Penguins have a more than 60 million year long evolutionary history. Thus, stem lineage fossil taxa are key to understanding their evolution. Here, we present data on three virtual endocasts from stem penguin skulls collected from the Eocene La Meseta Formation of Seymour Island (Antarctica), along with comparative data from extant penguins and outgroups. These fossils appear to belong to three distinct species, and represent both the oldest (34.2 Ma) and the most basal penguin taxa that have yielded endocast data. Data collected from the fossils provide new support for several important shifts in neuroanatomy and cranial skeletal anatomy along the transition from stem to crown penguins, including (1) caudal expansion of the eminentia sagittalis, (2) an increase in the overlap of the telencephalon onto the cerebellum, (3) reduction of the bulbus olfactorius, and (4) loss of the interaural pathway. The large semicircular canal diameters of the Antarctic fossils as well as the more crownward stem penguin Paraptenodytes antarcticus together suggest that canal size increased in basal penguins relative to outgroup taxa but later decreased near the crown radiation. As in most other wing-propelled diving birds, the endocasts lack evidence of cerebellar folds and possess a relatively large floccular recess. Several aspects of the endocast morphology, including the exposure of the tectum opticum in dorsal view and the rostral displacement of the eminentia sagittalis away from the border of the cerebellum, are seen neither in crown penguins nor in Procellariiformes (the extant sister clade to Sphenisciformes) and so appear to represent unique characters of these stem taxa.

Highlights d Dinosaurs and early birds had similar relative brain sizes d Major shifts in brain-body integration occur in the aftermath of the K-Pg extinction d Rates of brain-body evolution are highest in non-avian dinosaurs, early-diverging birds, parrots, and crows d Corvids, like hominins, evolved larger relative brains and bodies simultaneously

Mourasuchus is a Miocene alligatorid endemic to South America, and is represented by four species. Together with the closely related Purussaurus, it is a peculiar crocodylian taxon of neogene Caimaninae and one of the most bizarre forms among eusuchian crocodiles. The phylogenetic relationships between Mourasuchus species have not been explored, and detailed skull descriptions are scarce. The goal of this study is to provide new data on skull morphology and cranial recesses in Mourasuchus nativus, including a new tomography analysis (3D modeling). We observed that several diagnostic characters of Purussaurus, such as lack of contact between the nasal and lacrimal, separation of the nasal and frontal by the prefrontals, and the posterior dorsal margin of the skull table, are shared with Mourasuchus. M. nativus is characterized by the presence of solid transverse squamosal eminences, large posttemporal fenestrae, and a quadrate laterocaudal bridge separating V 2 -V 3 trigeminal openings. Compared with other crocodylians, the endocast of M. nativus is similar in shape but quite sigmoid in lateral view, the canal of the supraorbital ramus of V 2 is more vertically oriented, the thick tympanic branch canal opens in a large foramen aligned with trigeminal foramen, and the canal of the vagal (X) tympanic ramus is also very wide. Contrary to extant alligatorids, the median pharyngeal recess remains paired throughout its course and only connects its opposite fellow near the external ventral opening. The knowledge of the internal skull anatomy of Mourasuchus contributes to the understanding of the general morphology of alligatorids, Caimaninae, and their variation. Anat Rec,

Psittaciform birds exhibit novelties in jaw bone structure and musculature that are associated with strong bite forces. These features include an ossified arcus suborbitalis and the muscles ethmomandibularis and pseudomasseter. We analyse the jaw musculature of the monk parakeet (Myiopsitta monachus) to enable future studies aimed at understanding craniofacial development, morphology, function and evolution. We estimate bite force based on muscle dissections, physiological cross‐sectional area and skull biomechanical modelling. We also compare our results with available data for other birds and traced the evolutionary origin of the three novel diagnostic traits. Our results indicate that, in Myiopsitta, (i) the arcus suborbitalis is absent and the orbit is ventrally closed by an elongate processus orbitalis and a short ligamentum suborbitale; (ii) the ethmomandibularis muscle is a conspicuous muscle with two bellies, with its origin on the anterior portion of the septum interorbitale and insertion on the medial aspect of the mandible; (iii) the pseudomasseter muscle consists of some fibers arising from the m. adductor mandibulae externus superficialis, covering the lateral surface of the arcus jugalis and attaches by an aponeurotic sheet on the processus orbitalis; (iv) a well‐developed adductor mandibulae complex is present; (v) the bite force estimation relative to body mass is higher than that calculated for other non‐psittaciform species; and (vi) character evolution analysis revealed that the absence of the arcus suborbitalis and the presence of the m. pseudomassseter are the ancestral conditions, and mapping is inconclusive about presence of one or two bellies of the m. ethmomandibularis.

