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See detailSupplementary Information for "Global ecomorphological restructuring of dominant marine reptiles prior to the K/Pg mass extinction"
Maclaren, James ULiege; Bennion, Rebecca ULiege; Bardet, Nathalie ULiege et al

in Maclaren, James; Bennion, Rebecca; Bardet, Nathalie (Eds.) et al Global ecomorphological restructuring of dominant marine reptiles prior to the K/Pg mass extinction (2022)

Mosasaurid squamates were the dominant amniote predators in marine ecosystems during most of the Late Cretaceous. Evidence from multiple sites worldwide of a global mosasaurid community restructuring ... [more ▼]

Mosasaurid squamates were the dominant amniote predators in marine ecosystems during most of the Late Cretaceous. Evidence from multiple sites worldwide of a global mosasaurid community restructuring across the Campanian–Maastrichtian transition may have wide-ranging implications for the evolution of diversity of these top oceanic predators. In this study, we use a suite of biomechanical traits and functionally descriptive ratios to investigate how the morphofunctional disparity of mosasaurids evolved through time and space prior to the Cretaceous-Palaeogene (K/Pg) mass extinction. Our results suggest that the worldwide taxonomic turnover in mosasaurid community composition from Campanian to Maastrichtian is reflected by a notable increase in morphofunctional disparity on a global scale, but especially driven the North American record. Ecomorphospace occupation becomes more polarised during the late Maastrichtian, as the morphofunctional disparity of mosasaurids plateaus in the Southern Hemisphere and decreases in the Northern Hemisphere. We show that these changes are not associated with strong modifications in mosasaurid size, but rather with the functional capacities of their skulls, and that mosasaurid morphofunctional disparity was in decline in several provincial communities before the K-Pg mass extinction. Our study highlights region-specific patterns of disparity evolution, and the importance of assessing vertebrate extinctions both globally and regionally. Ecomorphological differentiation in mosasaurid communities, coupled with declines in other formerly abundant marine reptile groups, indicates widespread restructuring of higher trophic levels in marine food webs was well underway when the K-Pg mass extinction took place. [less ▲]

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See detailMarine tetrapod feeding guilds using automated high-density 3D geometric morphometrics
Fischer, Valentin ULiege; Maclaren, James ULiege; Bennion, Rebecca ULiege et al

Conference (2021, December)

Defining feeding guilds based on tooth morphology is an entire subfield in marine tetrapod science. However, these assessments have mostly been qualitative, relying on gross tooth shape, rarely preserved ... [more ▼]

Defining feeding guilds based on tooth morphology is an entire subfield in marine tetrapod science. However, these assessments have mostly been qualitative, relying on gross tooth shape, rarely preserved gut content, and killing behaviour. Moreover, some of the data at the foundation of these guilds have proven to be debatable and there is an ever clearer need for a testable, quantitative framework to assess feeding guilds. We develop a novel protocol that incorporates the pseudo-landmarking technique into high-density geometric morphometrics procedures, sampling 3D surface models of tooth crowns automatically and densely (e.g. 2000 surface landmarks) after placing just 5 fixed landmarks on each tooth. A crushing-to-piercing transition is evident along the first axis of the PCA-based morphospace, while the presence and shape of carinae, as well as crown curvature, is captured by the second axis. This allows an efficient visualisation of tooth shapes with just two axes. Peculiar structures such as strong crown curvature or carinae are mostly recorded on medium-sized teeth, suggesting that a scaling factor is at play. We attempt a new definition of marine reptile feeding guilds based on tooth morphology and size, using extant polarizers. [less ▲]

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See detailLeaving the triangle: marine tetrapod tooth guilds using automated high-density 3d geometric morphometrics
Fischer, Valentin ULiege; Maclaren, James ULiege; Bennion, Rebecca ULiege et al

Conference (2021, April 20)

Defining feeding guilds based on tooth morphology is an entire subfield in marine tetrapod science, which essentially started with a seminal paper by Judy Massare in 1987. However, these assessments have ... [more ▼]

