New digital braincase endocasts of two species of Desmatosuchus and neurocranial diversity within Aetosauria (Archosauria: Pseudosuchia).

In the present contribution we revise, figure, and redescribe several isolated braincases of the iconic aetosaur Desmatosuchus from the Placerias Quarry locality, Chinle Formation, Arizona, United States. The detailed study of the isolated braincases from the UCMP collection allowed us to assign them at the species-level and recognize two species of Desmatosuchus for the Placerias Quarry: D. spurensis and D. smalli. The former can be distinguished from the latter by the presence of a transverse sulcus on the parietals, deep median pharyngeal recess on the basisphenoid, almost no gap between the basal tubera and the basipterygoid processes, and the exoccipitals meeting at the midline. The presence of D. smalli at the Placerias Quarry has not been previously reported. Based on the braincases UCMP 27408, 27410, 27407, three new brain endocasts were developed through CT scan images, reconstructing the most complete endocranial casts known for an aetosaur, including the encephalon, cranial nerves, inner ear, and endocranial vasculature. The cranial endocasts also exhibited some differences between both species of Desmatosuchus, with D. spurensis having a distinguishable dural expansion and markedly asymmetric anterior and posterior semicircular canals of the labyrinth. Additionally, the combination of osteological features and the endocranial casts allowed us to identify and discuss the presence of an ossified orbitosphenoid on the anteriormost region of the braincase among aetosaurs. Furthermore, we were able to reinterpret some of the observations made by previous authors on the endocast of the holotype of Desmatosuchus spurensis (UMMP VP 7476) and provide some insight into their neurosensory capabilities.

Endocast structures are reliable proxies for the sizes of corresponding regions of the brain in extant birds.

Endocasts are increasingly relied upon to examine avian brain evolution because they can be used across extant and extinct species. The endocasts of birds appear to be relatively faithful representatives of the external morphology of their brains, but it is unclear how well the size of a surface feature visible on endocasts reflects the volume of the underlying brain region. The optic lobe and the Wulst are two endocast structures that are clearly visible on the external surface of avian endocasts. As they overlie two major visual regions of the brain, the optic tectum and hyperpallium, the surface areas of the optic lobe and Wulst, respectively, are often used to infer visual abilities. To determine whether the surface area of these features reflects the volume of the underlying brain regions, we compared the surface areas of the optic lobes and Wulsts from digital endocasts with the volumes of the optic tecta and hyperpallia from the literature or measured from histological series of brains of the same species. Regression analyses revealed strong, statistically significant correlations between the volumes of the brain regions and the surface areas of the overlying endocast structures. In other words, the size of the hyperpallium and optic tectum can be reliably inferred from the surface areas of the Wulst and optic lobe, respectively. This validation opens the possibility of estimating brain-region volumes for extinct species in order to gain better insights in their visual ecology. It also emphasizes the importance of adopting a quantitative approach to the analysis of endocasts in the study of brain evolution.

Tempo and Pattern of Avian Brain Size Evolution.

Relative brain sizes in birds can rival those of primates, but large-scale patterns and drivers of avian brain evolution remain elusive. Here, we explore the evolution of the fundamental brain-body scaling relationship across the origin and evolution of birds. Using a comprehensive dataset sampling> 2,000 modern birds, fossil birds, and theropod dinosaurs, we infer patterns of brain-body co-variation in deep time. Our study confirms that no significant increase in relative brain size accompanied the trend toward miniaturization or evolution of flight during the theropod-bird transition. Critically, however, theropods and basal birds show weaker integration between brain size and body size, allowing for rapid changes in the brain-body relationship that set the stage for dramatic shifts in early crown birds. We infer that major shifts occurred rapidly in the aftermath of the Cretaceous-Paleogene mass extinction within Neoaves, in which multiple clades achieved higher relative brain sizes because of a reduction in body size. Parrots and corvids achieved the largest brains observed in birds via markedly different patterns. Parrots primarily reduced their body size, whereas corvids increased body and brain size simultaneously (with rates of brain size evolution outpacing rates of body size evolution). Collectively, these patterns suggest that an early adaptive radiation in brain size laid the foundation for subsequent selection and stabilization.

A Dome-Headed Stem Archosaur Exemplifies Convergence among Dinosaurs and Their Distant Relatives.

