Select Year
Abstracts: Birmingham 2017 - University of Birmingham
 
 
If you copy the HTML page and paste it into a blank Word document it will retain the formattng.
SVPCA Conference
Platform presentation (15 mins)
Tooth-like patterning in the scales of early 'sharks'
 
*
Plamen Andreev
1
 
1
University of Birmingham
In chondrichthyan fish odontodes are ubiquitous skeletal elements, components of dermal and oropharyngeal scales as well as the dentition. Despite years of intense study of odontode development from molecular, anatomical and evolutionary perspectives, there has not been a major breakthrough in understanding the origins of teeth and tooth pattering.
This study presents a new phylogenetic tree of jawed fish that puts at the base of the Chondrichthyes a plethora of mongolepid and other polyodontode scale species (e.g. Tezakia, Canyonlepis) that predate the first appearance of teeth in the Upper Silurian.
Investigations of their scales via tomography, SEM and histological sections identify mechanisms of odontode addition/crown growth related to current theories of pattern formation in the skeleton of early fish.
Common for mongolepid scale crowns are linear rows of odontodes (odontocomplexes) remarkably similar in structure to the tooth whorls of 'acanthodians' and more conventional stem chondrichthyans such as Doliodus. These odontocomplexes are suggested to be capable of continuous orderly growth at one end by generation of ever larger elements in the absence of dental lamina. The dermal skeleton of chondrichthyans can thus be viewed to have primitively possessed the competence to organise odontodes in a manner characteristic of tooth families.
Teeth, development, and hyraxes
 
*
Robert J Asher
1
 
1
 
The life history of animals includes aspects of growth and development such as attainment of maturity and gestation time. Such features are difficult to infer among fossils, but not impossible. I investigated dental homologies, development, and growth in living and fossil hyracoids and tested if hyracoids and other mammals show correlations between eruption patterns and aspects of life history. Most lower cheek teeth finish eruption during growth in hyracoids, not after growth as in most other afrotherians. All hyracoids show the m1 at (lower) or near (upper) the beginning of eruption of permanent teeth; M3 is the last permanent tooth to erupt. The living P. capensis erupts most lower antemolar loci before m2. In contrast, fossil hyraxes erupt lower antemolars after m2. The early eruption of antemolars correlates with increased gestation time and age at maturity in primates and Tupaia (i.e., ‘Schultz’s rule’), and modern hyraxes resemble some anthropoid primates in exhibiting long gestation and eruption of antemolars at or before molars. Among other afrotherians sampled so far, eruption patterns do not significantly covary with either life history parameter.
Upper Karoo vertebrate assemblages (Late Triassic/Early Jurassic) of the Late Kariba region, Zimbabwe
 
*
Paul M Barrett
1,2
 
Tim Broderick
3
 
Kimberley Chapelle
2
 
Jonah Choiniere
2
 
Steve Edwards
6
 
Darlington Munyikwa
7
 
Pia Viglietti
2
 
Michel Zondo
9
 
1
Natural History Museum, London
 
2
University of the Witwatersrand, Johannesburg
 
3
Broderick Geological Consulting, Harare
 
6
Musango Safari Camp, Kariba
 
7
National Museums and Monuments of Zimbabwe, Harare
 
9
Natural History Museum of Zimbabwe, Bulawayo
Southern Africa has provided critical information on the composition of terrestrial tetrapod faunas during the Late Triassic–Early Jurassic, but most of the fossil localities in this region are limited to the
Elliot and Clarens formations of South Africa and Lesotho. By contrast, other countries in the region have been explored less thoroughly, although preliminary work has revealed significant potential. Previous work led to reports of several Early Jurassic localities in Zimbabwe, which have yielded specimens of the basal sauropodomorph Massospondylus, the early sauropod Vulcanodon and some theropod material. In addition, several (largely unreported) Late Triassic localities are also known, which had yielded a small number of fragmentary specimens.

In early 2017, a joint South African-Zimbabwean-UK team conducted fieldwork in the upper Karoo-aged deposits cropping out along the shores of Lake Kariba, northern Zimbabwe (Mid-Zambesi Basin). We were able to relocate the Vulcanodon type locality on Island 126/127 and to make new stratigraphic observations. Contrary to the majority of previous reports, which suggested a Toarcian age for this taxon, the quarry is situated in a horizon that significantly pre-dates the onset of Drakensburg volcanism (= Batoka Basalts) and is located instead within the earlier Lower Jurassic Forest Sandstone. This indicates that Vulcanodon is 10–15 million years older than previously thought, recalibrating several nodes within Sauropoda and indicating extensive ecological overlap between true sauropods and 'prosauropods'.

Exploration of other islands in Lake Kariba and along its southern shore revealed new sites in the Pebbly Arkose (part of the Tashinga Formation: Late Triassic), as well as in the Forest Sandstone. Each of these units provides evidence of terrestrial and freshwater aquatic vertebrate faunas. Sauropodomorphs are present in the Forest Sandstone and upper Tashinga formations, but a grey mudstone facies within the Pebbly Arkose yields a more aquatic fauna, that includes lungfish, carnivorous archosaurs, and phytosaurs, although sauropodomorphs appear to be absent. Thus far, phytosaurs are represented by craniodental material and osteoderms only, but these specimens are significant as they document the first occurrence of the clade on Africa to the south of the Sahara. The combination of numerous phytosaur and lungfish discoveries with sedimentological evidence indicates that the Late Triassic and Early Jurassic sites in this area were deposited in more mesic environments than their lateral equivalents in South Africa.
Bite performance in Tyrannosaurus rex: Consensus or controversy?
 
*
Karl T Bates
1
 
Peter L Falkingham
2
 
1
University of Liverpool
 
2
Liverpool John Moores University
Bite performance predictions for Tyrannosaurus rex vary widely, despite recent modelling studies using near-identical methods. Here we examine previously proposed explanations for the disparity in estimates, in addition to presenting new analyses of model input data. We find no support for suggestions that the highly disparate estimates are the result of muscle volume reconstructions or specific assumptions of muscle contraction dynamics in earlier studies: in both instances standardising these parameters across studies increases, rather than decreases, the difference in predicted bite force. New analyses of model inputs unambiguously indicate that the difference between studies results from subjective selection of muscle fibre lengths; longer fibre lengths in more recent studies result in smaller muscle cross sectional areas and subsequently lower muscle and bite force estimates. To examine this specific issue, and to guide future estimates of muscle fibre lengths in extinct animals more generally, we analysed the ratio of muscle fibre length (FL) to muscle length (ML) in over 1200 muscles of extant terrestrial vertebrates from various body regions. We find that FL:ML varies widely within and across body regions, but with strong convergence on a mean ratio of 0.45-0.55. Application of this ratio to T. rex jaw muscles results in higher-end muscle and bite forces favoured by earlier studies, though considerable uncertainty remains. Future work might constrain higher-end estimates through further analysis of jaw muscle architecture in living vertebrates and by incorporation of bone loading limits into computer simulations.
Adaptive radiation of Pelagia (Teleostei: Acanthomorpha) indicated by 3D morphometry
 
*
Hermione Beckett
1
 
Zerina A Johanson
2
 
Sam Giles
1
 
Matt Friedman
1,5
 
1
University of Oxford, Department of Earth Sciences, South Parks Road, Oxford, OX1 3AN
 
2
Department of Earth Sciences, The Natural History Museum, Cromwell Road, London SW7 5BD, UK
 
5
Museum of Paleontology and Department of Earth and Environmental Science, University of Michigan, 1109 Geddes Ave, Ann Arbor, MI 48109-1079, USA
Pelagia or Pelagaria is an anatomically disparate clade of open-ocean fishes identified through molecular studies. It has been postulated that the ecologically distinct lineages of Pelagia diverged rapidly early in the group’s history, perhaps shortly after the end-Cretaceous extinction. This differs from model ‘adaptive radiations’ like cichlid fishes and Anolis lizards in both age and geographic scale, but corresponds closely to the pattern for which the concept was first defined: Cenozoic mammals. We collected computed tomographic data for the crania of 56 extant species of Pelagia, sampling all 14 confirmed extant families and a quarter of living species. We use three-dimensional geometric morphometrics to generate a shape space for this sample of skulls. Cranial anatomy clearly separates families within Pelagia, paralleling conspicuous differences in body shape. Of note are the gempylids (snake mackerels) and trichiurids (scabbardfishes), which have elongate skulls, and the deep-sea chiasmodontids (black swallowers) with largely reduced opercular series, posteriorly directed suspensoria, and long gapes. We conducted a disparity-through-time analysis based on these shape data and existing molecular phylogenies to test for deviations from a Brownian motion model of phenotypic evolution. We find substantially lower-than-expected levels of subtree disparity early in the history of the clade, a pattern often interpreted as consistent with theoretical models of adaptive radiation. These results provide quantitative support for an adaptive radiation in Pelagia early in the Cenozoic. This morphometric framework will allow the inclusion of three-dimensionally preserved fossils of this group, allowing us to further constrain patterns of phenotypic diversification in Pelagia.
New data on Moeritherium from the Birket Qarun Formation (Eocene), Fayum Depression, Egypt
 
*
Gemma Louise Benevento
1
 
William Sanders
2
 
Matt Friedman
1,2
 
Erik R Seiffert
5
 
1
University of Oxford, Department of Earth Sciences, South Parks Road, Oxford, OX1 3AN
 
2
University of Michigan, Museum of Palaeontology, 1109 Geddes Road, Ann Arbor, MI, 48109
 
5
University of Southern California, Department of Integrative Anatomical Sciences, Keck School of Medicine of USC, LA, California, 90033
Moeritherium is an early diverging proboscidean from the late Eocene-early Oligocene of Africa. Unlike other proboscideans, Moeritherium was characterised by a relatively small body size, a likely semiaquatic lifestyle, and is often reconstructed with either a small, tapir-like proboscis, or lacking one altogether. At least three species are currently recognised from isolated teeth, partial skulls and partial postcrania. Here we present new cranial and mandibular material of Moeritherium from the early Priabonian (~37Ma) of Birket Qarun, Fayum Depression, Egypt, increasing our knowledge of its anatomy. These fossils include the oldest known cranial and mandibular material of Moeritherium. The skull appears distinct from previously described cranial material, but the teeth are morphologically similar to isolated teeth described from the broadly contemporaneous Bir El Ater locality, Algeria, suggesting an affinity with M. chehbeurameuri. We present the first digital reconstruction and analysis of the internal structures of the skull and mandible of Moeritherium based on µCT data for this specimen. From the cranium, a near complete three-dimensional model of the endocast was produced, permitting investigation of brain anatomy. Within the mandible, of particular interest is the large size of the mandibular canal, which extends labially to the tooth row, and occupies much of the internal space within the mandible. Among Paenungulata, sirenians share this feature with Moeritherium. Extant proboscideans, however, have smaller mandibular canals, more similar to hyraxes, indicating a complicated evolutionary—and perhaps functional—history of this feature.
The functional morphology of the carnivoran baculum
 
*
Charlotte Brassey
1
 
1
Manchester Metropolitan University
The mechanical behaviour of an individual’s genitals directly impacts upon their fitness. If genitalia cannot function mechanically, the individual cannot deliver/receive sperm, and cannot produce viable offspring. Within carnivores, males possess a mineralized element within the glans of the penis, which is extremely morphologically diverse. Baculum size and shape has previously been found to be under direct sexual selection. Yet the adaptive advantage of such a structure remains unclear.

I present a comparative analysis of modern carnivoran bacula with the aim of testing functional hypotheses, including their role in facilitating intromission, prolonging copulation duration and/or removal of competitors’ sperm. 3D models of museum specimens (n=77, families=11) were generated from microCT. In the absence of numerous definitive homologous landmarks for GMM, alternative means of quantifying ‘shape’ are explored, including elliptical fourier analysis, thickness mapping and curvature analysis. Biomechanical performance is derived from modified beam theory.

All shape metrics are found to contain a strong phylogenetic signal. Curvature does not scale to body mass (r2=0.01, p>0.05), and baculum ‘relative strength’ decreases in larger taxa (r2=0.12, p=0.002). Baculum tip is more variable in shape than the shaft (t=4.97, p<0.001). Tip shape correlates to relative testes mass (a proxy for post-copulatory sexual selection; p=0.013, λ=-0.56), suggesting a role for the tip in biasing paternity via sperm competition or cryptic female choice.

Future research will quantify the rate of shape evolution in the tip versus base. Additionally, cadaveric material will be incorporated in order to consider the biomechanics of the whole soft tissue structure.
A re-examination of Milosaurus mccordi, and the evolution of large body size in Carboniferous synapsids
 
*
Neil Brocklehurst
1
 
Jörg Fröbisch
1
 
1
Museum für Naturkunde - Leibniz-Institut für Evolutions- und Biodiversitätsforschung
Milosaurus mccordi was described in 1970 as a large pelycosaurian-grade synapsid from the late Carboniferous of Illinois, but has since received little attention. Here the holotype and referred material of Milosaurus is re-examined and incorporated into a recent phylogenetic analysis. Milosaurus is found to be the sister to Sphenacomorpha, sharing with this clade the posterodorsally expanded ischium and the calcaneum with a greater length than width, but the more plesiomorphic ischium indicates it is an outgroup to the clade rather than within it. Most of the referred material shows very little overlap with the holotype, and so most was not included in the cladistic analysis. However, what was originally described as a dorsal rib is judged to be a femur, sharing a highly distinctive morphology with the holotype.
With an estimated body mass of 41 kg, Milosaurus represents one of the largest Carboniferous synapsids. Large size evolved at least twice independently in Sphenacomorpha during the Pennsylvanian: once in Sphenacodontidae and once in Edaphosauridae. Using comparative phylogenetic methods, we re-examined body size evolution in sphenacomorphs with Milosaurus included. Milosaurus is found to represent an independent evolution of large size, but with its trait evolution considerably slower than was observed in the other sphenacomorph clades, indistinguishable from evolution under Brownian Motion. Nevertheless, as one of the earliest appearances of large body size in the amniote fossil record and the outgroup to the most diverse Paleozoic synapsid lineages, Milosaurus represents a crucial taxon for understanding early terrestrial ecosystems.
Making Morphology Move: XROMM Ventilation Kinematics of Extant Archosaurs and Reconstructing Rib Motion in Fossils
 
*
Robert Brocklehurst
1
 
Sabine Moritz
2
 
Jonathan Codd
3
 
Phillip L Manning
1,5
 
William I Sellers
1
 
Elizabeth Brainerd
2
 
1
School of Earth and Environmental Sciences, University of Manchester, Manchester, M13 9PT, UK
 
2
Department of Ecology and Evolutionary Biology, Brown University, Providence, RI 02912, USA
 
3
Faculty of Biology, Medicine and Health, University of Manchester, Manchester, M13 9PT, UK
 
5
Department of Geology and Environmental Geosciences, College of Charleston, Charleston, SC 29424, USA
Modern birds have a highly efficient respiratory system, and a highly derived ribcage with which they ventilate their lungs via costosternal pumping. However, understanding the co-evolution of ribcage morphology and ventilation mechanics in birds has been hindered by a lack of integration between kinematic and anatomical data. In particular, birds differ significantly from their nearest living relatives, the crocodilians, in terms of their costovertebral anatomy. This makes it difficult to directly compare patterns of rib motion, which in turn prevents reconstruction of motion in fossil archosaurs. Here, we employ a joint-based approach to represent both bone morphology and motion. In vivo rib kinematics were recorded during ventilation in living crocodilians and birds using XROMM (X-ray reconstruction of moving morphology), with motion measured relative to the articular surfaces of the costovertebral joint; axes of rib translation and rotation were defined with respect to specific homologous landmarks on the ribs and vertebrae. This approach then served as the basis for “scientific motion transfer”, applying motion patterns observed in modern archosaurs to their fossil relatives- specifically, non-avian dinosaurs- in a framework grounded in the anatomy of the costovertebral joint. Whether to apply a rib motion pattern more similar to birds or crocodilians to dinosaurs was determined through detailed morphometric analysis of the vertebrae and ribs. This approach represents a testable and repeatable way of predicting ventilation kinematics in extinct taxa, and so should greatly improve our understanding of respiratory evolution along the avian stem.
A case of mistaken identity: 'Latvius' obrutus and new dipnoans from the Frasnian of Stolbovo, Russia.
 
*
Tom Challands
1
 
Oleg Lebedev
2
 
Matt Friedman
3
 
Natasha Krupina
2
 
1
University of Edinburgh
 
2
Palaeontological Institute of the Russian Academy of Sciences
 
3
University of Michigan

The village of Stolbovo, western Russia has previously yielded a dipnoan fauna comprising the long-snouted Rhinodipterus stolbovi but also a three-dimensionally preserved otoccipital region of a skull previously identified as the osteolepiform ‘Latvius obrutus’. This identification was made from the density of lateral line canal pores in the skull roof bones in addition to what Vorobyeva described as the post-parietal, tabular and extratabular bones. Reexamination of this specimen does not recognise the osteolepiform dermal elements previously identified but rather a dipnoan configuration including a characteristic B-bone, lateral dermal elements and a dipnoan parasphenoid. MicroCT-scanning reveals an enlarged utricular recess in the vestibular system, the derived condition for lungfishes, and separate sacculae and lagenae also present in rhinodipterid and dipterid dipnoans. Phylogenetic analysis resolves ‘Latvius’ obrutus as the primitive sister taxon to the ‘phaneropleurid-fleurantiid’ clade – a clade comprising derived Middle-Upper Devonian lungfishes. Rhinodipterus stolbovi resolves among the more primitive griphognathids, typical Gondwanan forms, and is not a rhinodipterid. Additional isolated dipnoan skeletal elements from the site include tooth plates, parasphenoids and dermal elements and indicate at least an additional three lungfish taxa from Stolbovo. Of particular note is a ctenodiform toothplate more characteristic of Carboniferous lungfishes. The new material and analyses not only demonstrate increased lungfish diversity from the Frasnian of Baltica but also probable interchange between Gondwana and Baltica. Most importantly, Carboniferous-like forms appear more common in the Devonian than previously realised indicating that Carboniferous lungfishes likely represent a survival assemblage in addition to a new radiation of lungfish.