Defining feeding guilds based on tooth morphology is an entire subfield in marine tetrapod science, which essentially started with a seminal paper by Judy Massare in 1987. However, these assessments have mostly been qualitative, subjectively placing teeth on a triangle using a series of criteria that draw both from tooth shape, rarely preserved gut content, and killing/hunting behaviour. Moreover, some of the data at the foundation of these guilds have proven to be debatable and there is an ever clearer need for a testable, quantitative framework to assess feeding guilds. We develop a novel protocol that incorporates the pseudo-landmarking technique into high-density geometric morphometrics procedures, sampling 3D surface models of tooth crowns automatically and densely (e.g. 2000 surface landmarks) after placing just 5 fixed landmarks on each tooth. This very dense sampling of shape permits full characterisation of crown shape, and we provide the first ever quantification of dental morphospace occupation among a sample of mosasaurs, ichthyosaurs, plesiosaurs, archaeocetes, and odontocetes with conical (i.e., non-multicuspid) teeth. A crushing-to-piercing transition is evident along the first axis of the PCA-based morphospace, while the presence and shape of carinae, as well as crown curvature, is captured by the second axis. This allows an efficient visualisation of tooth shape differences and definition of regions/guilds with just two axes that explain ca. 90% of the total variance. Because crown shape has a direct functional signal, our results provide important data to better understand how marine tetrapods evolved and functioned. Despite their gigantic size, the teeth of the largest marine macropredator ever, the physeteroid Livyatan melvillei, have crowns with an unremarkable shape, occupying a position close to the center of the morphospace. In fact, peculiar structures such as crown curvature or carinae are mostly recorded on medium-sized teeth, suggesting that a scaling factor is at play. Similarly, crown shapes that have rarely or never been evolved, either by certain groups or during certain time periods, also carries information. For example, the perfectly straight teeth of the Early Jurassic ichthyosaurian Temnodontosaurus platyodon with protruding flange-like carinae are unique and indicate neoichthyosaurians developed a unique pathway towards hypercarnivory, albeit only once. Further work will incorporate as many different taxa and tooth-shapes into the morphospace as possible to usher in a new, quantitative paradigm for understanding marine tetrapod feeding ecology. [less ▲]

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See detailA European mosasaurine (Squamata: Mosasauridae) with affinities to a clade from new zealand
Street, Hallie; Caldwell, Michael; Maclaren, James ULiege et al

Conference (2021)

The Royal Belgian Institute of Natural Sciences houses an extensive collection of Maastrichtian-aged mosasaur fossils. Among the numerous specimens of these marine reptiles is a skull and associated ... [more ▼]

The Royal Belgian Institute of Natural Sciences houses an extensive collection of Maastrichtian-aged mosasaur fossils. Among the numerous specimens of these marine reptiles is a skull and associated vertebral column, IRSNB 3211. This specimen was originally assigned to Plioplatecarpus houzeaui, but more recent studies have suggested that it represents an immature individual of Mosasaurus lemonnieri. New three-dimensional surface scans of IRSNB 3211 contribute to a detailed description of the specimen. Several characters strongly disagree with the diagnosis for Plioplatecarpus houzeaui, including the height of the coronoid process and the presence of zygosphenes and zygrantra. The proportions of the quadrate, the complex frontal/parietal suture, and the morphology of the posterior mandibular elements identify this specimen as belonging to Mosasaurinae. However, the morphology of the skull and vertebrae differs from that of Mosasaurus lemonnieri. Skeletal immaturity could account for some of these differences, such as the relatively longer rostrum or shorter quadrate of IRSNB 3211. However, ontogenetic change does not explain the differences in the morphology of the maxilla or braincase elements. Phylogenetic analyses do not find a sister-group relationship between Mosasaurus lemonnieri and IRSNB 3211. The results of these analyses instead indicate a relationship between this specimen and Moanasaurus mangahouangae within a clade of mosasaurines from the Pacific Ocean. Characters that unite IRSNB 3211 with Moanasaurus mangahouangae include dorsal excavation of the maxilla for the external naris and cervical vertebrae with laterally extended postzygapophyses. However, skull proportions and differences in dental morphology do not support assigning this specimen to Moanasaurus mangahouangae. The qualitative results of morphological comparisons and the quantitative results of phylogenetic analysis support assigning IRSNB 3211 to a new taxon. Moanasaurus mangahouangae and its closest relatives have previously been considered to have been endemic to New Zealand. The New Zealand fossils are from Campanian strata, indicating the clade originated in the Southern Pacific and migrated to the Tethys during the Maastrichtian. Other fossils in European collections are likely representatives of this new taxon, but general similarities between this moanasaur-like taxon and Mosasaurus lemonnieri masked the diversity of latest Cretaceous mosasaurines from Europe. [less ▲]

