Angela C. Milner

The Cretaceous-Tertiary biotic transition

Mass extinctions are recognized through the study of fossil groups across event horizons, and from analyses of long-term trends in taxonomic richness and diversity. Both approaches have inherent flaws, and data that once seemed reliable can be readily superseded by the discovery of new fossils and/or the application of new analytical techniques. Herein the current state of the Cretaceous-Tertiary (K-T) biostratigraphical record is reviewed for most major fossil clades, including: calcareous nannoplankton, dinoflagellates, diatoms, radiolaria, foraminifera, ostracodes, scleractinian corals, bryozoans, brachio-pods, molluscs, echinoderms, fish, amphibians, reptiles and terrestrial plants (macrofossils and palynomorphs). These reviews take account of possible biasing factors in the fossil record in order to extract the most comprehensive picture of the K-T biotic crisis available. Results suggest that many faunal and floral groups (ostracodes, bryozoa, ammonite cephalopods, bivalves, archosaurs) were in decline throughout the latest Maastrichtian while others (diatoms, radiolaria, benthic foraminifera, brachiopods, gastropods, fish, amphibians, lepidosaurs, terrestrial plants) passed through the K-T event horizon with only minor taxonomic richness and/or diversity changes. A few microfossil groups (calcareous nannoplankton, dinoflagellates, planktonic foraminifera) did experience a turnover of varying magnitudes in the latest Maastrichtian-earliest Danian. However, many of these turnovers, along with changes in ecological dominance patterns among benthic foraminifera, began in the latest Maastrichtian. Improved taxonomic estimates of the overall pattern and magnitude of the K-T extinction event must await the development of more reliable systematic and phylogenetic data for all Upper Cretaceous clades.

The avian nature of the brain and inner ear of Archaeopteryx

Archaeopteryx, the earliest known flying bird (avialan) from the Late Jurassic period, exhibits many shared primitive characters with more basal coelurosaurian dinosaurs (the clade including all theropods more bird-like than Allosaurus), such as teeth, a long bony tail and pinnate feathers. However, Archaeopteryx possessed asymmetrical flight feathers on its wings and tail, together with a wing feather arrangement shared with modern birds. This suggests some degree of powered flight capability but, until now, little was understood about the extent to which its brain and special senses were adapted for flight. We investigated this problem by computed tomography scanning and three-dimensional reconstruction of the braincase of the London specimen of Archaeopteryx. Here we show the reconstruction of the braincase from which we derived endocasts of the brain and inner ear. These suggest that Archaeopteryx closely resembled modern birds in the dominance of the sense of vision and in the possession of expanded auditory and spatial sensory perception in the ear. We conclude that Archaeopteryx had acquired the derived neurological and structural adaptations necessary for flight. An enlarged forebrain suggests that it had also developed enhanced somatosensory integration with these special senses demanded by a lifestyle involving flying ability.

Baryonyx, a remarkable new theropod dinosaur

An extremely large claw bone, some 30 cm long, was found in Wealden (Lower Cretaceous) deposits in a Surrey claypit in January 1983. This led to the discovery the following month of the well-preserved skeleton of a new large theropod dinosaur. Only one other theropod specimen comprising more than a few bones had ever been found in Britain, and that discovery was more than a century ago. Indeed, no large theropod, reasonably complete, had previously been discovered in Lower Cretaceous rocks anywhere in the world. Our study so far suggests that the Surrey dinosaur was a typical large theropod in certain respects, resembling, for example Allosaurus1. In several other respects, however, it differs sufficiently from all known dinosaurs to merit designation as the representative of a new species, genus and family.

Cretaceous Extinctions: Multiple Causes

![Figure][1] Deccan plateau basalts. Lava from Deccan volcanism formed distinct layering. CREDIT: GSFC/NASA In the Review “The Chicxulub Asteroid Impact and Mass Extinction at the Cretaceous-Paleogene boundary” (P. Schulte et al. , 5 March, p. [1214][2]), the terminal Cretaceous

Inner ear anatomy is a proxy for deducing auditory capability and behaviour in reptiles and birds