The effect of spatial fossil bias on dinosaur palaeodiversity estimates in the latest Cretaceous of North America
 
*
Alfio Alessandro Chiarenza
1
 
Philip D Mannion
1
 
Peter A. Allison
1
 
Dan J. Lunt
4
 
Paul J. Markwick
5
 
1
Department of Earth Science and Engineering, Imperial College London, South Kensington Campus, London, SW7 2AZ, UK
 
4
School of Geographical Sciences, Bristol University, University Road, Bristol, BS8 1SS, UK
 
5
Getech, Elmete Hall, Elmete Lane, Leeds, LS8 2LJ, UK
Analyses of dinosaur diversity patterns through time have offered different interpretations when focusing on time intervals of important biotic events, such as the lead up to the K-Pg mass extinction. Although the general consensus suggests a sudden demise at the K-Pg boundary, others have proposed a gradual, long-term diversity decline. One of the main methodological obstacles is the estimation of the impact of spatial bias when reconstructing diversity, especially where sampling is minimal or absent. A possible solution is the application of ecological niche modelling, which uses species occurrences and climatic data to reconstruct habitat distribution through space. We downloaded a comprehensive database (>4100) of latest Cretaceous dinosaur occurrences from the Paleobiology Database and combined this with a high-resolution atlas of palaeogeography. A time series analysis of species distribution models was developed from the early Campanian to the late Maastrichtian of North America. The ecological niches modelled on available outcrop area through time show a decline for herbivorous taxa from the Campanian to the Maastrichtian, with an initial increase followed by a slight decline for tyrannosaurids, mirroring trends based on previous diversity analyses. However, when continental projections are considered, taking into account the poorly sampled eastern half of North America, these patterns shift dramatically, with an overall increase in habitat suitability between the Campanian and Maastrichtian for all three clades. This novel approach suggests that dinosaur diversity was not in decline in the lead-up to the K-Pg boundary, and highlights the impact of spatial bias in interpreting macroevolutionary dynamics.
New Late Devonian lungfishes from Greenland morphologically and phylogenetically blur the D-C boundary
 
*
Jennifer A Clack
1
 
Timothy R. Smithson
1
 
Tom Challands
3
 
1
University of Cambridge
 
3
University of Edinburgh
Three small lungfish specimens were collected in 1987 from the late Famennian of Greenland by the Copenhagen-Cambridge expedition that also found the iconic specimens of Acanthostega. Study of these lungfish specimens including by microCT scanning show that they probably all belong to the same species. They show a number of derived characters uncommon in Devonian taxa, but which are more typical of Carboniferous examples. The most conspicuous of these is a long-stalked parasphenoid with a diamond-shaped anterior plate. In the new specimens, the stalk is about twice the length of the plate, as in many Carboniferous genera. This feature is also shown in an isolated parasphenoid figured by Lehman in 1959 and attributed to the early Famennian Oervigia. Future studies of his material are planned to determine whether our new material is also a species of Oervigia. Bayesian and parsimony-with-implied-weighting phylogenetic analyses place the new material among Carboniferous forms, notably with Ctenodus. Devonian taxa normally range through the base of the tree, with Carboniferous forms clustering together towards the top. In our analyses, only the Devonian Nielsenia, also from Greenland, falls within the Carboniferous cluster. Our new material thus demonstrates that more derived forms prefiguring those of the Carboniferous did exist in the Devonian, and help to fill the morphological and phylogenetic gaps and blur the stratigraphical boundary implied by the end-Devonian mass extinction. Some lungfish lineages, as with some tetrapods, appear not to have been as seriously affected by that extinction event as others.
Comparative skull biomechanics in Varanus and Salvator ‘Tupinambis’
 
*
Hugo Dutel
1
 
Alana Sharp
2
 
Marc E H Jones
3,4
 
Susan E Evans
2
 
Michael J Fagan
1
 
Flora Gröning
7
 
1
School of Engineering, Medical and Biological Engineering Research Group, University of Hull, Hull, UK
 
2
Research Department of Cell and Developmental Biology, University College London, London, UK
 
3
School of Earth and Environmental Sciences, The University of Adelaide, Adelaide, Australia
 
4
South Australian Museum, Adelaide, Australia
 
7
School of Medicine and Dentistry, University of Aberdeen, Aberdeen, UK
The lizard species Salvator “Tupinambis” merianae and Varanus ornatus evolved independently in South America and Africa but share similar ecology and feeding behaviour, despite having notable differences in their skull structure. Tupinambis has a compact, relatively short and wide snout, whereas that of Varanus is more slender and narrow. In addition, a postorbital bar (POB) is present in Tupinambis but absent in Varanus, and the former lacks the mid-frontal suture that is present in the latter. Here, we explore the biomechanical significance of these differences using 3D computer-based mechanical simulations based on micro-computed tomography, detailed muscle dissections, and in vivo data. First, we simulated muscle activity and joint-reaction forces during biting using Multibody Dynamics Analysis. Then, the forces calculated from these models were used as an input for Finite Element Analysis, to investigate and compare the strains of the skull in these two species. The effects of the presence/absence of structures, such as the POB, were investigated by constructing artificial models which geometry was altered. Our results indicate that strains in the skull bones are lower in Tupinambis than in Varanus, in particular at the back of the skull. The presence of a POB clearly reduces the strains in the bones during posterior biting in Tupinambis, but not in Varanus. Our results hence highlight how the morphological differences between these two taxa affect the mechanical behaviour of their respective skulls during feeding.
Ecomorphological disparity and evolution of marine reptiles of the Sub-Boreal Jurassic Seaway
 
*
Davide Foffa
1
 
Stephen L Brusatte
1,3
 
Mark T Young
1
 
1
School of Geosciences, University of Edinburgh
 
3
National Museums Scotland
The dominant consumers in Jurassic ocean ecosystems were distantly related clades of reptiles secondarily adapted to an aquatic lifestyle. Stratigraphic and fossil evidence suggests that, like their modern counterparts, different lineages such as plesiosaurs, ichthyosaurs, and thalattosuchian crocodylomorphs were able to coexist in the same environment. As in modern aquatic environments diet and trophic specialisation are the main drivers for convergent evolution, it is possible that marine reptiles occupied analogous ecological roles now held by cetaceans, sirenians, and large fishes such as sharks. However, these comparisons, supported by qualitative observations, have rarely been tested in a quantitative way. In this study we aim to rectify this using the rich fossil assemblage from the Middle-Late Jurassic Sub-Boreal Seaway (Callovian-Tithonian, 164–145 Mya). Using multivariate analyses, we analysed a novel dataset of continuous and discrete functionally relevant characters from the dentition and lower jaws of 62 specimens. Our results show that the transition from the Middle to the Late Jurassic involved a synchronous significant decrease in functional disparity for pliosaurids and teleosaurids, and an increase for ichthyosaurs and metriorhynchids. Late Jurassic metriorhynchids and ichthyosaurs shifted towards regions of ecomorphological space vacated by groups that had declined from the Middle Jurassic. Combined with novel fossil evidence from the so called ‘Corallian Gap’ (a poorly investigated interval of the Oxfordian deposited in between the fossil-rich Callovian and Kimmeridgian formations), our results indicate that a significant marine faunal turnover – maybe linked to environmental perturbations – affected the marine ecosystems through the early Late Jurassic.
A re-examination of Orovenator mayorum using μCT data, and its consequences for early amniote phylogeny.
 
*
David P. Ford
1
 
Roger B J Benson
1
 
1
University of Oxford, Dept. of Earth Sciences

Orovenator mayorum, from the early Permian (Sakmarian) of Oklahoma, has been described as the earliest representative of Neodiapsida, a clade that includes dinosaurs and birds, snakes, lizards, crocodiles and turtles. High-resolution CT scans of the two known specimens of this taxon reveal novel anatomical data. These include the arterial pathways in the basicranium and palate, the osseous canal enclosing the nasolacrimal duct, and well preserved septomaxillae. Exceptional detail of the cranial sutural anatomy is evident, including unusual medial buttressing on the lacrimal and maxilla which indicate a high structural integrity of the anteorbital region. By combining the elements of both specimens we have been able to reconstruct most of the skull in virtual 3-D, including a complete palate and basicranium. Several features of Orovenator are shared with Varanopidae, a generally accepted stem-ward clade of pelycosaur-grade synapsids. Substantial similarity between early diapsids and varanopid synapsids has previously been noted: four taxa currently assigned to the Varanopidae were originally described as diapsids (Archaeovenator, Heleosaurus, Apsisaurus) or archosauromorphs (Mesenosaurus). However, our study suggests that these similarities are greater than previously appreciated, raising questions about deep phylogenetic divergences within amniotes. Application of the new data on Orovenator into an existing phylogenetic matrix intended to discriminate between varanopids and diapsids resulted in an unresolved consensus. This has prompted work on a novel phylogeny, with greater taxonomic scope, to provide a working hypothesis for the interrelationships of Permo-Carboniferous amniotes.

Evolution of body size in Crocodylomorpha in a multi-regime evolutionary landscape
 
*
Pedro Godoy
1
 
Roger B J Benson
2
 
Mario Bronzati
3,4
 
Richard J. Butler
1
 
1
School of Geography, Earth and Environmental Sciences, University of Birmingham, Birmingham, United Kingdom
 
2
Department of Earth Sciences, University of Oxford, United Kingdom
 
3
Bayerische Staatssammlung für Paläontologie und Geologie, Staatlichen Naturwissenschaftlichen Sammlungen Bayerns, Munich, Germany
 
4
Department of Earth and Environmental Sciences, Palaeontology & Geobiology, Ludwig-Maximilians-Universität München, Munich, Germany
Body size is strongly related to physiology and ecology, and its evolution has been studied intensely for many animal groups. Crocodylomorphs occupy the intermediate–large range of body sizes among extant tetrapods, and have a rich and diverse fossil record, ranging in size from less than 30 centimetres to over 10 metres in length. To investigate crocodylomorph body size evolution, we used maximum-likelihood to fit macroevolutionary models to body length data within a phylogenetic context. We focused on non-uniform Ornstein-Uhlenbeck (OU) models (“SURFACE” model). Under an OU process, lineages are attracted towards an adaptive trait optimum (theta, θ) through time. SURFACE allows multiple adaptive regimes (i.e. shifts in trait optima). We also fitted other models, and conducted analyses using alternate body length proxies, tree topologies and time-calibration methods, to assess the influence of analytical choices on results. The SURFACE model fit best to our data, suggesting convergent evolution of body size among macroevolutionary adaptive zones. Although different trees identify regime shifts in phylogenetically distinct positions, we recognized some shared patterns. For instance, non-Mesoeucrocodylia crocodylomorphs maintained relatively small sizes (θ = 0.646 m), and experienced no or few shifts, thriving until the Late Cretaceous. During the Late Jurassic and Cretaceous, crocodylomorph body size disparity increased, with a high number of regime shifts, particularly within Notosuchia and Thalattosuchia. Another important regime shift, towards larger body sizes (θ = 5.561 m), occurred in the lineage leading to extant crocodylians (Eusuchia), potentially related to an adaptive radiation of the group after the Cretaceous-Paleogene extinction.
Using computational fluid dynamics to analyse the hydrodynamic properties of fossil marine reptiles
 
*
Susana Gutarra Diaz
1
 
Benjamin C. Moon
1
 
Imran A. Rahman
3,1
 
Stephan Lautenschlager
1,6
 
Alison J. Brimacombe
1
 
Colin Palmer
1
 
Michael J Benton
1
 
1
School of Earth Sciences, University of Bristol, Bristol, UK.
 
3
Oxford University Museum of Natural History, Oxford, UK.
 
6
School of Geography, Earth and Environmental Sciences, University of Birmingham, Birmingham, UK.
Various groups of reptiles recolonised aquatic environments and became apex predators of the Mesozoic seas; among them were the iconic ichthyosaurs, plesiosaurs and mosasaurs. During their evolution, their anatomies were subject to strong selective pressures imposed by the physical constraints of water, resulting in adaptations to locomotion such as the streamlining of body shapes, and the transformation of cursorial limbs into paddles or hydrofoils. Until now, the functional impact of these morphological changes has been difficult to study due to the lack of an objective, physics-based quantitative method.

Computational fluid dynamics (CFD) is a state-of-the-art engineering tool used to simulate fluid flows and calculate the forces resulting from fluid-solid interactions. I will present a case study in which this technique is used to analyse the effect of body plan evolution on the hydrodynamic properties of ichthyosaurs. Full-body reconstructions of various specimens were created using 3D modelling software, and the total drag, lift and the contribution of the limbs to these forces were evaluated by means of CFD. The results demonstrate that changes in the ichthyosaur body plan – from the narrow and elongated basal forms to the deep-bodied and highly streamlined parvipelvians – did not contribute to drag reduction, contrary to what has classically been suggested. The ecological relevance of these results will be discussed.

As a next step, and building upon this experimental strategy, I aim to evaluate the degree of functional convergence in hydrodynamic performance in the most derived species of Mesozoic marine reptiles, and also between these and aquatic mammals.
The Fort McMurray nodosaur – a three-dimensional dinosaur fossil with preserved scales, pigments, and stomach contents
 
*
Donald M Henderson
1
 
Caleb Brown
1
 
Jakob Vinther
3
 
1
Royal Tyrrell Museum of Palaeontology
 
3
University of Bristol. UK.
In the spring of 2011 a surprising discovery was made – a three-dimensional dinosaur fossil in the marine sediments of the Aptian-Albian age Clearwater Formation in northeastern Alberta, Canada. During the previous 20 years, two ichthyosaurs and nine plesiosaurs had been recovered from the tar sand mines in the area north of the town of Fort McMurray. Soon after the carcass arrived at the seabed, a very dense, well-cemented concretion grew around the carcass, and protected the specimen from subsequent crushing from overburden pressure over the next 112 million years. The only damage to the specimen occurred at the instant of discovery by heavy equipment, resulting in the loss of the tail and parts of the limbs. The concretion had the unfortunate side-effect of preventing permineralization of the armour and skeleton, leaving the fossil extremely soft. The unscavenged, uncrushed specimen shows all the dermal armour in place and in association with the underlying skeleton. Patterns of inter-locking, large polygonal scales, preserved as carbon films, demonstrate the relationships between osteoderms and squamation. Fossil pigment allows for the interpretation of a counter-shading on the body. The specimen arrived at the seabed on its back, and the rotting viscera collapsed downwards to rest on the spine. A collapsed, football-sized volume of densely packed, pea- to grape-sized concretions are interpreted to be animal’s last meal. The preserved armour and skin are so extensive that skeletal details are largely hidden from view.
The first virtual cranial endocast of a porolepiform fish and the evolution of the Dipnomorpha
 
*
Struan Henderson
1
 
Tom Challands
1
 
1
University of Edinburgh
The Dipnomorpha include the extinct Porolepiformes, in addition to Powichthys and Youngolepis and the extant Dipnoi (lungfish). As sister group to the Tetrapodomorpha the Dipnomorpha hold a valuable place in our understanding of early sarcopterygian evolution. With complete cranial endocasts now known from most other stem sarcopterygian groups including actinistians and tetrapodomorphs (which bracket the Dipnomorpha), a thorough understanding of the porolepiform endocranium remains one of the last pieces in the puzzle towards understanding evolution of the neurocranium in stem sarcopterygians.

We present the first virtual cranial endocast of a porolepiform fish (Glyptolepis paucidens) demonstrating that it displays predominantly primitive sarcopterygian endocast characters including: lack of a bifurcation of the olfactory nerves, separate pineal and parapineal recesses, circumvention of the nasal capsule by the orbitonasal canal, sessile olfactory bulbs and lack of ventral expansion of the telencephalon. The union of the profundus nerve and the nasal capsule is consistent with that seen in Powichthys, Youngolepis and the primitive dipnoan Dipnorhynchus.

Inconsistent with the general primitive nature of the endocast seen in Glyptolepis is the large, curved hypophyseal recess, a feature shared with Powicthys but unlike Youngolepis. Further similarity with Powichthys is found in the size and direction of the profundus canals suggesting greater endocast similarities of Powichthys with the Porolepiformes than previously assumed.

This new data enhances our understanding of brain and sensory evolution in sarcopterygians. In particular, it shows conservatism in stem Porolepiformes contrary to the liberal morphology of their sister-group the Dipnoi.
Fusion in the vertebral column of the pachyosteomorph arthrodire Dunkleosteus terrelli ('Placodermi')
 
*
Zerina A Johanson
 
Kate Trinajstic
1,2
 
Michael Ryan
3
 
Stephen Cumbaa
4
 
1
Department of Environment and Agriculture, Curtin University, Perth, Western Australia
 
2
Earth and Planetary Sciences, Western Australian Museum, Perth, Western Australia
 
3
Department of Vertebrate Paleontology, Cleveland Museum of Natural History, Cleveland, Ohio;
 
4
Paleobiology, Canadian Museum of Nature, Ottawa, Ontario, Canada

Fusion in the vertebral column occurs normally in jawed vertebrates, with structures like the sacrum and pygostyle providing rigidity, support and increased area for muscle attachment. The synarcual represents the fusion of the anterior part of the vertebral column and occurs in a number of jawed vertebrates, including a variety of placoderms and chondrichthyans. Placoderms are an entirely fossil group of armoured fishes (Silurian-Devonian), resolved to the base of the jawed vertebrate clade, with vertebrae comprising neural and haemal arches composed of perichondral bone. The placoderm synarcual preserves substantial developmental information from anterior (oldest) to posterior, where new vertebrae are incorporated. A subadult specimen of the pachyosteomorph arthrodire Dunkleosteus terrellli preserves vertebral elements showing varying degrees of anteroposterior fusion along the vertebral column. Synarcuals are rarely preserved in this group, and micro-CT scanning of this synarcual provides details of a transitional zone of vertebral fusion, providing unprecedented information on how each vertebrae is modified and incorporated into the synarcual. All elements in the synarcual retain vertebral identity, showing less fusion overall, more comparable to other arthrodires such as Compagopiscis. By comparison, synarcuals of other taxa such as ptyctodont placoderms, batoids, holocephalans (Chondrichthyes) and mammals (syncervical) show more complete fusion of vertebral elements.