The South American phorusrhacid bird radiation comprised at least 18 species of small to gigantic terrestrial predators for which there are no close modern analogs. Here we perform functional analyses of the skull of the medium-sized (∼40 kg) patagornithine phorusrhacid Andalgalornis steulleti (upper Miocene–lower Pliocene, Andalgalá Formation, Catamarca, Argentina) to assess its mechanical performance in a comparative context. Based on computed tomographic (CT) scanning and morphological analysis, the skull of Andalgalornis steulleti is interpreted as showing features reflecting loss of intracranial immobility. Discrete anatomical attributes permitting such cranial kinesis are widespread phorusrhacids outgroups, but this is the first clear evidence of loss of cranial kinesis in a gruiform bird and may be among the best documented cases among all birds. This apomorphic loss is interpreted as an adaptation for enhanced craniofacial rigidity, particularly with regard to sagittal loading. We apply a Finite Element approach to a three-dimensional (3D) model of the skull. Based on regression analysis we estimate the bite force of Andalgalornis at the bill tip to be 133 N. Relative to results obtained from Finite Element Analysis of one of its closest living relatives (seriema) and a large predatory bird (eagle), the phorusrhacid's skull shows relatively high stress under lateral loadings, but low stress where force is applied dorsoventrally (sagittally) and in “pullback” simulations. Given the relative weakness of the skull mediolaterally, it seems unlikely that Andalgalornis engaged in potentially risky behaviors that involved subduing large, struggling prey with its beak. We suggest that it either consumed smaller prey that could be killed and consumed more safely (e.g., swallowed whole) or that it used multiple well-targeted sagittal strikes with the beak in a repetitive attack-and-retreat strategy.

The extinct dromornithids, gastornithids and phorusrhacids are among the most spectacular birds to have ever lived, with some giants exceeding 500 kg. The affinities and evolution of these and other related extinct birds remain contentious, with previous phylogenetic analyses being affected by widespread convergence and limited taxon sampling. We address these problems using both parsimony and tip-dated Bayesian approaches on an expansive taxon set that includes all key extinct flightless and flighted (e.g. Vegavis and lithornithids) forms, an extensive array of extant fowl (Galloanseres), representative Neoaves and palaeognaths. The Paleogene volant Lithornithidae are recovered as stem palaeognaths in the Bayesian analyses. The Galloanseres comprise four clades inferred to have diverged in the Late Cretaceous on Gondwana. In addition to Anseriformes and Galliformes, we recognize a robust new clade (Gastornithiformes) for the giant flightless Dromornithidae (Australia) and Gastornithidae (Eurasia, North America). This clade exhibits parallels to ratite palaeognaths in that flight presumably was lost and giant size attained multiple times. A fourth clade is represented by the Cretaceous Vegavis (Antarctica), which was strongly excluded from Anseriformes; thus, a crucial molecular calibration point needs to be reconsidered. The presbyornithids Wilaru (Australia) and Presbyornis (Northern Hemisphere) are robustly found to be the sister group to Anatoidea (Anseranatidae + Anatidae), a relatively more basal position than hitherto recognized. South America's largest bird, Brontornis, is not a galloansere, but a member of Neoaves related to Cariamiformes; therefore, giant Galloanseres remain unknown from this continent. Trait analyses showed that while gigantism and flightlessness evolved repeatedly in groups, diet is constrained by phylogeny: all giant Galloanseres and palaeognaths are herbivores or mainly herbivorous, and giant neoavians are zoophagous or omnivorous.

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