Inferences of hearing capabilities and audition-related behaviours in extinct reptiles and birds have previously been based on comparing cochlear duct dimensions with those of living species. However, the relationship between inner-ear bony anatomy and hearing ability or vocalization has never been tested rigorously in extant or fossil taxa. Here, micro-computed tomographic analysis is used to investigate whether simple endosseous cochlear duct (ECD) measurements can be fitted to models of hearing sensitivity, vocalization, sociality and environmental preference in 59 extant reptile and bird species, selected based on their vocalization ability. Length, rostrocaudal/mediolateral width and volume measurements were taken from ECD virtual endocasts and scaled to basicranial length. Multiple regression of these data with measures of hearing sensitivity, vocal complexity, sociality and environmental preference recovered positive correlations between ECD length and hearing range/mean frequency, vocal complexity, the behavioural traits of pair bonding and living in large aggregations, and a negative correlation between ECD length/rostrocaudal width and aquatic environments. No other dimensions correlated with these variables. Our results suggest that ECD length can be used to predict mean hearing frequency and range in fossil taxa, and that this measure may also predict vocal complexity and large group sociality given comprehensive datasets.

Functional morphology of spinosaur ‘crocodile-mimic’ dinosaurs

Abstract Spinosaurid theropod dinosaurs appear to represent convergent morphological evolution toward a crocodylian-like cranial morphology, previously linked to the possibility that spinosaurs adopted a similar, partially piscivorous, trophic niche. Further conclusions are hindered by a lack of quantitative evidence, and an incomplete understanding of the functional significance of key crocodylian cranial characters. A comparative biomechanical analysis of function in the snout of the spinosaurid theropod Baryonyx walkeri has been performed, comparing B. walkeri with a generalised large theropod dinosaur and two extant crocodylians (Alligator, Gavialis) that represent different endpoints of extant crocodylian morphological diversity. The aims of the analysis were (a) to determine which group is the closest functional analogue to B. walkeri, and (b) investigate the mechanical influence on cranial function of the antorbital fenestra and the secondary palate; morphological characters that appear to be of importance in both crocodyliform and spinosaur functional morphology. Results demonstrate that the closest structural and biomechanical analogue to B. walkeri is the extant gharial, rather than the alligator or conventional theropods. The secondary palate confers strength to the alligator skull in torsion, but provides resistance to bending in gharials and B. walkeri. Loss of the antorbital fenestra strengthens narrow or tubular theropod and gharial snouts, but has limited influence on the broader-snouted alligator morphotypes. Consequently, with their large antorbital fenestrae and lack of secondary palate, most theropod skulls were surprisingly suboptimally constructed to resist feeding-related bite loads. The mechanical impetus for archosaur palatal development and fenestral closure appears more complex than previously thought.

Establishing a framework for archosaur cranial mechanics

Abstract The aim of this analysis was to establish the basic mechanical principles of simple archosaur cranial form. In particular we estimated the influence of two key archosaur innovations, the secondary palate and the antorbital fenestra, on the optimal resistance of biting-induced loads. Although such simplified models cannot substitute for more complex cranial geometries, they can act as a clearly derived benchmark that can serve as a reference point for future studies incorporating more complex geometry. We created finite element (FE) models comprising either a tall, domed (oreinirostral) snout or a broad, flat (platyrostral) archosaur snout. Peak von Mises stress was recorded in models with and without a secondary palate and/or antorbital fenestra after the application of bite loads to the tooth row. We examined bilateral bending and unilateral torsion-inducing bites for a series of bite positions along the jaw, and conducted a sensitivity analysis of material properties. Pairwise comparison between different FE morphotypes revealed that oreinirostral models are stronger than their platyrostral counterparts. Oreinirostral models are also stronger in bending than in torsion, whereas platyrostral models are equally susceptible to either load type. As expected, we found that models with a fenestra always have greatest peak stresses and by inference are “weaker,” significantly so in oreinirostral forms and anterior biting platyrostral forms. Surprisingly, although adding a palate always lowers peak stress, this is rarely by large magnitudes and is not significant in bilateral bending bites. The palate is more important in unilateral torsion-inducing biting. Two basic principles of archosaur cranial construction can be derived from these simple models: (1) forms with a fenestra are suboptimally constructed with respect to biting, and (2) the presence or absence of a palate is significant to cranial integrity in unilaterally biting animals. Extrapolating these results to archosaur cranial evolution, it appears that if mechanical optimization were the only criterion on which skull form is based, then most archosaurs could in theory strengthen their skulls to increase resistance to biting forces. These strengthened morphotypes are generally not observed in the fossil record, however, and therefore archosaurs appear subject to various non-mechanical morphological constraints. Carnivorous theropod dinosaurs, for example, may retain large suboptimal fenestra despite generating large bite forces, owing to an interplay between craniofacial ossification and pneumatization. Furthermore, living crocodylians appear to strengthen their skull with a palate and filled fenestral opening in the most efficient way possible, despite being constrained perhaps by hydrodynamic factors to the weaker platyrostral morphotype. The future challenge is to ascertain whether these simple predictions are maintained when the biomechanics of complex cranial geometries are explored in more detail.