Early Cambrian ostracoderms and the trials and tribulations of total evidence dating
 
*
Joseph Keating
1,2
 
Richard Dearden
1,4
 
Philip C.J. Donoghue
1
 
1
University of Bristol
 
2
University of Manchester
 
4
Imperial College London
The origin of jawed vertebrates is a formative episode in evolutionary history. Key to understanding this important interval are the armoured jawless ‘ostracoderms’, the relationships among which have previously been considered either in terms of clade intrarelations, or interrelations among clade exemplars. Thus, the monophyly of the putative ostracoderm plesions have yet to be adequately tested, principally because of computational limitations surmounted in the last decade. We constructed a supermatrix
of 560 characters and 240 taxa, which we analysed using the MK likelihood model. The ensuing trees were compared using Bayes factors. Our results support the majority of ostracoderm clades, but resolve galeaspids and heterostracans as paraphyletic grades of stem-gnathostomes, and stem-vertebrates, respectively. Finally, we considered the role of time in topology estimation, using the fossilized birth-death model (FBD). This approach resolved heterostracans, thelodonts and anaspids as a single clade, while suggesting that ostracoderms evolved in the earliest Cambrian. This result con icts strongly with the fossil record, suggesting that the FBD method may incorrectly estimate divergences of extinct taxa. Taken together, these results suggest that the consensus of early vertebrate relations, underpinned by the last 40 years of cladistics, does not stand up to deeper scrutiny afforded by likelihood-based phylogenetics.
The Recent Now: Post-Medieval Wood Mice
 
*
Elizabeth Kerr
1,2
 
Raphäel Cornette
3
 
Silvia Valenzuela
4,5
 
Benoît Clavel
2
 
Jean-Claude Marquet
7
 
Antony Herrel
1,9
 
Anne Tresset
2
 
1
UMR 7179 CNRS/MNHN, 57 rue Cuvier, Case postale 55, 75231, Paris Cedex 5, France.
 
2
UMR CNRS/MNHN 7209, 55 Rue Buffon, Case Postale 56, 75005 Paris, France
 
3
UMR 7205 CNRS/MNHN/UPMC/EPHE, (ISYEB), 45 rue Buffon, 75005 Paris, France
 
4
Universitat de Barcelona, (GRACPE), C/ Montalegre 6, E-08001 Barcelona, Spain
 
5
Universitat de Barcelona, (SERP), C/ Montalegre 6, E- 08001 Barcelona, Spain
 
7
University of Tours, Tours, France
 
9
Ghent University, Evolutionary Morphology of Vertebrates, K.L. Ledeganckstraat 35, B-9000 Gent, Belgium.
During the post-medieval era in Western Europe, arable land expanded and agricultural practices homogenised ecological and landscape diversity. There were also highly increased rates of vertebrate disappearance and extinction. Anthropogenic species such as wood mice (Apodemus spp.) have persisted, despite the increased intensity in human impacts. An analysis of Apodemus mandible shape variation in France does however show a dramatic shift in mandible shape between the end of the 16th century and the present day. This change is more distinctive than all the mandible shape variability present in the preceding 12,000 years. A biomechanical model suggests that present-day wood mice are less capable of producing a higher bite force. Implications for function probably include a reduced dietary range, for example due to restricted access to diverse natural resources or the increasing availability of homogenous anthropogenic resources. The changes in landscape structure therefore appear to have imposed new functional requirements on native species. These results indicate that there are pervasive changes in ecosystem function in response to human impacts from the post-medieval era. Comprehensive datasets of archaeological small mammal material therefore provide crucial insight into organism function as well as the dramatic impact of ecological and landscape changes during this transformative episode of the European Holocene.
Opportunities in disasters: molecular evidence for the rapid radiation of snakes following the Cretaceous-Paleogene mass extinction
 
*
Catherine Klein
1
 
Daniel J. Field
1
 
Davide Pisani
3
 
Nicholas R Longrich
1
 
1
Department of Biology & Biochemistry, and Milner Centre for Evolution, University of Bath, 4 South, Bath BA2 7AY, United Kingdom
 
3
School of Biological Sciences, and School of Earth Sciences, University of Bristol, Life Sciences Building, 24 Tyndall Avenue, Bristol BS8 1TQ, United Kingdom
Mass extinction events have shaped global biodiversity throughout Earth's history. The most recent major mass extinction, at the Cretaceous-Paleogene (K-Pg) boundary, took place 66 Mya and has been particularly well studied. This event witnessed the demise of non-avian dinosaurs, pterosaurs, and mosasaurs, amongst numerous other major groups. However, it also provided an opportunity for significant ecosystem restructuring, and has been suggested to have stimulated the rapid diversification of mammals, teleosts, frogs, and birds in its aftermath. The influence of the K-Pg mass extinction on the evolutionary history of snakes is less well understood, however. To address this understudied topic, we used a molecular clock approach to show that Alethinophidia, the clade comprising the majority of modern snake diversity and disparity, also underwent a rapid diversification in the aftermath of the K-Pg mass extinction event. We used a novel supermatrix representing 169 species coded for up to 52 loci, and a new set of 42 phylogenetically and stratigraphically constrained fossil calibration points. Divergence time analyses were performed under a variety of parameterisations in PhyloBayes and PAML, and numerous outgroup taxa were included to control for rate heterogeneity within the ingroup. Our inferred association between the K-Pg event and the rapid diversification of Alethinophidia provides further evidence of the importance of this mass extinction in shaping Earth’s modern vertebrate faunas. Our analyses also imply that crown snakes only explored arboreality after the K-Pg, suggesting some degree of ecological filtering induced by the mass extinction event.
The role of miniaturisation for cranial function across the cynodont-mammaliaform transition
 
*
Stephan Lautenschlager
1
 
Pamela Gill
2
 
Zhe-Xi Luo
3
 
Michael J Fagan
4
 
Emily J Rayfield
2
 
1
School of Geography, Earth and Environmental Sciences, University of Birmingham, UK
 
2
School of Earth Sciences, University of Bristol, UK
 
3
Department of Organismal Biology and Anatomy, University of Chicago, USA
 
4
School of Engineering, University of Hull, UK

The evolution of mammals from cynodonts is a key event in vertebrate history and is characterized by a number of modifications to the cranial skeleton. This is most prominently expressed in the emergence of a novel jaw joint and the reduction of the mandible to a single tooth-bearing bone in crown mammals. These skeletal changes are generally thought to be allied with increasing cranial strength in response to a more powerful and complex jaw musculature, and the evolution of a more ‘efficient’ feeding system. Using an integrated suite of digital reconstruction, visualization and biomechanical modelling techniques (FEA, MDA), we studied six key taxa (including Thrinxaodon, Probainognathus, Morganucodon, and Hadrocodium) across the cynodont-mammaliaform transition. This allowed testing the hypothesis that during feeding the cranial skeleton of cynodonts and mammaliaforms experienced progressively lower magnitudes of stress and strain and increased bite forces. Results show a slight decrease in average stress and strain across taxa, but do not support a large-scale trend for the increase of cranial strength towards crown mammals. However, we find that a reduction in size of the cranial system, as observed across the cynodont-mammaliaform transition, results in opposing trends of reducing bite forces, but also a substantial reduction of compressive and tensile stresses in the jaw joint. This suggests that presumed changes in the loading regime and/or the cranial morphology did not act as the main trigger for the emergence of a novel jaw joint. Rather, miniaturisation provided an evolutionary environment, in which substantial morphological modifications were possible.

Hips, Tips and Sweet Sweptback Rays: Looking beyond traditional cranial characters in Pachycormiformes
 
*
Jeff J Liston
1,2,3
 
Anthony Maltese
4
 
1
Bayerische Staatssammlung für Paläontologie und Geologie Richard-Wagner-Str. 10 80333 München,, Germany
 
2
National Museums Scotland, Chambers Street, Edinburgh, Scotland
 
3
Yunnan Key Laboratory for Palaeobiology, Yunnan University, North Cuihu Beilu 2#, Kunming, Yunnan Province, 650091, People’s Republic of China
 
4
Rocky Mountain Dinosaur Resource Center, Woodland Park, CO, United States of America, 80863

A focus on cranial characters for determining relatedness is a predominant trait in many datasets, however this overemphasis can lead to distortion from sampler bias. We report on revised analyses of pachycormids - a key group within Actinopterygii, as part of the Holostei-Teleostei Transition, which display a phyletic trend towards reduced skeletal ossification with the increased adult size of a pachycormid taxon. This reduced preservation potential for the axial skeleton makes it difficult not to base phylogenetic assumptions primarily on the limited skull material present. However, pachycormids show a remarkable conservatism in their dermatocranial anatomy, the few differences being useful for showing the separation of genera, but of little utility in working out broader intrafamilial relationships. The combination of a paucity of postcranial characters in the Late Cretaceous pursuit predator Protosphyraena with a poor knowledge about the skulls of suspension-feeding pachycormids (SFPs) had led to the absence of Early Cretaceous predatory pachycormids being interpreted as indicating a ghost lineage between Protosphyraena and the European Upper Jurassic taxa Orthocormus and Hypsocormus over an almost 50 million year gap. However, the inclusion of several features from the pectoral and pelvic fins, supplemented by splanchnocranial characters, produces a much clearer picture that questions the traditional perception of a single carnivore lineage: Protosphyraena emerges as secondarily carnivorous from the SFPs’ tribe, mirroring 130 years of misidentification of North American Bonnerichthys specimens as Protosphyraena. Confirmation of this will rely on the further recovery of data concerning the skull morphology of SFPs.

High Diversity of of Small Dinosaurs Preceding the Cretaceous-Paleogene (K-Pg) Mass Extinction
 
*
Nicholas R Longrich
1
 
1
University of Bath
Dinosaurs dominated terrestrial ecosystems for over 100 million years, before disappearing at the end of the Cretaceous. Based on the fauna of the Western Interior of North America, it has been assumed that latest Cretaceous faunas were low in diversity and dominated by large species, consistent with the hypothesis that there was a decline in dinosaur diversity preceding the K-Pg boundary that left dinosaurs vulnerable to extinction. However, small dinosaurs are rare and tend to be understudied, raising the possibility their apparent low diversity is an artifact of sampling and study biases. Here, it is shown that alongside the giant T. rex and Triceratops, a diverse fauna of small dinosaurs thrived in the late Maastrichtian, including previously unrecognized taxa of Dromaeosauridae, Troodontidae, Caenagnathidae, Alvarezsauridae, Thescelosauridae and Leptoceratopsidae. Total dinosaur diversity includes at least 29 species ranging from 2 kg to 50,000 kg, occupying carnivorous, herbivorous, insectivorous, and piscivorous niches. An analysis of functional diversity, using a combination of behavioral characters, morphological characters chosen for functional significance, and body size shows that ecological niche occupation increased rather than decreased from the Late Campanian to the Late Maastrichtian in the Western Interior, driven by the immigration of new taxa such as alvarezsaurids and titanosaurians. These patterns reject the hypothesis of a dinosaur decline, and suggest that the dinosaur fauna remained ecologically diverse until the end of the Cretaceous. These findings are consistent with a catastrophic extinction at the end of the Maastrichtian coinciding with, and driven by, the Chicxulub asteroid impact. The extinction of small dinosaurs also suggests that while size was a major contributing factor, size selectivity alone cannot explain dinosaur extinction.
Spatiotemporal patterns in fossil sampling bias and their impact on understanding the evolutionary history of terrestrial crocodylomorphs
 
*
Philip D Mannion
1
 
Alfio Alessandro Chiarenza
1
 
Pedro Godoy
3
 
Jonathan Tennant
1
 
Yung Nam Cheah
1
 
1
Imperial College London
 
3
University of Birmingham
The 24 species of living crocodylians are the remnants of a once much more diverse and widespread clade. Crocodylomorpha has an ~230 myr evolutionary history that was punctuated by a series of radiations and extinctions. Whereas previous studies have reconstructed sampling-standardised crocodylomorph diversity, the completeness of the fossil specimens themselves provides additional information that is not captured in those approaches. We compiled a dataset of all taxonomically diagnosable, non-marine species (393) of Crocodylomorpha, including fossil occurrences of extant taxa. Based on the number of phylogenetic characters (484) that can be scored for all known fossils of each species, we calculated a completeness value for each taxon. Average species completeness is 55%, and this value is largely consistent within subgroups (e.g. Crocodylia, Notosuchia, non-neosuchians) and for different body size classes, suggesting no significant biases across the crocodylomorph tree. In general, average completeness values are higher in the Mesozoic (60%) than the Cenozoic (50%). Completeness decreases across the Cretaceous/Paleogene boundary, which might explain why an uncorrected census of taxonomic diversity shows a crash at this mass extinction, whereas a sampling standardisation approach reveals little change in overall standing diversity. There is no evidence for a Pull of the Recent effect, with lower average completeness during the last ~10 myr, although many extant taxa are identified based on very incomplete fossil remains. Spatiotemporal sampling bias impedes our understanding of some Mesozoic radiations, including Neosuchia and Notosuchia, whereas divergences within Crocodylia based on molecular data are generally in close agreement with the fossil record.
Grabbing evolution by the throat: functional regionalisation of the avian cervical column
 
*
Ryan Marek
1
 
Peter L Falkingham
2
 
Karl T Bates
1
 
1
University of Liverpool
 
2
Liverpool John Moores University

Birds have evolved powerful forelimbs that are used primarily in flapping flight. This has rendered these appendages less useful for environmental manipulation than in many of their dinosaurian antecedents. Reliance on the head and neck for feeding and environmental interaction places high selective pressure on cervical form-function, potentially explaining the diversity in neck morphology seen in birds. However, to-date there has no been no systematic study of morphological diversity in the avian neck and its correlation with feeding habits. This study uses a combination of three-dimensional geometric morphometrics (GMM) and qualitative character coding to assess regionalisation within the cervical column of a wide variety of extant birds. These species represent a large diversity of feeding (carnivores, seed eaters) and functional (swimmers, flyers and terrestrial) ecology, cervical count (12-17) and body size. Results provide strong support for 5 cervical subregions (axis, anterior, middle, midposterior, posterior) in all species. The atlas subregion appears to show the strongest signal, with the axis (cervical 2) being clearly separate in all studied birds, possibly owing to its function into head stabilisation. Other subregions with a reasonably stable cervical count (anterior and posterior) also display a clear functional role. The remaining 2 regions (middle, midposterior) show much variability in cervical count between species (middle 2-6 cervicals, midposterior 1-4 cervicals). These results suggest that whilst the underlying Hox genetics may restrict avians to 5 cervical subregions, expansive variability in the middle and midposterior regions allow the cervical columns of birds to adapt to many different functional ecologies, and may be responsible for the large variety of neck morphologies observed in extant Aves.

Pterosaur wing bone geometry and its relationship to pneumaticity and palaeoecology
 
*
Elizabeth Martin-Silverstone
1,2
 
1
University of Southampton
 
2
University of Bristol. UK.
Pterosaurs are typically portrayed as having extremely thin-walled wing bones, much thinner than birds, which is assumed to lower mass, assisting large bodied forms in take-off and flight. However, there is a trade-off as thinner-walls are more likely to buckle or bend. Smaller, basal taxa are called ‘thick’-walled, while more derived pterodactyloids are described as thinner-walled, suggesting a size constraint. Here I present the first phylogenetically-grounded cross-clade study of pterosaur wing bone geometry. I test the phylogenetic affinity of pterosaur bone thickness, and its relationship to both pneumaticity and palaeoecology, using 120 wing bones. Cortical thickness (t) and diaphyseal radius (R) of wing bones was measured in order to characterize variation using R/t and K-values (inner to outer bone radius), and bending stiffness was estimated using second moment of area (I). Contrary to previous studies, wing bone thickness does not carry a phylogenetic signal, nor vary consistently between or within groups. R/t values similar to those seen in modern birds are common, with relatively few large-bodied pterodactyloids reaching extreme values upwards of 15. Incorporating size reveals a phylogenetic signal in K and R/t values. Bending stiffness varies from 30 to over 1 000 000 m4, increasing with bone size. These data show that smaller-bodied forms, were resistant to impact, compression and buckling, typical of animals frequently taking-off and landing. Conversely, larger pterosaurs were optimized for mass reduction and bending resistance, essential for wings under high loads. R/t values may indicate if a bone was pneumatic, but more data is needed.
New onychodontid (Osteichthyes ; Sarcopterygii) remains from the Middle Devonian of Morocco
 
*
Jorge Mond�ar Fern�ndez
1
 
1
Muséum national d'Histoire naturelle (MNHN) of Paris
Onychodontids are a specialized and phylogenetically important group of marine sarcopterygians (lobe-finned fishes), probably closely related to coelacanths. However, they are among the least well-understood groups of early osteichthyans (bony fishes). The principal reason for this is their partly cartilaginous and therefore perishable endoskeleton, which causes the bones of the head to disarticulate and scatter after death.
A new onychodontid species from the Eiffelian (Middle Devonian) of Morocco was found 40 years ago but never described in detail. New preparation of the specimens has revealed new features and bones that were not previously known. These fossil remains can be attributed to a single individual and have an extraordinary scientific potential: dermal skull bones are well preserved and allow a partially complete reconstruction of the snout, cheek and skull roof. Moreover, endoskeletal branchial and fin bones are exceptionally preserved, adding to our knowledge of early osteichthyan endoskeleton.
Preliminary phylogenetic placement supports its basal position among onychodontids, enabling to test evolutionary scenarios among the clade. This new species, along with well-preserved onychodontid fossils from Euramerica (Strunius from Germany) and Gondwana (Qingmenodus from China and specially Onychodus from Australia) adds to our knowledge of these elusive fishes. The Moroccan specimens, representing the first occurrence of onychodontids from Africa (Northern margin of Gondwana), thus constitute important material from a new locality that would furnish key information not only on onychodontid morphology and interrelationships, but also on their paleobiogeographical distribution and Devonian faunal affinities between Euramerica and Gondwana.
The Biomechanical Significance of the Rodent Mandibular Incisor “Root” – A Finite Element Analysis Study
 
*
Philip J R Morris
1
 
Philip G Cox
1,3
 
Samuel N Cobb
1,3
 
1
Centre for Anatomical and Human Sciences, Hull York Medical School, University of Hull, UK.
 
3
Department of Archaeology, University of York and Hull York Medical School, York, UK.
Rodents are an order of mammals characterised by a distinctive masticatory apparatus which includes a pair of enlarged and continually growing incisors, a feature termed diprotodonty. Diprotodonty is present in all living and extinct rodents and has independently evolved in a phylogenetically diverse range of non-rodent mammals; including the aye-aye, a primate which we have previously shown to be highly convergent in skull morphology to rodents.

