understanding of what processes drove their diversification, how they will respond to future environmental change, and how to minimize species' extinctions and losses of genetic diversity while balancing future sustainable development needs. In response to the need for conservation and management strategies to minimize the loss of Wallacea's unique biodiversity under future environmental change and future development scenarios, we will develop ForeWall, a genetically explicit individual-based model of the origin and future of the region's biodiversity. ForeWall will integrate state-of-the-art eco-evolutionary modelling with new and existing ecological and evolutionary data for terrestrial and aquatic taxa including mammals, reptiles, amphibians, freshwater fish, snails, damselflies and soil microbes, to deliver fresh understanding of the processes responsible for the generation, diversification, and persistence of Wallacea's endemic biodiversity. After testing and calibrating ForeWall against empirical data, we will forecast biodiversity dynamics across a suite of taxa under multiple environmental change and economic development scenarios. We will develop a set of alternative plausible biodiversity management/mitigation options to assess the effectiveness of these for preserving ecological and evolutionary patterns and processes across the region, allowing for policy-makers to minimise biodiversity losses during sustainable development. Our project will thus not only provide novel understanding of how geological and evolutionary processes have interacted to generate this biodiversity hotspot, but also provide policy- and decision-makers with tools and evidence to help preserve it.

Novel anatomical adaptations and mechanisms for feeding are often postulated as 'key innovations' that spark the diversification of major clades. However, the mechanics of these adaptations are rarely quantitatively or rigorously tested, seriously undermining the validity of these hypotheses. Moreover, the majority of biomechanical analyses are carried out on single exemplar organisms, whereas a comparative phylogenetic context is critical to understanding the impact of feeding on evolutionary history and testing macroevolutionary hypotheses. Dinosaurs dominated terrestrial ecosystems for >130 million years, exhibiting a tremendous range of body sizes, shapes and ecologies. The earliest dinosaurs and their ancestors were generalists and minor faunal components. Dramatic increases in body size, diversity and abundance occurred during the Late Triassic-Early Jurassic (230-180 million years ago), and various factors have been implicated in dinosaur success. It is thought that the appearance of novel feeding adaptations permitted ecological diversification. However, this engaging 'functional story' has not been tested in a quantitative, hypothesis-driven comparative framework and previous work has focused on derived dinosaur taxa with extreme morphologies (e.g., Tyrannosaurus, Diplodocus), ignoring forms close to the base of Dinosauria. For these reasons, dinosaurs are an ideal model system for integrating data on feeding biomechanics with phylogeny, allowing more rigorous investigation of the relationship between functional diversity and clade dynamics. In this project we aim to comprehensively understand the consequences of functional changes in dinosaur skull biomechanics during the origin and early evolution of dinosaurs, a key moment in life's history. The proposed project is particularly timely given the availability and integration of cutting-edge computational methods for biomechanical analyses and new discoveries of early dinosaurs and their ancestors. We will integrate principles and methods from palaeontology, biology and engineering to reconstruct skull anatomy and function in 15 early dinosaur and dinosauriform taxa. CT scans and visualization software will be used to create 3D computer models. Information from the original fossils and living crocodilians, birds and lizards will be used to reconstruct head musculature. Using these reconstructions and multi-body dynamics analysis, we will model jaw motions during feeding, estimate bite forces along the tooth row and calculate maximum jaw closing speed. We will integrate results from dynamic models with finite element analysis and geometric morphometrics to test how the skulls respond to feeding-induced loads. In addition, we will run simulations on three living species to ensure model predictions are accurate. Results from these analyses will provide evidence for the jaw function and potential diet of early dinosaurs, and whether they became more specialized in terms of feeding performance during their evolution. Finally, we will compare the appearance of feeding characters to dinosaur diversity patterns to determine what role feeding had in their early evolution and success. Palaeontologists, anatomists, biomechanists, evolutionary biologists and engineers will benefit from this work, which will set new benchmarks for performing evolutionary biomechanics in living and fossil animals and will establish new UK, European and overseas collaborations. This project will also generate new methodological advances that can be applied to other clades and other functional questions. Finally, the technological and visual aspects of this work and its focus on early dinosaurs will appeal to the general public, offering numerous engagement opportunities and media interest that will contribute to increased public understanding of scientific principles and methods, and will ensure wide dissemination of this work.