Stig A. Walsh

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.

Fossil Seals from Late Neogene Deposits in South America: A New Pinniped (Carnivora, Mammalia) Assemblage from Chile

A new monachine phocid pinniped assemblage from the north–central coast of Chile is described. The material was recovered from a marine bonebed of the Bahia Inglesa Formation which, based on macro– and microfossil evidence, is probably Late Miocene–Early Pliocene in age. At least two genera, Acrophoca and Piscophoca (both originally described from the Pisco Formation of Peru), are present. The Chilean material is significantly different from that of the two described species from the Pisco Formation and probably represents new species, though these are not named pending description of new material reported from Peru. The postcrania are morphologically intermediate between the northern Phocini and southern Lobodontini. The Bahia Inglesa Formation pinniped assemblage represents the first occurrence of fossil seals in Chile, and provides valuable information regarding the late Neogene radiation of monachines in the Southern Hemisphere. The occurrence of Acrophoca sp. in beds underlying a condensed Mio–Pliocene bonebed suggests that these strata are no older than Late Miocene.

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.

New penguin remains from the Pliocene of Northern Chile

Until recently, all described fossil penguin species from South America were recovered from the Atlantic coast. The description of three fossil species of Spheniscus from Peru and Chile now allows a clearer estimate of the historical diversity on the Pacific coast. Here we describe a further new species from a Pliocene level of the Bahía Inglesa Formation, northern Chile. This taxon, the first to be described from this sequence, is based on a partial skeleton lacking a skull. These remains are clearly referable to the living genus Pygoscelis, and share a mosaic of characters with extant species of the genus. Pygoscelis grandis sp. nov. was around the size of a King Penguin, and therefore much larger than any extant Pygoscelis species. Our cladistic analysis places P. grandis within the Pygoscelis clade as the sister taxon of living representatives. Living species of Pygoscelis have a sub-Antarctic distribution, and the presence of this species slightly south of the Tropics during the Pliocene may relate to end Neogene global cooling.

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.

Open data and digital morphology

Over the past two decades, the development of methods for visualizing and analysing specimens digitally, in three and even four dimensions, has transformed the study of living and fossil organisms. However, the initial promise that the widespread application of such methods would facilitate access to the underlying digital data has not been fully achieved. The underlying datasets for many published studies are not readily or freely available, introducing a barrier to verification and reproducibility, and the reuse of data. There is no current agreement or policy on the amount and type of data that should be made available alongside studies that use, and in some cases are wholly reliant on, digital morphology. Here, we propose a set of recommendations for minimum standards and additional best practice for three-dimensional digital data publication, and review the issues around data storage, management and accessibility.

Phylogenetic and environmental context of a Tournaisian tetrapod fauna

The end-Devonian to mid-Mississippian time interval has long been known for its depauperate palaeontological record, especially for tetrapods. This interval encapsulates the time of increasing terrestriality among tetrapods, but only two Tournaisian localities previously produced tetrapod fossils. Here we describe five new Tournaisian tetrapods (Perittodus apsconditus, Koilops herma, Ossirarus kierani, Diploradus austiumensis and Aytonerpeton microps) from two localities in their environmental context. A phylogenetic analysis retrieved three taxa as stem tetrapods, interspersed among Devonian and Carboniferous forms, and two as stem amphibians, suggesting a deep split among crown tetrapods. We also illustrate new tetrapod specimens from these and additional localities in the Scottish Borders region. The new taxa and specimens suggest that tetrapod diversification was well established by the Tournaisian. Sedimentary evidence indicates that the tetrapod fossils are usually associated with sandy siltstones overlying wetland palaeosols. Tetrapods were probably living on vegetated surfaces that were subsequently flooded. We show that atmospheric oxygen levels were stable across the Devonian/Carboniferous boundary, and did not inhibit the evolution of terrestriality. This wealth of tetrapods from Tournaisian localities highlights the potential for discoveries elsewhere.

Best practices for digitally constructing endocranial casts: examples from birds and their dinosaurian relatives

The rapidly expanding interest in, and availability of, digital tomography data to visualize casts of the vertebrate endocranial cavity housing the brain (endocasts) presents new opportunities and challenges to the field of comparative neuroanatomy. The opportunities are many, ranging from the relatively rapid acquisition of data to the unprecedented ability to integrate critically important fossil taxa. The challenges consist of navigating the logistical barriers that often separate a researcher from high‐quality data and minimizing the amount of non‐biological variation expressed in endocasts – variation that may confound meaningful and synthetic results. Our purpose here is to outline preferred approaches for acquiring digital tomographic data, converting those data to an endocast, and making those endocasts as meaningful as possible when considered in a comparative context. This review is intended to benefit those just getting started in the field but also serves to initiate further discussion between active endocast researchers regarding the best practices for advancing the discipline. Congruent with the theme of this volume, we draw our examples from birds and the highly encephalized non‐avian dinosaurs that comprise closely related outgroups along their phylogenetic stem lineage.

Modern Imaging Techniques as a Window to Prehistoric Auditory Worlds

Direct evidence regarding the evolution of vertebrate hearing is available only through the examination of fossils. The fossilized bony anatomy of the middle ear and otic capsule can provide information about auditory adaptations present in extinct animals, which can in turn be used to infer hearing capability in these taxa and to trace the evolutionary history of hearing. However, this internal anatomy is rarely visible in fossils. Historically, fortuitously broken or incomplete specimens, or destructive serial sectioning techniques, have been the only ways in which ear structures could be reconstructed. Modern nondestructive X-ray microcomputed-tomographic (μCT) approaches avoid this problem and allow ear morphology to be assessed in many more taxa, as virtually any three-dimensionally preserved skull can be analyzed reproducibly. This chapter reviews advances in our understanding of vertebrate hearing evolution that have occurred since the application of this imaging technology. In addition to insights gained from the analysis of key fossils, μCT also allows inferences to be made regarding the auditory and vestibular systems of extinct taxa through quantitative analyses of the ear regions of living species. Problems affecting μCT analyses of fossils are discussed and the authors comment on the outlook for future hearing-related μCT research.

Floccular fossa size is not a reliable proxy of ecology and behaviour in vertebrates

The cerebellar floccular and parafloccular lobes are housed in fossae of the periotic region of the skull of different vertebrates. Experimental evidence indicates that the lobes integrate visual and vestibular information and control the vestibulo-ocular reflex, vestibulo-collic reflex, smooth pursuit and gaze holding. Multiple paleoneuroanatomy studies have deduced the behaviour of fossil vertebrates by measuring the floccular fossae (FF). These studies assumed that there are correlations between FF volume and behaviour. However, these assumptions have not been fully tested. Here, we used micro-CT scans of extant mammals (47 species) and birds (59 species) to test six possible morphological-functional associations between FF volume and ecological/behavioural traits of extant animals. Behaviour and ecology do not explain FF volume variability in four out of six variables tested. Two variables with significant results require further empirical testing. Cerebellum plasticity may explain the lack of statistical evidence for the hypotheses tested. Therefore, variation in FF volume seems to be better explained by a combination of factors such as anatomical and phylogenetic evolutionary constraints, and further empirical testing is required.