This study examined the functional significance of the incredibly long rodent-like incisor. A sample of four rodents, and the non-rodent diprotodont aye-aye, were segmented to exhibit incrementally shorter internal incisor lengths, with variant models containing a solid crypt and a hollow crypt, and examined using finite element analysis to examine the resultant differences in principal strains. The primary aim of this study was to elucidate whether the internal length of the incisor is functionally significant to the resistance of mechanical forces encountered during incision.

The findings of this research indicate that strain increases substantially as a consequence of shortening the internal length of the incisor, with strain increasing by up to 35%. This study thus finds that the long internal incisor plays a substantial role in the resistance of bending forces, both by itself, and in tandem with an arch-like mandibular shape - itself a result of a long and curved internal incisor. Consequently, this study proposes that the evolution of these features together aid in strengthening the rodent-like mandible against the repetitive loadings and high forces encountered during the iconic rodent behaviour – gnawing.
A virtual reconstruction and volumetric mass estimate of AL288-1, Australopithecus afarensis
 
*
Thomas O’Mahoney
1
 
Charlotte Brassey
2
 
Andrew Chamberlain
1
 
William I Sellers
1
 
1
University of Manchester
 
2
Manchester Metropolitan University
Fossil body mass estimation is a well-established practice within the field of physical anthropology. Previous studies have relied upon traditional allometric approaches, where the relationship between skeletal dimensions and body mass in a range of modern taxa is used predictively. The lack of relatively complete skeletons has limited the potential application of alternative mass estimation techniques, such as volumetric reconstruction, to fossil hominins. Yet across vertebrate palaeontology, volumetric approaches are increasingly predicting values for fossil body mass very different to those estimated by traditional allometry. Here we present the results of a virtual reconstruction exercise for the iconic hominin AL288-1 and the calculation of its body mass, created using a combination of differing morphometric techniques. These reconstructions represent working hypotheses of the morphology of AL288-1, based upon what is currently known about Australopithecus anatomy.

The convex hull technique relies upon identifying a predictable relationship between the ‘shrinkwrapped’ volume of the skeleton and known body mass in a range of modern taxa, and subsequent application to an articulated model of the fossil taxa of interest. Here, our calibration dataset comprises whole body CT scans of 15 species of modern primate. Application of the convex hull technique to A. afarensis results in a relatively low body mass estimate of 20.1kg (95% prediction interval 13.3-30.6kg). A sensitivity analysis on the thorax, performed by 10 and 20% expansion of this area, suggests that the heaviest of previous body mass estimates would require the thorax to be expanded to an unlikely extent.
The dentary of Wareolestes rex and new fossil mammal material from the Middle Jurassic of Skye, Scotland
 
*
Elsa Panciroli
1,2
 
Roger B. J. Benson
3
 
Stig. A Walsh
2
 
1
University of Edinburgh
 
2
National Museum of Scotland
 
3
University of Oxford
The Middle Jurassic was a pivotal time for many vertebrate groups: recent research suggests mammals experienced a surge in ecological diversity at this time. However, vertebrate fossils from the Middle Jurassic are rare. Until now, Middle Jurassic mammals in the UK came almost exclusively from Bathonian deposits in England, while globally the most complete specimens are mainly found in China. Ongoing fieldwork on the Isle of Skye suggests this locality is of international significance for microvertebrate skeletal remains. So far, Skye has yielded skeletal associations and fragmentary material including early mammals, tritylodontids, salamanders and basal squamates.

We report the most complete specimen of the Middle Jurassic morganucodontan Wareolestes rex, from the Bathonian Kilmaluag Formation of Skye, Scotland. The specimen was digitally reconstructed using microCT scan data. It comprises a partial left dentary with two erupted molars, one unerupted molar, and three unerupted premolars. Empty alveoli for a canine, p1 and p3 are also present.

Wareolestes was previously known from four isolated molars from Kirtlington, England, and there was debate over the position of the holotype tooth as an upper or lower molar. Comparing our new material with the holotype, we support the original diagnosis of the holotype as a lower molar, most likely m1. In the Scottish specimen, unerupted and erupted premolars, and presence of permanent molars, supports diphyodonty in Wareolestes. This has previously been suggested for other morganucodontans. Damage to the dentary of Wareolestes means questions remain regarding the sequence of replacement along the tooth row in this genus.
A novel phylogeny for the Heterostraci: evolutionary relationships of extinct jawless vertebrates on the gnathostome stem
 
*
Emma Randle
1
 
Robert S Sansom
1
 
1
University of Manchester
Heterostracans - the largest clade of extinct boney jawless vertebrates (ostracoderms) - are key member of the jawed vertebrate (gnathostome) stem, thus fundamental to our understanding of early vertebrate evolutionary patterns. There are many different and disparate clades within the Heterostraci, but all are united in having a single pair of external branchial openings. Phylogenetic relationships of the Heterostraci are poorly understood, meaning the pleisomorphic state for heterostracans and most likely vertebrates as a whole is unknown. Here, we propose a new phylogeny including all described heterostracan genera (137 taxa), including taxa from Pteraspidiformes, Cyathaspididae, Psammosteidae, Amphiaspididae, Traquariaspididae and the problematica. Our results indicate the Cyathaspididae are paraphyletic and the Psammsoteidae fall within the Pteraspidiformes. Inclusion of quantitative characters improves the resolution of phylogenetic trees for this morphologically homogenous group. This novel phylogeny provides a framework to investigate major evolutionary transitions such as dispersal histories. Using taxa's stratigraphical occurrence, we also investigate the stratigraphical congruence of phylogenies and find that heterostracans originate in Laurentia and appear to disperse into other palaeocontinents from this provenance.
Morphological data partitioning and phylogenetic signal in rodents.
 
*
Aime H Rankin
1
 
Robert J Asher
1
 
1
University Museum of Zoology, University of Cambridge
Different morphological character partitions (e.g., teeth vs. postcrania) sometimes vary in the extent to which they reflect phylogenetic history. They may also vary in terms of their impact on deep vs. superficial nodes. For example, in at least some clades, dental data might reflect accurate, historical signal among recently diverged species and genera, but become noisier with deeper nodes and older divergences. Here, we measure how signal provided by different partitions in Glires (Rodentia + Lagomorpha) varies with node depth across a tree, from superficial clades at the tips to basal clades near the root. Glires comprises over 40% of mammalian species and has an excellent fossil record and our understanding of their phylogenetic history has greatly improved in recent years. This makes it a good sample with which to examine hard-tissue partitions. By measuring median distances between unambiguous character changes and the root of a tree across several morphological partitions, we found no significant differences between the depths of clades supported by the different data partitions in Glires. Finer detail suggests a slight bias in axial skeleton characters to support more basal clades and for braincase and basicranium characters to optimize as synapomorphies for clades near the tips of the tree. Overall, our study suggests that commonly used, hard-tissue morphological characters in Glires contain at least some signal to support clades throughout the tree, both deep and superficial.
How do mechanical loads influence cranial shape and function during development?
 
*
Emily J Rayfield
1
 
Lucy Brunt
2
 
Jen A Bright
3
 
Karen Roddy
2
 
Chrissy Hammond
2
 
1
School of Earth Sciences, University of Bristol
 
2
School of Physiology and Pharmacology, University of Bristol
 
3
School of Geosciences, University of South Florida, Tampa, Florida
The shape of the vertebrate skeleton is dictated by a mixture of genetic patterning and environmental influences. Mechanical loading via muscle activity is required for normal skeletal development, including in the skull. Despite this, we have little idea of the extent to which mechanical loads control embryonic and adult morphology and how this subsequently influences skull function. To address this question we used zebrafish, a model vertebrate system, to investigate how disrupted mechanical loading modifies craniofacial form and function. We compared control fish developing normally to fish anaesthetized for varying time periods between 3-5 days postfertilization and mutant fish that lack adductor musculature. Control fish began to voluntarily open and close their mouths at days 4 to 5. Removal of muscle loads at days 4 to 5 resulted in abnormal Meckel’s cartilage and jaw joint shapes. Removal of loads earlier during development had little effect on element shape and joint formation. After recording muscle fibres, fibre area and estimated muscle force, we used finite element modelling to reconstruct strains within the developing cartilages. Mutant and anesthetized jaws functioned differently to control jaws, and when muscle-induced strain is absent, cells on the medial side of the joint changed orientation. Our results suggest that muscle-induced strain regulates cell orientation at the developing joint and that biomechanical loading via adductor muscle contraction plays a key role in normal zebrafish jaw formation. Modification of the developmental loading environment leads to cranial shape changes and offers a mechanism by which evolutionary shape change can occur.
Differences between hard and soft data in phylogenetic analyses
 
*
Robert S Sansom
1
 
Matthew Wills
2
 
1
University of Manchester
 
2
University of Bath
Heterogeneity of phylogenetic signal is often tested for and accounted for when analysing molecular data, but not when analysing morphological data. Differences between types of morphological data will prove problematic, especially if those differences align with the missing data biases intrinsic to the fossil record. Here we demonstrate through meta-analysis of 59 empirical morphological datasets that significant differences exist between the phylogenetic signal conveyed by ‘hard’ biomineralized characters and ‘soft’ characters (i.e. bones, teeth and shells versus myology, integument, neurology etc.). Those differences are demonstrated through application of partition heterogeneity tests (incongruence length difference test and incongruence relationship distance test). Furthermore, hard and soft morphological partitions are found to differ in their consistency relative to independent molecular trees. Differences between readily fossilizable and less fossilizable characters could prove problematic for drawing evolutionary inferences from fossil data, especially in those clades in which the hard character partitions have been demonstrated to be less consistent with molecular trees than the soft character partitions (mammals, bivalves). Other clades are found to exhibit fewer differences (non-acanthopterygian fishes, gastropods) or demonstrate elevated consistency of the hard character partitions on molecular trees relative to soft character partitions (birds, potentially including dinosaurs, and acanthopterygian fishes). In which case, the later clades maybe a more appropriate focus for macroevolutionary studies that utilize fossil data. In all cases it will be necessary to consider the role that missing data plays in shaping empirical data, and consider the differences that exist between morphological modules.
Testing adaptive radiation scenarios in marine fishes by combining phylogenomic and paleobiological data
 
*
Francesco Santini
1
 
1
Associazione Italiana per lo Studio della Biodiversita'
Adaptive radiation scenarios have been invoked to explain the diversity of some of the best studied groups of organisms (e.g., Rift lake cichlids, Hawaiian Silversword Alliance, passerine birds). Under the most traditional adaptive radiation model numerous lineages start diverging within a brief period of time from an ancestral adaptive type, with each new lineage filling an available ecological niche; subsequently this rapid initial morphological evolution is replaced by relative stasis due to most available niches having already been filled. A number of recent studies, based on molecular phylogenies, questioned the generality of this model and found little evidence of an early burst of morphological diversification in most studies. For most of these clades, however, it is not known if inclusion of the paleodiversity would have modified the results. I will compare the results of our study of several major groups of marine teleosts, such as tetraodontiforms (puffers, triggerfishes and allies), carangoids (jacks, remoras), acanthuroids (surgeonfishes, luvar) and sparoids (seabreams, emperors and allies). All of these groups possess a rich fossil record - mostly known from European sites - which to date has rarely been used in evolutionary studies. I will show how the results based on extant taxa and those based on extant plus extinct species differ, and how inclusion of fossil data can alter the conclusion of studies based on molecular phylogenies.
Improving morphological character choice in phylogenetic inference - data from modern crocodilians
 
*
Roland B. Sookias
1
 
1
Museum für Naturkunde, Berlin
Molecular phylogenies are increasingly reliable, with larger datasets and improved analytical techniques. Morphological phylogenies often continue to conflict with them, due to convergent morphological evolution masking phylogenetic signal and the difficulties of character delineation. While molecular analyses can be increasingly relied upon for modern taxa, phylogenetic placement of fossil taxa must continue rely on morphology, and this raises the concern that phylogenetic placement of fossils will remain less accurate. In order to asses how better to avoid these problems, homoplasy of characters from a recent morphological phylogenetic analysis including modern crocodilians was assessed against a tree based on recent molecular phylogenies. The relative homoplasy of different character groups was assessed using the consistency index, retention index and rescaled consisteny index. Cranial characters were significantly less homoplastic than postcranial characters (p=0.05 level), as were multistate characters compared to binary characters. Differences between braincase characters and the rest of the skull, and between “naturally” delineated characters and other characters were nonsignificant. Each morphological character in the matrix was reassessed first hand. Those characters in which homology across taxa within a state was not doubtful tended to be consistent with the molecular tree. This was confounded by less robust characters possibly having been created/scored with a preexisting grouping in mind; these characters were nonhomoplastic but could not be reliably observed. This work may begin to assist in improving fossil archosaur, and potentially vertebrate, phylogenetics; further work with a wider taxonomic scope is required.
The complicated and surprisingly early origin of the pterodactyloid bauplan
 
*
David M Unwin
1
 
1
School of Museum Studies, University of Leicester, 19 University Road, Leicester LE1 7RF
The origin of the pterodactyloid bauplan from that of non-monofenestratan (‘rhamphorhynchoid’) pterosaurs involved extensive anatomical changes and had profound consequences for the evolutionary history of Pterodactyloidea, a clade that dominated the aerial realm throughout the Cretaceous. This important evolutionary transformation, about which almost nothing was known for almost two centuries, is now rapidly coming into focus thanks to a plethora of new pterosaur fossils from the Middle and Upper Jurassic of South America, Europe and China. So far, however, these finds have largely been considered in isolation from each other and few have been thoroughly evaluated. Phylogenetic analysis, combined with improved stratigraphic data for all potentially relevant taxa including putative non-pterodactyloid monofenstratans (NPMs) and the oldest known pterodactyloids (e.g. Liaodactylus) was used to generate a new map of the anatomical transformations and temporal history of the non-monofenestratan–pterodactyloid transition. Evolution of the pterodactyloid skull construction predates the Middle Jurassic, but remains almost completely undocumented by fossils. Liaodactylus reveals that innovation in pterodactyloid skull anatomy and the appearance of derived features was well underway prior to the Upper Jurassic. Douzhanopterus, a derived NPM, demonstrates that elongation of the metacarpus and reduction of the tail and fifth toe (classic pterodactyloid synapomorphies) also predates the Upper Jurassic, but disjunction in the degree of their development across taxa is not consistent with simple explanations such as ‘adaptation for flight’. Overall, late Early to early Late Jurassic pterosaurs were much more diverse and had a far more complex evolutionary history than heretofore recognised.
Range of motion and hydrodynamic implications of the long-neck in plesiosaurs.
 