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See detailLeviathans Unleashed: Skull Ecomorphological Evolution During The Initial Aquatic Radiations Of Mosasaurs And Cetaceans
Bennion, Rebecca ULiege; Maclaren, James ULiege; Coombs, Ellen et al

Conference (2021)

The repeated return of tetrapods to aquatic environments provides many iconic examples of convergent evolution, with various groups of mammals and reptiles independently evolving streamlined body shapes ... [more ▼]

The repeated return of tetrapods to aquatic environments provides many iconic examples of convergent evolution, with various groups of mammals and reptiles independently evolving streamlined body shapes and similar feeding strategies. One comparison that has received little attention is that between mosasaurs (a group of Late Cretaceous marine squamates) and early cetaceans (middle to late Eocene ancestors of modern whales and dolphins). These two groups share broad similarities in skull morphology, filling a wide range of niches and achieving global distributions. The earliest fully aquatic members of both groups had serpentine bodies and swam by axial undulation, before evolving more efficient caudal oscillatory locomotion and colonising open ocean niches. Cetaceans continued to diversify after reaching this form whereas the evolutionary history of the mosasaurs was cut short by the end-Cretaceous mass extinction. Here, we investigate possible parallel evolutionary trajectories of skull morphology that occurred during these initial aquatic radiations. A series of functionally informative ratios were calculated from 32 species of mosasaurs and early cetaceans. These were subjected to ordination techniques to reconstruct patterns of functional ecomorphospace occupation, and putative examples of convergence were tested statistically. Preliminary results show that the earliest mosasaurs had gracile skulls, specialised for smaller prey, from which they radiated in several waves across the ecomorphospace. There is considerable variation within certain genera, such as Mosasaurus. By contrast, basilosaurid cetaceans occupy a relatively constrained megapredatory niche and cetaceans only evolved new ecomorphologies after the late Eocene split into odontocetes and mysticetes. Oligocene odontocetes explore a new area of morphospace away from the basilosaurids, evolving a long, narrow snout with an increased number of small teeth. The earliest toothed mysticetes have a similar ecomorphology to the basilosaurids, with aetiocetids appearing to radiate in a similar direction to the odontocetes. The late Oligocene Janjucetus, which has a highly unusual ecomorphology, plots away from other cetaceans. Despite showing striking similarities to the mosasaur Prognathodon (e.g short robust snout and large eyes), the two taxa were not found to be statistically convergent. However, cranial convergence was found between the mosasaur Mosasaurus hoffmanni and the basilosaurid Dorudon atrox. Future work will investigate these results using 3D landmark analyses, and the evolutionary trajectories in early mysticetes will be extrapolated by including toothless species. [less ▲]

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See detailInfluences on cranial morphology in whales: Investigating the evolutionary history and diversity of the cetacean skull
Coombs, Ellen; Clavel, Julien; Felice, Ryan et al

Conference (2021)

Odontoceti (toothed whales) and Mysticeti (baleen whales) diverged ~39 Ma. Odontocetes evolved high-frequency echolocation and shifted cranial bones posteriorly, while mysticetes evolved larger masses and ... [more ▼]