Avian cerebellar floccular fossa size is not a proxy for flying ability in birds

Extinct animal behavior has often been inferred from qualitative assessments of relative brain region size in fossil endocranial casts. For instance, flight capability in pterosaurs and early birds has been inferred from the relative size of the cerebellar flocculus, which in life protrudes from the lateral surface of the cerebellum. A primary role of the flocculus is to integrate sensory information about head rotation and translation to stabilize visual gaze via the vestibulo-occular reflex (VOR). Because gaze stabilization is a critical aspect of flight, some authors have suggested that the flocculus is enlarged in flying species. Whether this can be further extended to a floccular expansion in highly maneuverable flying species or floccular reduction in flightless species is unknown. Here, we used micro computed-tomography to reconstruct “virtual” endocranial casts of 60 extant bird species, to extract the same level of anatomical information offered by fossils. Volumes of the floccular fossa and entire brain cavity were measured and these values correlated with four indices of flying behavior. Although a weak positive relationship was found between floccular fossa size and brachial index, no significant relationship was found between floccular fossa size and any other flight mode classification. These findings could be the result of the bony endocranium inaccurately reflecting the size of the neural flocculus, but might also reflect the importance of the flocculus for all modes of locomotion in birds. We therefore conclude that the relative size of the flocculus of endocranial casts is an unreliable predictor of locomotor behavior in extinct birds, and probably also pterosaurs and non-avian dinosaurs.

Halcyornis toliapicus (aves: Lower Eocene, England) indicates advanced neuromorphology in Mesozoic Neornithes

Our recent X-ray micro computer-tomographic (μCT) investigations of Prophaethon shrubsolei and Odontopteryx toliapica from the Lower Eocene London Clay Formation of England revealed the avian brain to have been essentially modern in form by 55 Ma, but that an important vision-related synapomorphy of living birds, the eminentia sagittalis of the telencephalon, was poorly developed. This evidence suggested that the feature probably appeared close to the end of the Mesozoic. Here we use μCT analysis to describe the endocranium of Halcyornis toliapicus, also from the London Clay Formation. The affinities of Halcyornis have been hotly debated, with the taxon referred to the Charadriiformes (Laridae), Coraciiformes (Alcedinidae, and its own family Halcyornithidae) and most recently that Halcyornithidae may be a possible senior synonym of Pseudasturidae (Pan-Psittaciformes). Unlike Prophaethon and Odontopteryx, the eminentia sagittalis of Halcyornis is strongly developed and comparable to that of living species. Like those London Clay taxa, the eminentia sagittalis occupies a rostral position on the telencephalon. The senses of Halcyornis appear to have been well developed. The length of the cochlear duct of the inner ear indicates a hearing sensitivity within the upper range of living species, and enlarged olfactory lobes suggest a reasonable reliance on sense of smell. The optic nerves were especially well developed which, together with the strong development of the eminentia sagittalis, indicates a high degree of visual specialization in Halcyornis. The advanced development of the eminentia sagittalis further supports a Mesozoic age for the appearance of this structure and associated neural architectural complexity found in extant Aves. The eminentia sagittalis of living Psittaciformes is situated caudally on the telencephalon, making a Pan-Psittaciformes relationship unlikely for Halcyornis.

A new Pliensbachian ichthyosaur from Dorset, England

The first ichthyosaur to be recorded from the Pliensbachian Stage of the English Lower Liassic is described as Leptonectes moorei sp. nov., extending the geological range of Leptonectes to the Pliensbachian. According to criteria for assessing the maturity of ichthyosaurs, it is concluded that L. moorei is an immature individual of a relatively small, slender and short snouted species close to the earlier long-snouted L. tenuirostris (Conybeare) which ranges from the Rhaetian to the Sinemurian. The presence of a short-snouted leptonectid in the Pliensbachian suggests two contrasting patterns of rostral morphology within the clade Leptonectidae in the latest Early Jurassic, rostral reduction within the genus Leptonectes and rostral elongation in ExcalibosaurusEurhinosaurus.