*
Pernille Venø Troelsen
1
 
Peter L. Falkingham
1
 
1
Liverpool John Moores University
Plesiosaurs are iconic extinct marine reptiles, and they possessed some of the longest necks among all vertebrates. The biomechanical implications of such a long neck on fundamental functions such as forward swimming remain relatively understudied. This study explored range of motion between cervical vertebrae from two plesiosaur species (Mureanosaurus sp. and Cryptoclidus eurymerus) using 3D digitization and modelling. The 3D models were manually aligned and then rotated until contact with the posterior vertebrae in order to measure maximal lateral, dorsal and ventral bending limits. Both species showed greatest bending dorsally (10.7° and 10.2° mean intervertebral rotations for Muraenosaurus and C. eurymerus). Mean ventral rotations were 5.3° and 7.5°, and lateral rotations 7.9° and 7.5° for Muraenosaurus and C. eurymerus. Summed for all vertebrae in the neck, this implies total dorsal/ventral/lateral flexion of 428°/212°/316° for Muraenosaurus and 326°/240°/240° for C. eurymerus. While soft tissue would reduce these ranges considerable, this provides an osteological maximum. Additionally, computational fluid dynamics (CFD) was used to simulate hydrodynamic effects of the long neck during forward locomotion. The CFD analyses included simulations of an idealised 3D plesiosaur model with the neck bent at different degrees (0-90) and locations (head, middle, and evenly). Intuitively, the CFD results showed higher drag as degree of bending was increased (0°; Cd = 0.041 vs. at 90°; Cd = 0.204). Combining digitization of real fossil specimens with idealized simulation can help shed light on the biomechanical implications of the long neck in plesiosaurs.
Glide analysis and bone strength tests indicate powered flight capabilities in hatchling pterosaurs
 
*
Mark P Witton
1
 
Elizabeth Martin-Silverstone
2
 
Darren Naish
2
 
1
School of Earth and Environmental Sciences, University of Portsmouth, Burnaby Building, Burnaby Road, Portsmouth, UK, PO1 3QL
 
2
School of Ocean and Earth Sciences, National Oceanography Centre, University of Southampton, European Way, Southampton, UK, SO14 3ZH
Pterosaur embryos and ‘hatchling’ specimens show a surprising level of skeletal development including well-ossified skeletons and large wings. This has prompted interpretations of pterosaurs as being flight-capable from the earliest ontogenetic stages, contrasting them against the majority of other flying animals, living or extinct. Though popular, this hypothesis is not universally accepted. Some authors propose that pterosaurs only became flight capable once they reached 50% of maximum size, explaining a slowing of growth rate in later ontogeny as metabolic resources were diverted into an energy-demanding form of locomotion. We investigated these hypotheses through glide performance and wing bone strength analysis on hatchling-grade specimens of two pterosaurs, Pterodaustro guinazui and Sinopterus dongi. We found that hatchling pterosaurs were excellent gliders, but with a wing ecomorphology more comparable to powered fliers than obligate gliders. Bone strength analysis shows that hatchling pterosaur wing bones are structurally identical to those of larger pterosaurs and – because of their very low body masses – their bending strength relative to body weight is very high, comparable to or exceeding the greatest values estimated for larger, more mature pterosaurs. Hatchling pterosaurs are thus as mechanically adapted to powered flight stresses as other pterosaurs, if not moreso. Together with our glide tests, this result supports interpretations of hatchling pterosaurs as flight-capable. Size differences between pterosaur hatchlings and larger members of their species dictate differences in wing ecomorphology and flight capabilities at different life stages, which might have bearing on pterosaur ontogenetic niching.

Poster
A revision of the morphology and systematics of geoemydids (Testudines: Cryptodira) from the Messel Pit quarry
 
*
Eduardo Ascarrunz
1
 
Walter G Joyce
1
 
1
University of Fribourg, Switzerland
We present preliminary results of our revision of Eocene European geoemydids, concentrating on two putative species from the Konservat-Lagerstätte of the Messel Pit quarry, in the state of Hesse, Germany. The two species, herein referred to as Palaeoemys kehreri and Palaeoemys messeliana, are the dominant components of the abundant and well-preserved testudinoid material from Messel being represented by nearly 100 specimens, often complete shells with associated skeletal material. Despite this, there are not clear diagnoses for these species or a firm knowledge of their relationships within geoemydids, in good part because of the extensive morphological variation within the populations, and the difficulty of finding phylogenetically informative characters for geoemydids in general. We aim to remedy that situation with new descriptions based on observations of multiple specimens from the two largest collections for these species, which we integrate in the context of more general geoemydid relationships thanks to a larger body of data that we have collected from the extant species of the group.
Systematics of Late Cretaceous European ankylosaurs: the importance of the dermal armour
 
*
Attila Ősi
1,2
 
Jonatán Rudolf
1
 
1
Eötvös University, Department of Paleontology, Budapest, Hungary
 
2
Hungarian Natural History Museum, Budapest, Hungary
Dermal armour composition and/or the morphology of a specific osteoderm can be diagnostic at generic or even species level in many lineages of armoured tetrapods. In ankylosaurs, besides cranial ornamentation, the cervical and pelvic armour elements are the most complex, diagnostic and easily recognizable parts. Whereas cervical elements are usually fused in half-rings bearing plates and/or spikes, pelvic armour shows a diverse composite of oval, polygonal or rounded osteoderms being completely or partially fused or unfused.
The ankylosaur record from the Upper Cretaceous of Europe is relatively poor known only by a few occurrences from Cenomanian and Santonian to Maastrichtian deposits, but osteoderms are present in most assemblages. Two different morphotypes of cervical half-rings, representing Hungarosaurus tormai and Struthiosaurus sp., are present in the Santonian of Hungary. The half-ring of the Early Campanian S. austriacus differs from the Hungarian remains. From the Upper Cretacoeus of the Ibero-Armorican landmass cervical half-rings are known from Cruzy (southern France), Laño and Chera (Spain). Though they are generally similar to each other and to those of the Central European elements, the shape of the dorsally projected plates and the number and morphology of the ossified elements between the plates are different. In addition, the recognition of massive conical spikes both in Hungarosaurus and in the Austrian ankylosaur material suggests that the latter is also represented by two different taxa. This suggests that ankylosaur diversity during the last 20 My of the Late Cretaceous European archipelago was more diverse than previously thought.
Evaluating the Performance of Morphological Partitions in Recovering Avian Evolutionary History
 
*
Leah Callender-Crowe
1
 
Robert S Sansom
1
 
1
The University of Manchester
Despite increased use of genomic data in phylogenetics, morphological information remains important in resolving evolutionary relationships, particularly for rare and fossil species where molecular data is unavailable. Accordingly, methods of phylogenetic reconstruction using morphological data have been widely discussed in recent literature, including whether the use of sophisticated model-based methods is better than traditional parsimony. Another important issue with molecular data, however, is the nature of the morphological characters themselves. Selection and evolutionary lability can make some sets of characters more prone to homoplasy than others. Moreover, when these characters evolve in semi-autonomous modules, they have the potential to overwhelm genuine phylogenetic signal contained within the data. The present study uses a meta-analysis approach to compare osteological and soft-character partitions in birds to determine whether these partitions contain different levels of homoplasy when compared to a molecular phylogeny. I find that soft-tissue characters are less consistent with molecular data in general, but that they are more consistent with this phylogeny for a single, large morphological dataset. This may partly reflect different evolutionary rates at different taxonomic levels, but generally osteological data appear to track evolutionary history better than soft characters in birds.
The Anatomy and Relationships of Eucamerotus foxi (Dinosauria, Sauropoda) from the early Cretaceous of England
 
*
Amy Campbell
1
 
1
University College London
Eucamerotus foxi is a macronarian sauropod from the Wessex Formation (Early Cretaceous, Barremian) of the Isle of Wight. It is represented by NHMUK R2522, a partial dorsal neural arch, and differs from other known sauropods by the presence of a stout robust prezygaparapophyseal lamina (PRPL) which bifurcates distally. NHMUK R90 are a pair of dorsal vertebrae also from the Wessex Formation which have been designated as paratypes of E. foxi, both R2522 and R90 have traditionally been regarded as basal titanosauriformes or of brachiosaurid affinities. Here phylogenetic analysis using the LSDM recovered R90 as a non-titanosaurian somphospondylan nested in a clade made up of ((Tastavinsaurus (Chubutisaurus, Angolatitan, R90)) though Bremer support for this group was low. Analyses recovered E. foxi as a neosauropod of uncertain affinities with little resolution throughout the tree. R90 is recognised as being distinct from E. foxi based on the lack of a robust distally bifurcated PRPL and is likely representative of a new sauropod taxon. E. foxi is here retained as a valid species of basal macronarian.
Tooth damage, loss and replacement in Pycnodont fishes.
 
*
Sally Collins
1
 
Charlie J Underwood
1
 
Martha Richter
3
 
Zerina A Johanson
3
 
Alison Longbottom
3
 
1
Birkbeck, University of London
 
3
Natural History Museum, London
Pycnodonts are an extinct group of neopterygian fishes that lived from the late Triassic (Norian) to Eocene (Lutetian-Ypresian). They were primarily marine and often reef dwelling, with heterodont dentitions, including crushing teeth on the vomer and prearticular bones. Two species, Pycnodus maliensis and Pycnodus zeaformis from the Late Palaeocene/Early Eocene of Mali have highly unusual dentitions, which appear to show tooth replacement where damage or loss of teeth has occurred. Micro-CT scans of the fossils show that small, round teeth, very different to the molariform teeth they have replaced, appear to have developed at an abrasion or wound surface; possibly constituting wound-healing tooth addition. We are comparing tooth replacement mechanisms in modern fish, to infer those present in these species. In the morphologically similar vomer of the common sea-bream (Pagrus pagrus) we found the mechanism to be different, with replacement teeth located beneath and associated with the functional tooth, as normal for actinopterygians. Tooth replacement studies on Atlantic salmon (Salmo salar) have identified a middle dental epithelium in this and two other teleost species (O. mykiss and P. senegalus). Our micro-CT scans of the salmon show unusual positioning of new teeth in spawning males, providing further scope for comparison. Studies on Cichlid pharyngeal jaws show that changes in an individual’s tooth type, for crushing or not, can be epigenetically induced depending on the availability of hard and soft food. Next steps are therefore to search for genetic control mechanisms in these and other vertebrates, that could produce this unusual morphology.
New data on the ophthalmosaurid basicranium
 
*
Lene Liebe Delsett
 
Aubrey Jane Roberts
1
 
Patrick Scott Druckenmiller
2
 
Jørn Harald Hurum
 
1
The National Oceanography Centre, University of Southampton
 
2
University of Alaska Museum
Ophthalmosaurids were the last ichthyosaur group and the morphology of the basicranium is important for phylogeny, especially the amount of basioccipital extracondylar area. However, this character has some challenges in phylogenetics, because its presence is inconsistent on family and subfamily level. Although articulating to the basioccipital, the basisphenoid is rarely found, as few specimens preserve both elements. Material from the Slottsmøya Member Lagerstätte (SML), Spitsbergen, can contribute to an evaluation of basicranium characters. Excavations (2004-2012) in the SML (Tithonian- Berriasian) have yielded 29 ichthyosaur specimens. Seven are previously described with a new specimen presented here, as well as Cryopterygius kristiansenae basicranial material. The size relationship between the basioccipital and basisphenoid for SML material and 16 specimens from other localities was calculated in order to establish whether there is any systematic variation interesting for phylogenetic relationships or functional morphology.

The new specimen preserves basicranium, jaw remains, an articulated dorsal vertebral column and pectoral girdle. It shows similarities to SML genera Janusaurus and Palvennia, but also to the widespread Arthropterygius, with possible biogeographical implications. Material from the Cryopterygius kristiansenae holotype includes basioccipital, basisphenoid and stapes. The subfamily Ophthalmosaurinae is characterized by a large extracondylar area, but the supposedly ophthalmosaurine SML assemblage includes specimens with zero and another with a very extensive extracondylar area. The basisphenoid morphology also varies, most importantly in size and shape of the pterygoid processes. The results from the calculation of basioccipital-basisphenoid pairs indicate that the size relationship is stable within Ophthalmosauridae and considerably less variable than the morphology.
Deltapodus footprints provide evidence of stegosaurs on Isle of Skye (Scotland, UK) in the Middle Jurassic
 
*
Paige dePolo
1
 
Stephen L Brusatte
1
 
Tom Challands
1
 
Davide Foffa
1
 
Mark Wilkinson
1
 
Paulo Pereira
1
 
Dugald Ross
7
 
1
University of Edinburgh
 
7
Staffin Museum
Although the Middle Jurassic (ca. 174-164 Ma) marks a pivotal time of diversification within dinosaur lineages, the global dinosaur body fossil record during this time is sparse. Dinosaur footprints supplement this scrappy body fossil record. Two new in-situ dinosaur track sites from the Lealt Shale Formation of the Great Estuarine Group (Bathonian) on the Isle of Skye present a diverse ichnoassemblage that includes footprints associated with ornithopod, theropod, and thyreophoran track makers. The tracks are predominantly impressions in a dark gray, thinly laminated shale with abundant desiccation cracks that were later infilled with casts of a tan bioclastic limestone. While the theropod and ornithopod tracks are consistent with previous footprint discoveries on Skye, a short quadrupedal trackway consisting of five prints of Deltapodus morphology is of particular interest as the first thyreophoran trackway from this area. Although the overall morphology is consistent with the type Deltapodus tracks from Yorkshire, the Skye Deltapodus tracks are significantly smaller (with an average length 16 cm). The tracks extend the known range of Deltapodus which is primarily known from Western Europe, the western United States, and China and are the most northerly occurrence of this ichnogenus. These tracks provide evidence that stegosaurs were present on Skye during the Middle Jurassic and underscore the Isle of Skye’s potential as location for better understanding this period of geologic history.
Correlates of mammalian tooth eruption patterns
 
*
Madeleine Geiger
1
 
Robert J Asher
1
 
1
University of Cambridge, Department of Zoology
Variation of tooth eruption and replacement patterns among mammalian species has been hypothesised to be related to various factors, including phylogenetic relatedness, diet, crown height, brain size, and pace of growth. For example, Schultz’s rule predicts that replacement teeth (second generation teeth) tend to erupt earlier in respect to the molar teeth in relatively slow growing mammals. Here, we present a study which compares tooth eruption patterns across the whole breadth of Mammalia and relates potential changes of these patterns to a range of variables that have previously been hypothesised to influence tooth eruption. Our dataset comprises more than 1000 specimens, representing ontogenetic series of 40 mammalian species, including marsupials, xenarthrans, afrotheres, euarchontoglires, and laurasiatherians. These data show that the variability of tooth eruption patterns is influenced by phylogenetic relatedness, i.e., certain patterns are only found in certain clades. This includes a correlation between at least some life history patterns and "slow" development (i.e., Schultz's rule) in primates. Further, variation of body mass appears to affect variation of tooth eruption patterns across all species. Our study will help to elucidate the evolution of tooth eruption patterns on a broad taxonomic scale, not only within certain clades, and will therefore likely also inform inferences about extinct taxa.
AN ICHNOLOGICAL STUDY OF THE TERRESTRIAL ‘RED-BEDS’ FROM THE SALOP FORMATION (UPPER CARBONIFEROUS) OF SHROPSHIRE
 
*
Josh Hedge
1
 
Richard J. Butler
1
 
Ivan Sansom
1
 
1
University of Birmingham
This study involved a re-evaluation of exceptionally preserved ichnofossils from the Alveley Member of the Salop Formation, a terrestrial ‘red-bed’ formation from the Upper Carboniferous of Shropshire. It was originally described by Raw (1915), and noted for the contained hypo-relief tetrapod footprints. This study focussed on resting tetrapod impressions (cubichnia) and also epi-relief arthropod trackways (repichnia) that have been less extensively studied. Analysis of the material was undertaken through comprehensive cataloguing; quantitative measurement and qualitative examination of the fossil morphology; 3D computer modelling by photogrammetry, and comparison to global palaeontological analogues. The vertebrate cubichnia was identified as Hermundurichnus Walter and Werneburg, 1988, the second known example after the holotype. This is the resting impression attributed to the Diplocaulidae, largely based on the v-shaped head impression. The genus here was most closely associated with Diploceraspis Romer, 1952, representing the first known example of this genus in Europe. Preservation of this material was polyphase, with cubichnia and repichnia elements, revealing some insight into previously hypothesised life habits for longhorned diplocaulids. Previous work on diplocaulid skull function was incorporated, and applied to the events that produced this ichnology in order to derive a chronological sequence, and a shoreline lacustrine environment was interpreted. The occurrence of these fossils together combined with previously studied elements of the Alveley Member help better understand the terrestrial environments and vertebrate evolution of the Late Carboniferous.
Luke 23:34 – Do we know what we are doing?
 
*
Bernd Herkner
1
 
1
Senckenberg
At the beginning of the twentieth century a couple of far reaching theories arose, which changed the view about nature fundamentally: quantum physics, theory of relativity, and theories concerning the so called linguistic turn in philosophy. Whilst in the field of physics and even chemistry these theories are nowadays well established, in other fields of natural sciences, they had less or almost no impact. The latter is the case in biology as well as in Palaeontology. Some may ask: what has quantum physics, theory of relativity and linguistics to do with biology and palaeontology? The answer is: a lot. All what I have learned in my career as a biologist, working mainly on the field of palaeontology, is that nature generally has to be seen as a unbroken continuum as well as in a constant state of dynamic. Everything is connected to each other somehow in a way. This view matches well with the modern view of theoretical physics. In the field of biology and palaeontology we inevitably disintegrate the continuum, and dynamic processes are cut in to pieces by building categories and by the act of classification. As far as we know, there is no other option to handle scientific research on nature, but we should know what we are doing. At present none of us reflects the consequences of classification and of building categories. But both determine the range of our results. We still work with a Newtonian view of nature where things are existing independent from each other and scientific results are independent from the observer. And we still ignore that our categories and terms are built by ourselves and that they are not just found in nature. So we should be aware what we are doing. And we should not say more as we can say on the basis of our methods.
Evolution of lower jaw shape disparity in pisciform gnathostomes
 
*
Jennifer (JJ) Hill
1
 
Philip C.J. Donoghue
1
 
Emily J Rayfield
1
 
1
University of Bristol. UK.
The vertebrate lower jaw is an effective proxy for inferring feeding ecology. As such, transformations in mandibular shape and structure may have facilitated the emergence of different feeding behaviours in vertebrate evolution, such as the transition from ram, filter and rake feeding to suction and protrusion feeding and biting. Here we present an elliptical Fourier shape analysis of modern and Palaeozoic primitively aquatic jawed vertebrates to characterize and compare disparity of jaw shape in early gnathostomes and their modern counterparts. 83% of shape variation is summarized on the first three axes and all component clades of early gnathostomes exhibit overlapping morphological variation. Non-tetrapodomorph Palaeozoic sarcopterygians are more disparate than their extant counterparts whereas extant chondrichthyans are more disparate than their Palaeozoic counterparts. More generally, extant pisciform gnathostomes are more disparate than their Palaeozoic relatives largely because of the extensive shape variation exhibited by mandibles of extant actinopterygians. Only some ecological niches vacated by Palaeozoic gnathostomes have been convergently refilled by living taxa. Our results are incompatible with the early burst model of clade evolution and contradict the hypothesis that maximum disparity is reached early in gnathostome evolutionary history.
Palaeopathological Trends in Jurassic Marine Reptiles
 
*
Ally Irwin
1
 
1
University of Southampton
Palaeopathology, the study of disease and injury in ancient animals, has contributed significantly to our current understanding of population dynamics and disease in extinct taxa. The principal aim of this study is to investigate whether there are significant differences in pathologies between Jurassic crocodilians, plesiosaurs and ichthyosaurs. This study also aims to describe pathological specimens of particular interest in order to develop our knowledge of bone healing in these animals.