Odontoceti (toothed whales) and Mysticeti (baleen whales) diverged ~39 Ma. Odontocetes evolved high-frequency echolocation and shifted cranial bones posteriorly, while mysticetes evolved larger masses and filter-feeding. Despite an excellent fossil record, little quantitative study of shape evolution spanning cetacean diversity has been done. Here, we quantify morphological disparity and evolutionary rate in cranial shape to identify the major factors driving its evolution, including ecological traits and palaeoclimate. We scanned 88 living and 111 Eocene to Pliocene fossil cetaceans, representing the largest 3D dataset for cetaceans ever constructed. We used 123 3D coordinate-based landmarks and 1905 curve semilandmarks to capture cranial shape and analyzed data within a phylogenetic framework. The largest component of cranial variation (PC1 = 41.6%) reflects a posterior shift in the nares, followed by rostrum length (PC2 = 23.0%) with dolichocephalic (e.g., Pontoporia blainvillei) and brachycephalic (e.g., Kogia sima) crania representing the extremes. After accounting for phylogeny, habitat had no significant effect on skull shape, but diet, feeding, and tooth type did. The highest morphological disparity is in the premaxilla and nasal, with the highest evolutionary rate ( 2 mult) in the frontal. Disparity rises rapidly in the Middle-Late Oligocene and peaks in the Middle-Late Miocene. A model with rate of cranial evolution tracking paleotemperature outperformed Brownian motion and Early Burst models and suggests highest rates in the Latest Eocene and Early-Mid Oligocene (during periods of global cooling) with a further peak in evolutionary rates during the Mid-Miocene warming period. [less ▲]

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See detailHow Whitby got its whale jaw arch (evolutionarily speaking)
Bennion, Rebecca ULiege

Conference given outside the academic context (2020)

The whalebone arch at the top of Whitby’s West Cliff is an iconic tourist attraction and landmark of the town. It pays homage to the town’s history as a whaling port in the 18th and 19th Centuries when ... [more ▼]

The whalebone arch at the top of Whitby’s West Cliff is an iconic tourist attraction and landmark of the town. It pays homage to the town’s history as a whaling port in the 18th and 19th Centuries when over two thousand Arctic whales were hunted for their blubber and oil. The bones that form the arch are the jaws of a bowhead whale (Balaena mysticetus), one of the main species targeted by whalers. It belongs to a group of whales that use a unique feeding mechanism: instead of catching prey with teeth, they filter them from the water using a specialised hair-like structure called baleen. Their skulls and jaws are specially adapted for this style of feeding and are very different to those of the earliest whales which swam the Late Eocene seas. In this talk, I shall endeavour to give a palaeontological perspective on the bowhead and other baleen whales. I will discuss how recent fossil finds from around the world are providing insight into how baleen filter-feeding evolved from toothed ancestors, as well as some of my own research investigating trends in marine vertebrate evolution using 3D models of fossil skulls. [less ▲]

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See detailCharting new waters: changes in skull ecomorphology during the initial aquatic radiations of mosasaurs and cetaceans
Bennion, Rebecca ULiege; Maclaren, James ULiege; Coombs, Ellen et al

Conference (2020, December)

The repeated return of tetrapods to water provides many iconic examples of convergent evolution, with various groups of mammals and reptiles independently evolving streamlined body shapes and similar ... [more ▼]

The repeated return of tetrapods to water provides many iconic examples of convergent evolution, with various groups of mammals and reptiles independently evolving streamlined body shapes and similar feeding strategies. One comparison which has received little attention is that of cetaceans (whales and dolphins) and mosasaurs (a group of Late Cretaceous marine squamates). The earliest fully aquatic members of both groups had serpentine bodies and swam by axial undulation, before evolving more efficient caudal oscillatory locomotion and colonizing open ocean niches. Here we investigate possible parallel evolutionary trajectories of skull morphology that occurred during these initial aquatic radiations. A series of functionally informative ratios were calculated from 32 species of mosasaur and early cetacean. These were subjected to ordination techniques to reconstruct patterns of functional morphospace occupation. Preliminary results show that the earliest mosasaurs had gracile skulls specialized for smaller prey, from which they radiated in several waves across the morphospace. By contrast, basilosaurid cetaceans occupied a relatively constrained megapredatory niche, and only evolved new ecomorphologies after the Late Eocene split into odontocetes and mysticetes. Theresults also suggest cranial convergence between the toothed mysticete Janjucetus and the mosasaur Prognathodon. Future work will investigate these results further using 3D landmarks. [less ▲]

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See detailCharting new waters: changes in skull ecomorphology during the initial aquatic radiations of mosasaurs and cetaceans
Bennion, Rebecca ULiege; Maclaren, James ULiege; Coombs, Ellen et al

Poster (2020, June)