Similarities in body plan evolution, such as wings in pterosaurs, birds, and bats or limblessness in snakes and amphisbaenians, can be recognized as classical examples of convergence among animals [1-3]. We introduce a new Triassic stem archosaur that is unexpectedly and remarkably convergent with the "dome-headed" pachycephalosaur dinosaurs that lived over 100 million years later. Surprisingly, numerous additional taxa in the same assemblage (the Otis Chalk assemblage from the Dockum Group of Texas) demonstrate the early acquisition of morphological novelties that were later convergently evolved by post-Triassic dinosaurs. As one of the most successful clades of terrestrial vertebrates, dinosaurs came to occupy an extensive morphospace throughout their diversification in the Mesozoic Era [4, 5], but their distant relatives were first to evolve many of those "dinosaurian" body plans in the Triassic Period [6-8]. Our analysis of convergence between archosauromorphs from the Triassic Period and post-Triassic archosaurs demonstrates the early and extensive exploration of morphospace captured in a single Late Triassic assemblage, and we hypothesize that many of the "novel" morphotypes interpreted to occur among archosaurs later in the Mesozoic already were in place during the initial Triassic archosauromorph, largely non-dinosaurian, radiation and only later convergently evolved in diverse dinosaurian lineages.

A Basal Lithostrotian Titanosaur (Dinosauria: Sauropoda) with a Complete Skull: Implications for the Evolution and Paleobiology of Titanosauria.

We describe Sarmientosaurus musacchioi gen. et sp. nov., a titanosaurian sauropod dinosaur from the Upper Cretaceous (Cenomanian-Turonian) Lower Member of the Bajo Barreal Formation of southern Chubut Province in central Patagonia, Argentina. The holotypic and only known specimen consists of an articulated, virtually complete skull and part of the cranial and middle cervical series. Sarmientosaurus exhibits the following distinctive features that we interpret as autapomorphies: (1) maximum diameter of orbit nearly 40% rostrocaudal length of cranium; (2) complex maxilla-lacrimal articulation, in which the lacrimal clasps the ascending ramus of the maxilla; (3) medial edge of caudal sector of maxillary ascending ramus bordering bony nasal aperture with low but distinct ridge; (4) 'tongue-like' ventral process of quadratojugal that overlaps quadrate caudally; (5) separate foramina for all three branches of the trigeminal nerve; (6) absence of median venous canal connecting infundibular region to ventral part of brainstem; (7) subvertical premaxillary, procumbent maxillary, and recumbent dentary teeth; (8) cervical vertebrae with 'strut-like' centroprezygapophyseal laminae; (9) extremely elongate and slender ossified tendon positioned ventrolateral to cervical vertebrae and ribs. The cranial endocast of Sarmientosaurus preserves some of the most complete information obtained to date regarding the brain and sensory systems of sauropods. Phylogenetic analysis recovers the new taxon as a basal member of Lithostrotia, as the most plesiomorphic titanosaurian to be preserved with a complete skull. Sarmientosaurus provides a wealth of new cranial evidence that reaffirms the close relationship of titanosaurs to Brachiosauridae. Moreover, the presence of the relatively derived lithostrotian Tapuiasaurus in Aptian deposits indicates that the new Patagonian genus represents a 'ghost lineage' with a comparatively plesiomorphic craniodental form, the evolutionary history of which is missing for at least 13 million years of the Cretaceous. The skull anatomy of Sarmientosaurus suggests that multiple titanosaurian species with dissimilar cranial structures coexisted in the early Late Cretaceous of southern South America. Furthermore, the new taxon possesses a number of distinctive morphologies-such as the ossified cervical tendon, extremely pneumatized cervical vertebrae, and a habitually downward-facing snout-that have rarely, if ever, been documented in other titanosaurs, thus broadening our understanding of the anatomical diversity of this remarkable sauropod clade. The latter two features were convergently acquired by at least one penecontemporaneous diplodocoid, and may represent mutual specializations for consuming low-growing vegetation.

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.

A New Titanosaurian Braincase from the Cretaceous "Lo Hueco" Locality in Spain Sheds Light on Neuroanatomical Evolution within Titanosauria.