Results showed a large assortment of bite marks in the Kimmeridge Clay Formation (Upper Jurassic) and Oxford Clay Formation (Middle to Upper Jurassic) material, particularly on the appendicular region of plesiosaurs. Key findings also included well-healed fractures, predominantly in ichthyosaur and plesiosaur ribs, which indicated that Jurassic marine reptiles were resilient against trauma. However, complications arising from a non-fatal bite on a plesiosaur humerus also showed a susceptibility towards infection after injury. A defect of vertebral formation and segmentation, causing congenital scoliosis of the spine, was described for the first time in a marine reptile specimen. This may have had severe implications for the animals’ swimming capabilities. Though there is some evidence to suggest differences in pathology type and location in Jurassic marine reptile groups, a strong emphasis can also be placed on the extraordinary ability of these animals to heal bones subsequent to trauma. This may reflect the high growth rates suggested for some of these clades, as well as the many ecological pressures faced from predation and competition in their respective habitats.
Identification of small carnivore fossils using brain endocasts
 
*
Kotaro Ishikawa
1
 
Tomohiko HORI
2
 
1
Okayama Univ. Sci.
 
2
Tokyo Gas Technology Research Institute
One classification trait that distinguishes species is the shape of the brain; more specifically, shapes of the sulci differ in many types of carnivore. Because of these differences, this brain-form classification method may be used to identify fossil species. In this study, brain form was studied using the cranial fossils of small carnivores (unidentified specimens) and extant species from the Mustelidae (Mustela itatsi, Martes melampus, Meles meles) and Mephitidae (Spilogale putorius, Mephitis mephitis) families. Using these materials, the study considered whether the analysis of brain form is an effective method of fossil identification. The materials used herein included silicone brain endocasts. Identification focused on the sulci in the cerebrum’s surface and involved a comparison of the materials. Differences among species were identified in three sulci: the lateral sulcus (la), located in the upper temporal lobe of the cerebral hemisphere; the suprasylvian sulcus (ss), located in the middle temporal lobe of the cerebral hemisphere; and the sylvian sulcus (sy), located in the lower temporal lobe of the cerebral hemisphere. As a result of comparison, the small carnivore fossil morphotypes were more similar to the sulci of the Mephitidae rather than those of the Mustelidae. Based on the identification of molars, the small carnivore fossils may be those of a spotted skunk (S. putorius). Therefore, the target fossils had the same result as the identification by the molars.
Ecometrics and macroevolutionary patterns in Cenozoic mammals: horse teeth and dog elbows
 
*
Christine M. Janis
1
 
1
University of Bristol. UK.
Ecometrics is the analysis of taxon-free traits – how the distribution of ecomorphological traits in a community reflects organisms’ environmental interactions. Mammalian evolutionary history was profoundly affected by late Neogene cooling and aridification. Ecometric studies provide reciprocal illumination between mammals and their environment, illuminating evolutionary response to climatic change.
Hypsodonty (tooth crown height) not only reflects individual dietary adaptations, but ungulate (hoofed mammals) community hypsodonty levels also correlate well with rainfall. Comparison of Neogene communities shows that North America preceded Eurasia in aridification. This may explain why large mammals were more successful in migrating from North America to Eurasia than vice versa, because they were preadapted for the aridification that would later affect Eurasia.
Dogs’ elbows show changes in predatory behavior associated with Neogene aridification and the spread of grasslands. Elbow morphology reflects forearm rotation ability, restricted in fast runners such as pursuit predators. Dogs switched from ambushing or pouncing to pursuing prey in the late Miocene, possibly giving them a behavioural advantage over dog-like hyenas when they emigrated from North America to the Old World around 7 Ma.
Ecometrics of communities of savanna ungulates, North American late Miocene savannas compared with those of present day East Africa, also illuminates mammalian evolution in the context of climatic change. The North American “Clarendonian chronofauna” contained a large diversity of camelids, which show ecomorphological similarities to the extant African arid-adapted eland, providing further evidence for more arid conditions in the North American Miocene than in the Old World, even in comparison to today.
The Applicability of Aerodynamic Power Curves to Fossil Avians
 
*
Carolina Karoullas
1
 
Robert Nudds
1
 
1
University of Manchester
Aerodynamic power curves are one of the key methods of assessing avian flight capability as they allow for flight style to be quantified. As such, it would be informative if they were applied to fossil taxa. However, the applicability of these curves is unknown as the regression equations used to estimate the variables used in the power curve equation are subject to error- error that could make aerodynamic power curves completely inappropriate for palaeontological use. Here, the best and worst regression equations for each variable were determined by selecting equations with the widest and narrowest confidence and prediction intervals. Using these, the best-case and worst-case scenario aerodynamic power curves were created for various fossil taxa. Overall, it was found that the estimations derived from the regression equations and their associated confidence intervals were meaningless to fossil birds since the equations were based on modern taxa, which are not necessarily representative of fossil avians. Prediction intervals are better predictors of where the aerodynamic power curves of fossil taxa could possibly lay as they are designed to cover all possible bases. However, the prediction intervals showed a huge range of possibilities to the point where it was impossible to tell if the fossil taxon in question could fly at all. Consequently, it was shown that the errors associated with the estimations derived from regression equations created using modern taxa are too large, which leads to the aerodynamic power curves not being accurate enough to be representative of the capabilities of fossil taxa.
Recording the uncollectable with low cost low tech: successful photogrammetry in the field using a mobile phone to create digital 3D models.
 
*
Nigel R Larkin
1
 
Steven Dey
 
1
Cambridge University Museum of Zoology, Downing St, Cambridge, CB2 3EJ, UK.
There are many different ways to record the three-dimensional morphology of a specimen in detail. Most techniques rely on expensive, cumbersome and delicate equipment requiring a power supply. However, if a very large and heavy fossil or geological feature in the field cannot easily be removed to a museum or is in danger of imminent loss it would be very useful to be able record the three-dimensional morphology of the find accurately and in detail there and then with readily available low-cost equipment and with simple techniques.

Fortunately, experimentation has shown that photogrammetry using a standard 5-megapixel mobile phone digital camera can produce very good quality digital 3D models of specimens in the field (such as short dinosaur trackways) that are useful for research. Unlike some scanning methods this also provides a photographic overlay to the morphological model creating, effectively, a 3D photographic record of the specimen and surrounding context.

Even specimens larger than 1m2 can be recorded to a level of sub-millimetre resolution with such a device in reasonable lighting conditions and if the relevant photographic techniques and image processing techniques are well understood. The software required to convert the photographs into 3D models is readily available and low cost. Current mobile phone cameras can even produce results that are better than digital SLR cameras in some low light conditions and they are improving every year. As most people have a mobile phone with them on fieldwork, with a little training anyone can undertake scientifically useful 3D scans.
An unusual ichthyosaur composite: An ‘iffyosaur’ comprising British and German material of different geological stages
 
*
Dean R Lomax
1
 
1
School of Earth, Atmospheric and Environmental Sciences, The University of Manchester, Oxford Road, Manchester, M13 9PL, UK
Composite fossils, if unrecognised, can distort our evaluation of the morphology and variation of taxa. Many specimens of Lower Jurassic ichthyosaurs collected during the 19th century have been identified as composites, but the problem is not restricted to historic specimens. More recently collected material, including some specimens for sale on the fossil market, are also composites, or have been modified. One such specimen (acquired ~1980s) said to be from Lyme Regis, Dorset (Lower Jurassic), and purchased as genuine, is not. The specimen appears to be an almost complete skeleton, but it comprises at least three individuals, along with a carefully reconstructed and carved skull, a coracoid, and a replica forefin. At least three and as many as four sets of vertebrae are pieced together to represent a complete, articulated vertebral column. The only portions that appear genuine in terms of provenance are the hindfins and pelvic girdle, which probably belong together and can be identified as Ichthyosaurus conybeari. The block of matrix containing the hindfins and pelvis and the rest of the skeleton is set into a large block of matrix which is from the Lower Jurassic (Toarcian) Posidonia Shale near Holzmaden, Germany. Therefore, this specimen represents a composite of material from multiple specimens, from two different countries and two different geological stages.
A quantitative evaluation of physical and digital approaches to centre of mass estimation
 
*
Sophie Macaulay
1
 
John R. Hutchinson
2
 
Karl T Bates
1
 
1
Department of Musculoskeletal Biology, Institute of Ageing and Chronic Disease, University of Liverpool
 
2
Structure & Motion Laboratory, Department of Comparative Biomedical Sciences, The Royal Veterinary College
Centre of mass is a fundamental anatomical and biomechanical parameter. Knowledge of centre of mass is essential to inform studies investigating locomotion and other behaviours, through its implications for segment movements, and on whole body factors such as posture. Previous studies have estimated centre of mass position for a range of organisms, using various methodologies. However, few studies assess the accuracy of the methods that they employ, and often provide only brief details on their methodologies. As such, no rigorous, detailed comparisons of accuracy and repeatability within and between methods currently exist. This project therefore applied three methods common in the literature (suspension, scales and digital modelling) to three ‘calibration objects’ in the form of bricks, and to three birds to determine centre of mass position. Application to bricks enabled conclusions to be drawn on the absolute accuracy of each method, in addition to their relative values. Application to birds provided insights into the logistical challenges of applying these methods to more complex, biological specimens. For bricks, we found that, provided appropriate repeats were conducted, the scales methods yielded the most accurate predictions of centre of mass (within 1.49mm), closely followed by digital modelling (within 2.39mm), with results from suspension being the most distant (within 38.5mm). Scales and digital methods both also displayed low variability between estimates, suggesting they can accurately and consistently predict centre of mass position. Our suspension method resulted not only in high margins of error, but also substantial variability, highlighting problems with this method.
New spinosaurid dinosaur finds from the Wessex Formation (Wealden Group, Early Cretaceous) of the Isle of Wight
 
*
Martin Munt
1
 
Steven C Sweetman
2
 
Jeremy Lockwood
1
 
Neil J Gostling
4
 
Alex Peaker
1
 
Gary Blackwell
1
 
Brian Foster
7
 
Josh Clark
4
 
Katy Rankin
9
 
1
Dinosaur Isle Museum, Culver Parade, Sandown, Isle of Wight PO36 8QA
 
2
Palaeobiology Research Group, School of Earth and Environmental Sciences, University of Portsmouth, Burnaby Building, Burnaby
 
4
Biological Sciences 4041/4, Life Sciences Building 85,Highfield Campus, University of Southampton, Southampton, SO17 1BJ
 
7
4 Blueberry Close, Driffield, East Yorkshire, YO25 6SE
 
9
� - IS X-ray Imaging Centre, Faculty of Engineering and the Environment Building 5 (Eustice) Room 1015, Mailpoint M7 Highfield Campus, University of Southampton, Southampton, SO17 1BJ UK
The discovery in Surrey of Baryonyx walkeri in 1983 was a highly significant event which reinvigorated the study of British dinosaurs, at a time when theropod material from the Wealden was usually assigned to the genus Megalosaurus. In the 1990’s the Museum of Isle of Wight Geology (subsequently Dinosaur Isle Museum) acquired many new specimens, including material from the theropod dinosaurs Neovenator salerii and Eotyrannus lengi, however vertebrae, a phalanx and denticle counts of isolated teeth, confirmed the presence of a Baryonyx-like spinosaurid on the Island.
The Wessex Formation exposed on the Isle of Wight is the richest dinosaur assemblage in Europe, with a diverse flora and fauna dominated by dinosaurs. Systematic sieving of the plant debris beds has yielded many baryonichid teeth. However, recently a number of collectors have made significant finds including a substantial proportion of the crania of at least two individuals on the Island’s south-west coast. The area is heavily collected and unfortunately some material has not come into the collection; however many significant finds are now in the care of Dinosaur Isle Museum.
Besides the palaeoenvironment and cranial osteology, future research will include a comprehensive study of the dental morphology, including tooth counts, replacement and variation; CT scanning to establish the endocranial anatomy and examination of the relationship of the trigeminal nerve to the numerous pits on the premaxillae, maxillae and dentaries. Our studies will test the hypothesis that the new material can be assigned to Baryonyx walkeri.
Reviewing the Vertebrate Fossil Record
 
*
Darren Naish
1
 
1
Ocean and Earth Science, University of Southampton
The vastness of the vertebrate fossil record and its literature makes any effort to review it in entirety a difficult task; ‘a review’ is understood to be a work that discusses the evolution and diversity of a group, drawing in knowledge on taxonomy, morphology, ecology and distribution, with representative illustrations. Existing reviews of the entire vertebrate record have mostly been designed with teaching in mind and have focused on groups, trends and processes deemed of interest to students. As goes more specific reviews, some groups (Mesozoic dinosaurs in particular) are frequently reviewed; others are afflicted by their association with idiosyncratic authors, others have only been given partial treatment, and others are woefully under-represented. Biographical information on vertebrate palaeontologists themselves is scattered. An additional issue is that several excellent volumes (notably the Handbook volumes) are prohibitively expensive, rare or otherwise hard to obtain. Cenozoic fishes – in particular actinopterygians, the largest and richest vertebrate radiation – remain essentially untouched and it might not be obvious to non-specialists just how rich the fish record is. Indeed, an enormous number of lineages scarcely known to experts on modern fishes are present in the fossil record.

The result of this skewed coverage is that both popular and technical perception of the vertebrate fossil record is biased. Not only are there a huge number of groups that fail to attract students, a large number of research questions relevant to these groups remain un- or under-investigated because so few researchers are aware of their existence.
Elephant dung, chewed antlers, weathered bones: documenting a unique taphonomic collection.
 
*
Spyridoula Pappa
1
 
Neil Adams
1,3
 
Philippa Brewer
1
 
Simon Parfitt
1,6
 
Gillian Carnegie
7
 
Mark Lewis
1
 
1
Department of Earth Sciences, Natural History Museum, London, SW7 5BD
 
3
School of Earth Sciences, University of Bristol, BS8 1RJ
 
6
Institute of Archaeology, University College London, WC1E 6BT
 
7
Department of Archaeology, University of Durham, DH1 3LE
The Antony John Sutcliffe Collection is a unique taphonomic reference collection held within the Department of Earth Sciences at the Natural History Museum (NHM). Sutcliffe was the Curator of Fossil Mammals at the NHM from 1957 to 1987, during which time he collected a large assemblage of modern comparative vertebrate material, including complete skulls, bones, teeth and soft tissues (skin and faeces) of numerous species from across the world including sites in East Africa, Canada, Alaska, Siberia and northwest Europe. Sutcliffe studied Pleistocene mammals and was particularly interested in the alterations to their remains after death and how they come to be fossilised, i.e., their taphonomy.

The specimens were either collected during Sutcliffe’s own fieldwork or gifted to him by international colleagues, and are often accompanied by detailed notes on their provenance and original letters of donation.

After a preliminary study in 2013, and given the value of the collection for future taphonomic studies, a curatorial project ran from 2015 to 2017 aiming to document the entire collection on EMu (the museum’s collection management system), recording the anatomy, taxonomy, previous and new locations within the museum, and any associated documentation of over 1500 specimens. Photographs of each object were included and all specimens have been registered, safely repacked and rehoused at Wandsworth museum store, where they are now accessible to researchers. The specimens have the potential to form curated taphonomic reference standards for use by researchers around the world.
The most complete ankylosaur skull ever found in the Wessex Sub-basin (Lower Cretaceous) of the Isle of Wight.
 