The earliest fully aquatic cetaceans (Middle to Late Eocene), had a serpentine body shape which was adapted for locomotion by axial undulation. This has drawn comparisons with the mosasaurs, a group of ... [more ▼]

The earliest fully aquatic cetaceans (Middle to Late Eocene), had a serpentine body shape which was adapted for locomotion by axial undulation. This has drawn comparisons with the mosasaurs, a group of Late Cretaceous marine squamates, with whom these early cetaceans also share broad similarities in skull morphology, including a triangular snout with widely spaced teeth and an unfused mandibular symphysis. Both groups filled a wide range of niches and achieved global distributions. As they became increasingly adapted to aquatic life, each clade modified their body plan to allow more efficient tail-based ‘thunniform’ locomotion, which enabled colonisation of the open ocean. Cetaceans continued to diversify after reaching this form whereas the evolutionary history of the mosasaurs was cut short by the end-Cretaceous mass extinction. We investigated the evolution of skull ecomorphology through the initial aquatic radiations of mosasaurs and cetaceans. Thirteen ecologically informative craniodental measurements and ratios were taken from 3D models of well-preserved skulls. The initial dataset consisted of ten cetaceans and eleven mosasaurs, sampling across the first twenty million years of evolution in each group. These data were subjected to ordination techniques in R to reconstruct patterns of morphospace occupation. Preliminary results show that Eocene cetaceans had a conserved skull morphology, only branching out into new morphologies in the Oligocene (early odontocetes significantly more so than early toothed mysticetes). By comparison, mosasaurs show much more ecomorphological variation. Future work will use 3D landmarks to accurately compare differences in skull shape through the evolution of each group. [less ▲]

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See detailThe macroevolutionary landscape of short-necked plesiosaurians
Fischer, Valentin ULiege; Maclaren, James ULiege; Soul, Laura C. et al

in Scientific Reports (2020), 10(16434), 1--12

Throughout their evolution, tetrapods have repeatedly colonised a series of ecological niches in marine ecosystems, producing textbook examples of convergent evolution. However, this evolutionary ... [more ▼]

Throughout their evolution, tetrapods have repeatedly colonised a series of ecological niches in marine ecosystems, producing textbook examples of convergent evolution. However, this evolutionary phenomenon has typically been assessed qualitatively and in broad-brush frameworks that imply simplistic macroevolutionary landscapes. We establish a protocol to visualize the density of trait space occupancy and thoroughly test for the existence of macroevolutionary landscapes. We apply this protocol to a new phenotypic dataset describing the morphology of short-necked plesiosaurians, a major component of the Mesozoic marine food webs (ca. 201 to 66 Mya). Plesiosaurians evolved this body plan multiple times during their 135-million-year history, making them an ideal test case for the existence of macroevolutionary landscapes. We find ample evidence for a bimodal craniodental macroevolutionary landscape separating latirostrines from longirostrine taxa, providing the first phylogenetically-explicit quantitative assessment of trophic diversity in extinct marine reptiles. This bimodal pattern was established as early as the Middle Jurassic and was maintained in evolutionary patterns of short-necked plesiosaurians until a Late Cretaceous (Turonian) collapse to a unimodal landscape comprising longirostrine forms with novel morphologies. This study highlights the potential of severe environmental perturbations to profoundly alter the macroevolutionary dynamics of animals occupying the top of food chains. [less ▲]

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See detailTaxonomic and ecomorphological reassessment of the Lower Jurassic ichthyosaur Temnodontosaurus
Bennion, Rebecca ULiege; Fischer, Valentin ULiege

Conference given outside the academic context (2019)

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See detailMandibular shape disparity and convergence in ichthyosaurs and toothed cetaceans
Bennion, Rebecca ULiege; Lambert, Olivier ULiege; Fischer, Valentin ULiege

Poster (2019, July)

Numerous tetrapod lineages have independently evolved to live in the water during the Mesozoic and Cenozoic, offering some of the best-known examples of convergent evolution. In particular, modern toothed ... [more ▼]