*
Stuart Pond
1
 
1
 
Ankylosaur remains are frequently recovered from the Lower Cretaceous Wealden deposits of the Isle of Wight, although the vast majority of these fossils represent post-cranial elements and osteoderms. The rarity of ankylosaur cranial material means any new specimens are important for understanding the morphology, palaeoecology and evolution of these taxa. Here we describe a well-preserved partial ankylosaur cranium recovered with associated ankylosaur remains from the Wessex Formation at Compton Bay. This is the most complete ankylosaur skull ever recovered from the Wessex Sub-basin and is now held at Dinosaur Isle Museum (DI), Sandown, IOW. A highly water worn specimen held at the Sedgwick Museum of Earth Sciences was found at Chilton Chine in the early 1990s and assigned to cf. Polacanthus, and the two crania are compared here.
The specimen consists of the posterior part of the basicranium and skull roof including the proximal paraoccipital processes, occipital condyle and basal tuberosity. Both skulls share characters including the position of some of the foramina exiting the endocranium and lateral curvature of the skull roof. The DI skull differs from the Sedgwick specimen by the presence of a well defined, notched border of the supraoccipital and foramen magnum , flat rostal-caudal dorsal cranial surface, the occipital condyle being more bulbous and angled more ventrally, a well defined nuchal shelf and being smaller and less robust. Differences between the specimens may be due to sexual dimorphism, ontogeny or they may represent different taxa.
Applying the fossil record of Crocodylia to assess extinction risk today
 
*
Jonathan Rio
1
 
Philip D Mannion
1
 
1
Imperial College London
The 24 living species of crocodylians (crocodiles, alligators, caimans and gharials) are the sole remnants of a once much more diverse group that evolved 250 million years ago (Ma), with crocodylians first appearing in the fossil record 80 Ma. Approximately 50% of living crocodylians are vulnerable to extinction or critically endangered, reflecting the wider global biodiversity crisis. In response, the field of conservation biology has encompassed palaeontological data, giving rise to conservation palaeobiology. Phylogenetic data can be used to account for bias in studies of extinction selectivity and to test for taxonomic patterns of extinction, particularly whether extinction clusters in certain taxa. Although studies show that extinction risk is not randomly distributed across species, these studies are based on living species only. This leads to a bias in assessments of extinction risk: living groups might seem less threatened because the most susceptible species are already extinct. This project will generate a comprehensive new global phylogeny for Crocodylia that will form the baseline for analysing patterns of extinction in this group. Correlates of extinction susceptibility will be tested, in addition to the taxonomic clustering of extinction. The critically endangered Chinese alligator will be used as a case study. Local ecological information collected from southern Anhui Province, China will be combined with historical, archaeological and palaeontological data to reconstruct spatiotemporal distribution patterns and assess future threats to reintroduced alligators. This work will demonstrate the application of the fossil record and evolutionary history of organisms to modern conservation and assessing extinction risk today.
The Early Triassic bone beds of Spitsbergen – marine ecosystem recovery after the P/T extinction event
 
*
Aubrey Jane Roberts
1,2
 
Lene Liebe Delsett
2
 
Patrick Scott Druckenmiller
4
 
Victoria E. Nash
5
 
Janne Bratvold
2
 
Christine Ekeheien
2
 
Jørn Harald Hurum
2
 
1
Ocean and Earth Science, University of Southampton, United Kingdom
 
2
The Natural History Museum, University of Oslo, Norway
 
4
University of Alaska, Fairbanks, United States
 
5
The Norwegian University of Science and Technology, Trondheim, Norway
The Early Triassic bone beds of Spitsbergen represent some of the earliest records of marine vertebrate fauna recovery after the Permo-Triassic Mass Extinction Event for the Northern embayment of Pangea. Limited systematic and stratigraphic control of Early Triassic vertebrate fossils has been undertaken since their discovery in 1860s and as such additional collection and detailed stratigraphic work was required. The Spitsbergen Mesozoic Research Group has over three field seasons collected new material from two of these levels; the Grippia niveau and the Lower Saurian niveau, from the Early Triassic (Olenekian) Vendomdalen Member, Vikinghøgda Formation at Marmierfjellet, central Spitsbergen. We present the findings from these excavations with preliminary descriptions of this new material, referable to many known taxa. Both bone beds include large- and small bodied ichthyopterygians, as well as a significant amount of chondrichthyan material. From the Lower Saurian niveau, 571 complete to fragmentary vertebrae and over 750 appendicular and cranial fragments from ichthyopterygians were collected. From the Grippia niveau, disarticulated mandibular, rostral and axial skeleton elements from the debated ichthyopterygian genus Omphalosaurus were identified. Additionally, this section includes disarticulated skeletons of the poorly understood, small bodied ichthyopterygian Grippia. Other vertebrate material from these bone beds include, teeth and cephalic spines from fifteen identified chondrichthyan taxa, as well as teleost, coelacanthian and dipnoi skeletal elements. This material is significant for understanding the faunal recovery at the Northern embayment of Pangea and for the palaeobiogeographic studies between the Tethys and Panthalassa.
Bored bones from the terrestrial mid Cretaceous Kem-Kem beds of south east Morocco
 
*
Christina Shears-Ozeki
1
 
1
School of Earth and Environmental Sciences, University of Portsmouth
During the last two decades, bioerosion of bone has become more widely investigated, with the earliest identified coming from mid Triassic strata. Bioerosion has even been identified in bone fragments from a Jurassic theropod coprolite! Although the effects of this destructive process can lead to a loss of important information from the fossil record, and understanding of bone bioerosion may be useful for identifying palaeoclimatic conditions in a variety of ancient fluvial settings.

Recent examination of bones of terrestrial fauna (mostly dinosaurs) from the mid Cretaceous Kem-Kem beds (?Albian - L. Cenomanian) of south east Morocco, identified several distinctive bioerosional traces (surface scratches, bite marks [up to 10 cm long] and borings in the form of channels and oval shaped chambers [~ 5 mm diameter]) from indeterminate producers. One heavily bored bones suggests successive infestation during prolonged subaerial exposure.

Silicon moulding, thin sections and SEM analysis enabled investigation of the bones to determine the producers responsible, and evaluate their role in the taphonomy of ancient fluvial systems.

The bite marks can be attributed to macro predators/scavengers, whereas the borings and surface scratches, are comparable to traces produced by living insects including isopterans and the larvae of coleopterans. Microscopic scratches (approx 1-2 mm long) within the chambers and channels were also identified and considered fodinichnia and domichnia in the form of pupation chambers.
A new Mississippian tetrapod from Fife, Scotland, and its environmental context.
 
*
Timothy R. Smithson
1
 
John Marsden
2
 
Michael A E Browne
3
 
Sarah J Davies
4
 
John E A Marshall
5
 
David Millward
3
 
Stig. A Walsh
7
 
Jennifer A Clack
1
 
1
University Museum of Zoology, Downing Street, Cambridge, CB2 3EJ, UK
 
2
Greenacres Lodge, Fordelhill, Leuchars, Fife, KY16 OBT, UK
 
3
British Geological Survey, The Lyell Centre, Research Avenue South, Edinburgh, EH14 4AP, UK
 
4
Department of Geology, University of Leicester, Leicester, LE1 7RH, UK
 
5
School of Ocean & Earth Science, National Oceanography Centre, University of Southampton, Waterfront Campus, European Way
 
7
Department of Natural Sciences, National Museums Scotland, Chambers Street, Edinburgh, EH1 1JF, UK
The Viséan stage of the Mississippian is a time of rapid tetrapod diversification which marks the earliest appearance of temnospondyls, microsaurs and the limbless aïstopods. Tetrapod finds from this stage are rare and only a dozen sites are known worldwide. Here we announce the discovery of a new Viséan site in Fife, Scotland. A number of tetrapod taxa are present. The most complete is a new species of the baphetoid Spathicephalus, represented by a skull and right mandible. Baphetoids are characterised by large openings on the skull roof, the antorbital vacuities. These are particularly large in the new material. Together the orbits and antorbital vacuities represent approximately 43% of the total skull length. This compares with c28% in the younger species S. mirus from the Serpukhovian of Scotland. The new specimens represent the oldest known baphetoids, yet belong to the most specialised members of the clade. Unlike typical baphetoids, which have the large marginal teeth and palatal fangs present in most early tetrapods, spathicephalids had very broad, flattened heads, with a dentition consisting of a large number of small, uniform teeth. This suggests that spathicephalids may have fed in a similar way to modern cryptobranchid salamanders, and were the earliest tetrapods to use suction feeding for small, aquatic prey. The associated fauna includes actinopterygians, gyracanthids, lungfish and rhizodonts. These are represented mainly by scales, spines and teeth. Palynological and sedimentological analysis indicates that the new fossil bed was deposited in a large, stratified, freshwater lake that became increasingly saline.
Temperature and variable rates of Crocodylomorph body size evolution
 
*
Max Stockdale
1
 
Michael J Benton
1
 
1
University of Bristol. UK.
Environmental temperature is an established constraint on the ecology, and geography of ectotherms. Body size interacts closely with temperature through surface area to volume ratio, with direct consequences for physiology. The crocodylomorpha have shown an enormous diversity of body size, from cat-sized sphenosuchia to the fearsome Sarcosuchus. How does crocodylomorph body size change through time, and how does it relate to climate change? We present an analysis of the interactions between body size, rates of body size evolution and temperature using time-series and comparative phylogenetic approaches. Both body size and rate of body size evolution are found to interact significantly with temperature. Increases in body size and evolutionary rate are associated with periods of cooling, possibly due to constraints imposed by physiology.
A unique Morrison-Formation sauropod specimen with biconcave dorsal vertebrae
 
*
Michael P Taylor
1
 
Mathew J Wedel
2
 
1
University of Bristol. UK.
 
2
Western University of Health Sciences
AMNH FARB 291 is a unique sauropod dinosaur specimen, excavated in 1898 from the Bone Cabin Quarry of Wyoming (Upper Jurassic; Morrison Formation). It consists of a sequence of five contiguous dorsal vertebrae, all with biconvex centra: a morphology not previously observed in any sauropod, and which is very rare even in other amniotes.

The specimen was initially catalogued as Brontosaurus, one of only half a dozen sauropods then known. But it lacks diplodocid features such as bifid neural spines, short centra and dual centroprezygapophyseal laminae. Instead, it resembles Brachiosaurus in its elongated centra, forward-shifted neural arches and lobe-like transverse processes. Centrum lengths (from anteriormost to posteriormost) are 207, 205, 232, 222 and 207 mm, about 60% those of the Brachiosaurus holotype.

This specimen could represent:

1. normal development: the condyle would have ossified separately but the individual died before this could happen;
2. a developmental anomaly;
3. a Brachiosaurus with a rare variation not previously seen due to paucity of specimens;
4. a bizarre taphonomic event;
5. a unique taxon in which this is a normal, inherited trait.

Vertebrae typically ossify as rings of bone around the notochord, which sometimes persists as a tunnel through the centrum. There is a developmental continuum from biconcave centra with a persistent notochordal tube (as in most fishes), through biconcave centra with the notochord persisting as intervertebral cartilage balls, to the normal amniote condition. However, all known baby sauropod presacrals have already ossified opisthocoelous or amphiplatyan centra, so the present specimen remains mysterious.
Growth series of one: case studies in time-transgressive morphology
 
*
Mathew J Wedel
1
 
Michael A Taylor
2
 
Brian Kraatz
1
 
Jann Vendetti
4
 
1
Department of Anatomy, Western University of Health Sciences
 
2
Department of Earth Sciences, University of Bristol
 
4
Malacology Department, Natural History Museum of Los Angeles County
The use of growth series to study ontogeny in the fossil record is complicated by two main factors: growth series are not available for many taxa, and correctly assigning juveniles and adults to the same taxon is often difficult, especially if several related taxa coexisted.

Ontogenetic change can also be revealed in single individuals whose morphology records characters from multiple ontogenetic stages. A snail shell is an intuitive example: the shell grows by accretion at its margin, starting from the protoconch, or larval shell, and moving outward. Larval shell shape varies predictably between planktotrophic and non-planktotrophic lineages, so larval ecology can be inferred from size and morphology of the protoconch, which is embedded in the adult shell (teleoconch).

In many extinct lagomorphs, the occlusal surface of the molars changed markedly over the lifespan of an individual, as features were revealed and then obliterated by wear. In this case, the complete ‘stack’ of potential occlusal morphologies was present in the adult tooth as soon as it was done mineralizing, and further change progressively erased the early character states.

In sauropodomorph dinosaurs, morphological complexity of the vertebrae increases along the cervical series. The simple morphology of anterior cervicals is reflective of both earlier ontogenetic stages, and more primitive character states. More posterior vertebrae reveal the sequential formation of complex structures.

Individuals that record multiple ontogenetic stages can help solve paleobiological problems, such as inferring life histories, determining ranges of variation, and describing how morphological characters come into being.

Lightining Talk (5 mins)
Morphological evolution of the caecilian skull
 
*
Carla Bardua
1
 
Mark Wilkinson
2
 
David J Gower
2
 
Anjali Goswami
1
 
1
University College London, UK
 
2
Natural History Museum, UK
With just over 200 species, Gymnophiona are the smallest amphibian clade. Despite their generally similar appearances (e.g. limblessness), caecilians exhibit a broad range of skull morphologies. However, very few studies explore morphological evolution within this clade. We used geometric morphometrics to quantify cranial morphological variation across Gymnophiona and reconstruct evolutionary mode and rate across the clade. An intensive landmarking approach was used to capture the skull morphology of 34 caecilians, with representatives from every genus. Anatomical landmarks and sliding-semilandmarks were placed onto the skull of each specimen and an automated procedure in the R package “Morpho” was then used to place patch points onto the surface of each bone, positioning points to minimise bending energy of a Thin Plate Spline deformation. These landmark data were subjected to Procrustes analyses to remove non-shape aspects, followed by a series of analyses to reconstruct shape evolution and morphological rates through time in a phylogenetic framework. It was found that a kappa model of evolution best fitted these shape data (κ=0.38), suggesting moderately punctuated evolution. Skull morphology had a strong phylogenetic signal (K=0.80, p=1×10-4), showing that closely related species had similar morphologies. Allometry accounted for 17% of shape variation (Rsq =0.17, p=1×10-4). Atretochoana eiselti exhibited the fastest rate of morphological evolution, and represents the most extreme cranial morphology. One rate shift was found at the base of Scolecomorphidae, revealing fast morphological evolution in this family. This landmark approach allowed accurate representation of skulls across Gymnophiona and new insights into skull morphological evolution.
The skull of the Middle Jurassic plesiosaurian Cryptoclidus revisited
 
*
Mark Evans
1
 
Stephanie Wright
2
 
Roger B J Benson
2
 
1
Department of Geology, University of Leicester, LE1 7RH, UK
 
2
Department of Earth Sciences, University of Oxford, OX1 3AN, UK
The Callovian (Middle Jurassic) cryptoclidid Cryptoclidus eurymerus is one of the best known plesiosaurians. Known from the Oxford Clay Formation of England, it has a character score of 95% in recent phylogenetic datasets.
A new specimen possesses an exceptionally complete, although dorsoventrally crushed, skull. The skull has been CT scanned, along with another previously described and similarly complete, but crushed, skull, allowing segmentation of individual bones. Our observations confirm previous descriptions of the jugal as a much reduced splint of bone. Although the best example is still incomplete ventrally, the material suggests the lack of a tight osteological connection, thus rendering the region structurally weak.
Previous interpretation of the orbital margin regarded it as incompletely known due to abrasion. The new specimen demonstrates that the orbital margin is in fact poorly ossified, with many of the bones displaying a thin, crenelated or fluted edge. The orbital outline deviates from the expected suboval shape, with significant excavations posteromedially and posterolaterally. A robust and concave orbital margin, formed by the ascending process of the maxilla, is only present anteriorly and contributes only 10% to the total. This contrasts with the plesiomorphic condition (e.g. Plesiosaurus) in which the entire margin of the orbit is concave. Data from other cryptoclidids (e.g. 30% concave margin in Muraenosaurus sp.) suggest that a poorly ossified orbital margin is a synapomorphy of the clade and that transmission of stress between the maxillary dentition and dorsal skull roof would have been limited to the region anterior to the orbit.
An anterior sauropod caudal from the Peterborough Oxford Clay: whose tail is it anyway?
 
*
Femke Holwerda
1
 
Jeff J Liston
1
 
1
Bayerische Staatssammlung für Paläontologie und Geologie, Munich, Germany
The Callovian Oxford Clay of England has yielded a rich and diverse marine fauna, mainly discovered and described by Alfred Leeds.
However, occasionally it also brought forth terrestrial fossils, including four isolated cases of sauropod remains, one of a stegosaurid, and another of a dryosaurid. Thus far, only Cetiosaurus oxoniensis and Cetiosauriscus stewarti are confirmed sauropod taxa from these beds.
Here, we describe an isolated sauropod anterior caudal vertebra from the Oxford Clay near Peterborough. Incomplete, it has been immersed in seawater post-mortem, indicated by the adhesion of mollusks, but some characters can nevertheless be used for diagnosis and comparisons.
The anterior articular surface is round, whereas the posterior articular surface is heart-shaped, as in many eusauropods, including Cetiosaurus. Interestingly, the ventral surface shows a keel-like structure; a character shared with neosauropods (e.g. Barosaurus) but also with an as yet unnamed Middle Jurassic sauropod caudal from York, UK.
The posterior side of the neural canal is teardrop-shaped, a characters shared with most basal eusauropods. Below the posterior neural canal, a lip-like structure seems to be present; a character which is shared with Cetiosaurus.
However, the anterior caudal transverse process (ACTP) complex is similar to, although more pronounced than Cetiosauriscus, being more similar to more derived sauropods (Neosauropoda).
Thus, an incomplete isolated element may help elucidate the sauropod species diversity and dispersal in the Middle Jurassic of England.
A unique exceptionally well-preserved pterosaur skull from the Crato Formation of Brazil
 
*
Hebert Bruno Nascimento Campos
1
 
Aude Cincotta
2,3
 
Johan Yans
2
 
Pascal Godefroit
3
 
Eberhard �Dino� Frey
1
 
1
Staatliches Museum für Naturkunde Karlsruhe, Geowissenschaftliche Abteilung, Erbprinzenstr. 13, D-76133 Karlsruhe, Germany.
 
2
University of Namur, Department of Geology, rue de Bruxelles 61, 5000 Namur, Belgium.
 
3
Royal Belgian Institute of Natural Sciences, Directorate ’Earth and History of Life’, rue Vautier 29, 1000 Brussels.
The posterior part of a skull from a tapejarid pterosaur was studied for the exceptional preservation of its soft-tissues. The specimen comes from the Early Cretaceous (Aptian) Crato Formation, in northeastern Brazil. It shows a peculiar soft sagittal crest that gives new information about the anatomy and the diversity of these tapejarid pterosaurs (Pterodactyloidea, Tapejaridae). The specimen lies on five limestone slabs. The bony part comprised the posterior part of the skull, with the braincase and one complex of three dorsally oriented and associated bony crests. The crest consists of a huge and elongated parietal crest, a trabecular frontal crest, and an occipital spine. The specimen preserves the posterior parts of the nasoantorbital fenestra and orbit. The soft cranial crest is characterized by a multilinear dorsal crest with almost parallel fibres, which curve posteriorly. Pigment organelles – melanosomes – were observed in the crest and ‘hair-like’ fibres (pycnofibres). They present diverse shapes and sizes, suggesting diverse colorations for the soft tissues. The new tapejarid shares certain features with the tapejarids Tupandactylus imperator and “Huaxiapterusbenxiensis, which include an elongated parieto-occipital crest. Nonetheless, it differs from Tupandactylus imperator and Ingridia navigans by the following characteristics: absence of a suprapremaxillary spine, very dorsally oriented posterior crests, as well as a specific morphology for the soft crest. Our specimen can therefore be assigned to a new taxon of Tapejaridae, a group previously reported in the Crato Formation.