Numerous tetrapod lineages have independently evolved to live in the water during the Mesozoic and Cenozoic, offering some of the best-known examples of convergent evolution. In particular, modern toothed cetaceans are often compared to ichthyosaurs, a diverse clade of extinct marine reptiles that also evolved a 'fish-shaped' body plan with tail-propelled locomotion. Both are predominantly raptorial marine tetrapods with long evolutionary histories and good fossil records, yet surprisingly the ecological convergences and the macroevolutionary pathways behind them are poorly understood. This project aims to investigate convergences of ichthyosaur and cetacean skulls on similar morphologies and ecological functions. Here we present results of a preliminary analysis focusing on mandible shape from a sample of archaeocete and odontocete cetaceans and parvipelvian ichthyosaurs. Landmarks and semi-landmarks were placed onto photographs of specimens or 3D models made with a handheld scanner. The resulting coordinates were subjected to a principal components analysis in R to show mandibular shape disparity, with preserved stomach contents and tooth shape data used to correlate how this morphological variation might relate to ecological function. Up-to-date phylogenies can be superimposed to show convergences and trajectories of evolutionary change in the two groups through time. Using these ordination techniques, an adaptive landscape can be created to show which areas of the morphospace ('peaks') are colonised more frequently. These results will form part of the first detailed quantitative analysis of ecomorphological convergence between ichthyosaurs and cetaceans. [less ▲]

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See detailSkull shape evolution in ichthyosaurs and cetaceans: detailed investigations into a classic example of convergence
Bennion, Rebecca ULiege; Lambert, Olivier ULiege; Fischer, Valentin ULiege

Conference (2018, October 11)

Ever since they first made the transition to life on land around 350 million years ago, more than 30 lineages of tetrapods have reinvaded the water independently, filling important roles in aquatic ... [more ▼]

Ever since they first made the transition to life on land around 350 million years ago, more than 30 lineages of tetrapods have reinvaded the water independently, filling important roles in aquatic ecosystems. The constraints that arose from living in water rather than air have forced the evolution of similar morphologies within these groups, making them some of the best-known examples of evolutionary convergence. In particular, modern day toothed cetaceans are often compared to the ichthyosaurs, a diverse clade of extinct marine reptiles which also evolved a ‘fish-shaped’ body plan with tail propelled locomotion. Both are groups of raptorial marine tetrapods with long evolutionary histories and good fossil records, yet surprisingly the ecological convergences and the macroevolutionary pathways behind them are poorly understood and lack a thorough, quantitative framework. The goal of this project is to investigate convergences of ichthyosaur and cetacean skulls on similar morphologies and ecological functions. We gathered a series of measurements and ratios that have direct ecological consequences (e.g. tooth and snout shape) from 2D and 3D preserved specimens. The initial dataset comprises ratios from 31 species of extinct and extant toothed cetaceans and 26 species of ichthyosaurs. These data were subjected to clustering and ordination techniques in R using a 50% completeness threshold. The results were corroborated with preserved stomach contents and modern ecological observations to reveal general convergence patterns within the two groups. These preliminary results will be investigated further in more detail by using them alongside analyses of 3D cranial and jaw landmarks. By combining these with the most up to date phylogenies for each group we can reveal parallel trajectories of convergence in marine tetrapods over time. [less ▲]

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See detailLes bacheliers du bloc 1 Géologie de l'ULiège. Journées de terrain, travaux pratiques: Évolution des environnements et de la tectonique de la région de Boulogne-sur-Mer, Wimereux et Wissant, sous la conduite du Prof. Valentin Fischer et de ses collaborateurs. Séquence video et photos
Marion, Jean-Marc ULiege; Fischer, Valentin ULiege; Denayer, Julien ULiege et al

Learning material (2018)

Field days are essential in learning and understanding geology. From April 16 to 18, 2018, graduates from block 1 Geology of the University of Liège (Belgium) studied the evolution of environments and ... [more ▼]

Field days are essential in learning and understanding geology. From April 16 to 18, 2018, graduates from block 1 Geology of the University of Liège (Belgium) studied the evolution of environments and tectonics in the region of Boulogne-sur-Mer, Wimereux and Wissant (France). [less ▲]

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See detailDiverse and durophagous: Early Carboniferous chondrichthyans from the Scottish Borders
Richards, Kelly; Sherwin, Janet; Smithson, Timothy et al

in Earth and Environmental Science Transactions of the Royal Society of Edinburgh (2018), 108(1), 67-87

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