Plenary Talk (30 mins)
Macroevolutionary patterns in Avialae: birdwatching through geological time
 
*
Daniel J. Field
1
 
1
 
Birds are among the most diverse and ubiquitously distributed tetrapod clades; they inhabit a myriad of different environments, and exhibit incredible disparity in their forms and lifestyles. Unraveling how, when, and why this diversity has arisen demands an appeal to the fossil record of crown birds (Neornithes), as fossils provide us with the only direct evidence of neornithine evolutionary history. Additionally, understanding the origins of the features that make birds unique—such as feather-assisted flight—forces us to look outside of Neornithes to the avian stem lineage, where the nature, timing, and order of character transformations are only accessible through fossils. I will discuss recent macroevolutionary advances related to crown birds and their close relatives among non-avian dinosaurs, covering questions like “how did the ancestors of modern birds survive the end-Cretaceous mass extinction event that eliminated their stem-group relatives?”; “how and why have the biogeographic distributions of modern bird groups changed throughout their evolutionary history?”; and “just how ancient is modern avian powered flying potential?”.
The macroevolutionary consequences of phenotypic integration: from development to deep time
 
*
Anjali Goswami
1
 
1
 
Phenotypic integration is a pervasive characteristic of organisms. Interactions among morphological traits, termed phenotypic integration, can be readily identified through quantitative analysis of geometric morphometric data from living and extinct organisms. These interactions have been hypothesized to be a fundamental influence on morphological evolution on small to large time scales. Simulations using covariance matrices derived from landmark data for diverse vertebrate taxa confirm that trait integration can influence the trajectory and magnitude of response to selection. At a macroevolutionary scale, phenotypic integration also produces both more and less disparate organisms than would be expected under random walk models, thereby increasing occupied morphospace range, but also homoplasy and convergence. However, this effect may not translate simply to evolutionary rates.

The role of integration in shaping morphological evolution is particularly interesting when combined with the observation from many studies that cranial integration changes through ontogeny and that postcranial integration is correlated with reproductive strategy in mammals. If integration directs morphological variation, functional pressures at various points in ontogeny may differ in consequence depending in part on level and pattern of integration. For example, high integration in early postnatal ontogeny in marsupials, combined with strong functional pressures for crawling and suckling, may have contributed to the low variance observed in early postnatal marsupials and low disparity across marsupials, relative to placentals. Here, I discuss the macroevolutionary consequences of interactions among phenotypic integration, ontogeny, and function with comparative data from living and extinct vertebrates.
The future of the past: Integrating fossil and extant anatomy with developmental data to infer vertebrate macroevolution.
 
*
Jason Head
1
 
1
Department of Zoology and University Museum of Zoology, University of Cambridge
Understanding vertebrate macroevolutionary transitions was based primarily on comparative anatomy in a pattern-based school of inference during the 19th-mid 20th centuries. Discoveries of regulatory genetic functions and molecular mechanisms in patterning morphology have dominated vertebrate studies since, incorporating embryological data from a few model taxa in a predominately process-based school of inference. Despite this disjunct, pattern- and process-based approaches both are necessary for a sophisticated understanding of macroevolutionary history, as typified by studies of the evolution of snake-like body forms within squamates.
Evolutionary changes in expression of Hox genes that pattern discrete anatomical regions of the axial skeleton in other vertebrates have been proposed to result in an apparent deregionalization of the axial skeleton in snake-like body forms, including the losses of a distinct cervico-thoracic transition and a ribless lumbar region. These hypotheses are based on embryological comparisons of a single model snake taxon to Mus and a small sample of lizard taxa, and do not examine the distribution of Hox-regulated morphologies across Squamata as tests of process. Placing axial morphologies of extant and fossil squamates in a phylogenetic context does not support the hypothesis that loss of a ribless lumbar region occurred in the evolution of snakes, and quantifying vertebral morphology throughout Squamata reveals regional differences associated with Hox expression boundaries that were previously thought to be non-functional. Thus, integration of morphological and developmental data reveals the evolution of both regulatory gene function and phenotypic expression, as well as the necessity of fossils for understanding both.
Anagenesis: the neglected half of macroevolution
 
*
Tom S Kemp
1
 
1
Oxford University
G. G. Simpson classically distinguished cladogenesis referring to taxon branching, from anagenesis relating to transformation of a single lineage. Modern systematic procedure of only recognising clades based on branch points reconstructed by apomorphy has resulted in almost all current work on macroevolution focussing on cladogenesis – the diversification of taxa over time and space. There are, however, equally valid questions about anagenesis at the macroevolutionary level.

How is the functional integration of the organism maintained even when changes in many characters occur? The correlated progression model of small changes in functionally interconnected traits offers the best account.

What controls the direction of travel of a long lineage through morphospace? Tracking long-lasting, multi-dimensional gradients in a metaphorical adaptive landscape models the process, but do they exist?

Why have so few lineages evolved beyond a low taxonomic level to achieve the conventional status of a new order, class or phylum? Because there are only a few sufficiently complex, long lasting ecological gradients available, such as water to land, ectothermic to endothermic niches, etc.

How does major anagenetic transformation relate to cladogenetic branching? In terms of the adaptive landscape metaphor, as periodic plateaus along a flat-topped ridge.

Why is there so little current interest in this particular area of evolution? Is it due to the constraints imposed by cladistic methodology excluding anagenesis from systematics; to too little evidence to support hypotheses (perjoratively called “scenarios”); to the decline of anatomical, functional morphological, and physiological teaching in favour of molecular and quantitative biology?
Journeying through discrete character morphospaces: synthesising tempo, disparity, and phylogeny
 
*
Graeme T Lloyd
1
 
1
School of Earth and Environment, Maths/Earth and Environment Building, The University of Leeds, Leeds. LS2 9JT
The morphological complexity of the vertebrate skeleton offers a rich data set that can offset deficiencies in their fossil record. One form of encoding such data is the discrete character-taxon matrix that first came to prominence in the middle of the twentieth century and has been utilised to understand evolutionary tempo, morphological diversity, and phylogenetic inference. The synthesis of these three approaches can best be understood in the context of an ordination space where empirical observations are connected by the branches of a phylogenetic hypothesis (previously scaled to geologic time) and separated by morphologic distances. Constructing such spaces is complicated by, amongst other things, the non-Euclidean nature of morphologic distances, the prevalence of phylogenetic uncertainty, and difficulties associated with visualizing hyperdimensional spaces. In addition, the literature contains at least three different approaches for reconstructing hypothetical ancestors (internal nodes) with no clear comparison or preference established. Here I outline current approaches for generating discrete character “chronophylomorphospaces”, apply some new metrics to understand how choices related to ancestral reconstructions affect our understanding of evolutionary tempo and morphological diversity, and identify some outstanding problems and their potential solutions. I show that some ancestral reconstruction approaches can lead to ordination spaces based more on phylogenetic rather than morphologic distances and that the choice to reconstruct ancestors pre- or post-ordination can dramatically alter our perception of evolution. Finally, I show that, complementary to a range of other studies, an early burst of morphological evolution is a common feature of vertebrate data sets.
The Pliocene marine megafauna extinction and its impact on functional diversity
 
*
Catalina Pimiento
1
 
John Griffin
2
 
Christopher Clements
3
 
Daniele Silvestro
4
 
Sara Varela
1
 
Mark Uhen
6
 
Carlos Jaramillo
7
 
1
Nturkunde Museum, Berlin
 
2
Swansea University
 
3
University of Zurich
 
4
University of Gothenburg
 
6
George Mason University
 
7
Smithsonian Tropical Research Institute
The end of the Pliocene marked the beginning of a period of great climatic variability and sea-level oscillations. Here, based on a new analysis of the fossil record, we identify a previously unrecognized extinction event among marine megafauna (mammals, seabirds, turtles and sharks) during this time, with extinction rates three times higher than in the rest of the Cenozoic, and with 36% of Pliocene genera failing to survive into the Pleistocene. To gauge the potential consequences of this event for ecosystem functioning, we evaluate its impacts on functional diversity, focusing on the 86% of the megafauna genera that are associated with coastal habitats. Seven (14%) coastal functional entities (unique trait combinations) disappeared, along with 17% of functional richness (volume of the functional space). The origination of new genera during the Pleistocene created new functional entities and contributed to a functional shift of 21%, but minimally compensated for the functional space lost. Reconstructions show that from the late Pliocene onwards, the global area of the neritic zone significantly diminished and exhibited amplified fluctuations. We hypothesize that the abrupt loss of productive coastal habitats, potentially acting alongside oceanographic alterations, was a key extinction driver. The importance of area loss is supported by model analyses showing that animals with high energy requirements (homeotherms) were more susceptible to extinction. The extinction event we uncover here demonstrates that marine megafauna were more vulnerable to global environmental changes in the recent geological past than previously thought.
Mass Extinction, Recovery, and the Assembly of Biodiversity in Fishes
 
*
Lauren Sallan
1
 
1
University of Pennsylvania
The term “Mass Extinction” brings to mind geologically-instantaneous devastation of the biosphere and global, thorough collapse of ecosystems, in which short-term survival determines the ultimate outcome. This view has been supported by previous work on the famous “Big Five” events, focused on marine invertebrates and on terrestrial tetrapods. Until recently, fishes were largely excluded from such studies due to a lack of appropriate databases, despite their importance in aquatic ecosystems, general diversity (>50% of vertebrate species) and extensive fossil record. Newly compiled databases for fossil fishes are upending our understanding of the role of global events in shaping vertebrate macroevolution. These have highlighted the selective nature of mass extinction, challenging the severity, scope and presumed causes of “Big Five” events. Indeed, second-order crises (e.g. end-Devonian, end-Jurassic) were likely more critical in shaping the evolution and current biodiversity of aquatic vertebrates. Investigations of fish faunas and traits across extinction boundaries have shown that predictable ecological processes during prolonged recovery intervals (15-25 My) are just as, if not more, important as the triggering events in structuring subsequent biodiversity. Mere survival is not enough; long-term responses determine the ultimate outcome of global events. Quantitative investigation of the fish fossil record has, and will continue to, rewrite our understanding of both mass extinction and vertebrate macroevolution as a whole.

SPPC/GCG Conference
Platform presentation (15 mins)
The Remedial Conservation and Support Jacketing of the Neotype dinosaur Massospondylus carinatus.
 
*
Mark R Graham
1
 
1
 
In March 2017 the Neotype specimen of the Early Jurassic South African prosauropod dinosaur Massospondylus carinatus was appraised and condition reported at the Evolutionary Studies Institute, University of the Witwatersrand (WITS), Johannesburg, in readiness for remedial conservation and re-storage.

The work was necessitated by deterioration of the specimen, which was caused by handling over a number of years and an inadequate and failing support mount.

Formally numbered BP/1/4934, but more affectionately known to staff as ‘Big Momma’, the specimen was contained within several individual blocks on flimsy support bases and presented various conservation challenges. These included treatment of fractures and cracking across several surfaces of the fossil and the production of clam shell supports to allow for articulated display within the constraints of an existing display cabinet. Part of the brief was to facilitate safer handling and access for researchers.

This project was led by the author (MRG) who also trained the curatorial and preparation staff at WITS in the methods and techniques employed.

The visit was funded by the Palaeontological Scientific Trust (PAST), the DST/NRF Centre of Excellence in Palaeosciences and The University of the Witwatersrand (WITS).
Finding and collecting a dinosaur in an open pit mine – the Fort McMurray nodosaur
 
*
Donald M Henderson
1
 
1
Royal Tyrrell Museum of Palaeontology
In 2011 a shovel operator working in the Suncor Millenium Mine in northern Alberta bumped into an exceptionally preserved armoured dinosaur contained in a single, very large concretion. The force of impact of the bucket knocked many pieces off the concretion, resulting in a period of 2 days of searching by hand through rubble for small pieces before collection of the remaining fossil could begin. The position of the remainder, located 8m up in a vertical cliff, required the use of a variety of heavy equipment to expose the specimen before it could be collected. After partly exposing the concretion, a high-pressure water and vacuum system was used to remove the last of the softer matrix, and to undercut the concretion. The rapid development of the concretion around the carcass soon after the animal’s death, prevented the fossil from becoming permineralized. This unexpected lack of mechanical strength of the specimen, due to the very soft nature of the skeleton and armour, led to the collapse of the concretion into multiple pieces when it was lifted. The remaining pieces were then collected by stabilizing exposed surfaces with a penetrant stabilizer, and then using standard burlap and plaster jackets. Some of the larger, fractured pieces were held together with various webbing and ratchet-straps prior to jacketing. An 800km return trip by road to the Museum required novel packing and stabilizing methods to avoid abrasion and vibration problems.
Replicating the 1.8m long skull of Pliosaurus carpenteri for display.
 
*
Nigel R Larkin
1
 
1
Cambridge University Museum of Zoology, Downing St, Cambridge, CB2 3EJ, UK.
The type specimen of Pliosaurus carpenteri from Westbury in Wiltshire is the most complete skeleton of its genus known, with an estimated body length of 8m. The whole skeleton was mounted for display at Bristol City Museum & Art Gallery in 2017 for the first time since its excavation in 1994. However, the skull is 1.8 m long, very heavy and consists of sixteen fragile pieces. Mounting the real skull in position would have required a large amount of unsightly supporting metalwork that would have obscured some very interesting pathology on the pallete.
One option was to CT-scan the skull pieces and mount 3D prints of the subsequent digital models. This would present less risk to the specimen than traditional moulding and casting and could possibly be quicker and cheaper. Importantly, the process would also provide 3D morphological data of the skull’s internal anatomy for research for the first time. But would the resulting replica look real or horribly fake?
After CT scanning the skull pieces, replicas were 3D-printed in gypsum and acrylic using a powder based 3D printer. After mounting these pieces together with internal metalwork, the replica was coloured with artists’ acrylic paints to match the fossil. Casts of the teeth were adhered in position, which would have been problematic if the real skull had been used.
Satisfyingly, at the opening of the exhibition many well-known palaeontologists viewing the specimen failed to recognise that the skull they were looking at was in fact a replica. It passed.
Project Airless: addressing the problem of pyrite oxidation in a large fossil collection
 
*
Kieran Miles
1
 
1
The Natural History Museum
The Natural History Museum in London holds around 7 million fossils, a diverse collection of huge scientific and historical importance. The Conservation Centre is responsible for a wide range of specimen care issues, including those affecting the palaeontology collections. One of the most serious of these problems is pyrite oxidation. Pyrite, a form of iron sulphide, can often be found in fossils or their surrounding matrix. Oxidation occurs when unstable pyrite, often in its microcrystalline form, reacts with atmospheric oxygen and water. This reaction is accelerated at relative humidity above 60%, and produces a variety of harmful by-products, usually comprising ferrous sulphate, hydrogen sulphide and sulphuric acid.

Airless, a three year project that started in August 2015, aims to address this problem. The goal is to identify, treat and prevent pyrite oxidation in the Earth Science collections. A small team of conservation technicians are surveying the collections looking for signs of pyrite decay. Affected specimens are taken to a dedicated lab space where remedial treatments, such as dry brushing or ammonia vapour treatment, are carried out, before re-storage of specimens in anoxic microenvironments. These are individually hand-made using barrier film, with oxygen scavenging sachets added to remove oxygen from the sealed bag. In addition, the project has a digitisation aspect, with the use of web-based applications, improving the museum’s database with high-quality photographs that partly compensate for the reduced physical access to the specimens. To date, the team has completed work on nearly 3000 specimens, including ichthyosaurs, plesiosaurs and pterosaurs.
Blood, sweat, and vinegar: Acetic acid preparation of cetacean fossils yields exceptional results
 
*
Tim Ziegler
1
 
Erich MG Fitzgerald
2,1
 
Matthew R McCurry
2,1,6
 
James L Goedert
7
 
1
Geosciences, Museum Victoria, Melbourne, Australia
 
2
National Museum of Natural History, Smithsonian Institution, Washington, DC, USA
 
6
Monash Biomedicine Discovery Institute and Centre for Human Anatomy Education, Department of Anatomy and Developmental Biology, Monash University, Melbourne, Australia
 
7
Burke Museum of Natural History and Culture, University of Washington, Seattle, Washington, USA
Apprehensions about long preparation time, cost of consumables and even personal safety often cause researchers and institutions to eschew the use of acid to prepare fossils in favour of a mechanical approach. Indeed, inconsistently applied chemical preparation protocols can rapidly permit irreversible fossil corrosion. This is seen as particularly pertinent in the preparation of cetacean fossils, which are vulnerable due to their large size, thin neocortex and abundant trabecular bone. We show these factors are mitigable by a methodology employing briefer acid immersions, longer neutralisation times and systematic consolidation and protection of exposed bone. Museums Victoria has accelerated a technical programme preparing Cenozoic marine tetrapods including cetaceans, pinnipeds and birds, following prior institutional advances in the preparation of three-dimensionally preserved Devonian placoderms and Cretaceous mammal microfossils. As an example, we present the cranium and tympanoperiotics of an allodelphinid odontocete, cf. Arktocara (NMV P252767), collected in a calcareous concretion by James L. Goedert from the Upper Oligocene Lincoln Creek Formation (Grays Harbor County, Washington, USA). Cetacean fossils from the North Pacific are notoriously difficult to prepare, due to bioerosion by siboglinid Osedax worms and subsequent tectonisation of sediments along the Pacific active margin. This specimen was prepared primarily with acetic acid over 13 months, with exceptional anatomical detail retained to a degree not achievable mechanically or by less-stringent acid protocols. This is shown by comparisons with examples of previously prepared North Pacific cetacean fossils, and of acid corrosion on an Australian Cretaceous ichthyosaur, Miocene marsupial megafauna, and an in-preparation Miocene odontocete.