Despite continuous improvements, our knowledge of the neurocranial anatomy of sauropod dinosaurs as a whole is still poor, which is especially true for titanosaurians even though their postcranial remains are common in many Upper Cretaceous sites worldwide. Here we describe a braincase from the uppermost Cretaceous locality of ''Lo Hueco" in Spain that is one of the most complete titanosaurian braincases found so far in Europe. Although the titanosaurian Ampelosaurus sp. is known from the same locality, this specimen is clearly a distinct taxon and presents a number of occipital characters found in Antarctosaurus and Jainosaurus, which are approximately coeval taxa from southern Gondwana. The specimen was subjected to X-ray computed tomographic (CT) scanning, allowing the generation of 3D renderings of the endocranial cavity enclosing the brain, cranial nerves, and blood vessels, as well as the labyrinth of the inner ear. These findings add considerable knowledge to the field of sauropod paleoneuroanatomy in general and titanosaurian endocast diversity in particular. Compared with that of many sauropodomorphs, the endocast appears only slightly flexed in lateral view and bears similarities (e.g., reduction of the rostral dural expansion) with Gondwanan titanosaurians such as Jainosaurus, Bonatitan, and Antarctosaurus. The vestibular system of the inner ear is somewhat contracted (i.e., the radius of the semicircular canals is small), but less so than expected in derived titanosaurians. However, as far as the new specimen and Jainosaurus can be contrasted, and with the necessary caution due to the small sample of comparative data currently available, the two taxa appear more similar to one another in endocast morphology than to other titanosaurians. Recent phylogenetic analyses of titanosaurians have not included virtually any of the taxa under consideration here, and thus the phylogenetic position of the new Spanish titanosaurian--even its generic, let alone specific, identification--is not possible at the moment. Nevertheless, both the braincase osteology and the endocast morphology suggest that the specimen represents a derived titanosaurian that presumably branched further from the base of Lithostrotia, potentially even near Saltasauridae, comparable in evolutionary terms with Jainosaurus.

Breathing life into dinosaurs: tackling challenges of soft-tissue restoration and nasal airflow in extinct species.

The nasal region plays a key role in sensory, thermal, and respiratory physiology, but exploring its evolution is hampered by a lack of preservation of soft-tissue structures in extinct vertebrates. As a test case, we investigated members of the "bony-headed" ornithischian dinosaur clade Pachycephalosauridae (particularly Stegoceras validum) because of their small body size (which mitigated allometric concerns) and their tendency to preserve nasal soft tissues within their hypermineralized skulls. Hypermineralization directly preserved portions of the olfactory turbinates along with an internal nasal ridge that we regard as potentially an osteological correlate for respiratory conchae. Fossil specimens were CT-scanned, and nasal cavities were segmented and restored. Soft-tissue reconstruction of the nasal capsule was functionally tested in a virtual environment using computational fluid dynamics by running air through multiple models differing in nasal soft-tissue conformation: a bony-bounded model (i.e., skull without soft tissue) and then models with soft tissues added, such as a paranasal septum, a scrolled concha, a branched concha, and a model combining the paranasal septum with a concha. Deviations in fluid flow in comparison to a phylogenetically constrained sample of extant diapsids were used as indicators of missing soft tissue. Models that restored aspects of airflow found in extant diapsids, such as appreciable airflow in the olfactory chamber, were judged as more likely. The model with a branched concha produced airflow patterns closest to those of extant diapsids. These results from both paleontological observation and airflow modeling indicate that S. validum and other pachycephalosaurids could have had both olfactory and respiratory conchae. Although respiratory conchae have been linked to endothermy, such conclusions require caution in that our re-evaluation of the reptilian nasal apparatus indicates that respiratory conchae may be more widespread than originally thought, and other functions, such as selective brain temperature regulation, could be important.

Divergence in skeletal mass and bone morphology in antarctic notothenioid fishes.

Cover illustration. Notothenioid fishes lack swim bladders, but species live temporarily or permanently in the water column. Without swim bladder, skeletal mass becomes a key determinant of buoyancy. In this issue of the Journal of Morphology, Joseph Eastman and coauthors (pp. 841-861 10.1002/jmor.20258) quantify the degree of skeletal ossification in notothenioid fish and discuss it in phylogenetic context. The cover image shows a volume rendered micro-CT image showing extensive bone in the adult skeleton of the non-Antarctic notothenioid Bovichtus diacanthus from Tristan da Cunha. Tissue density is mapped in false color on a spectrum ranging from less dense (cool colors) to more dense (warm colors). Bone is considerably more extensive in B. diacanthus than in Antarctic notothenioids.

Divergence in skeletal mass and bone morphology in antarctic notothenioid fishes.

Although notothenioid fishes lack swim bladders, some species live temporarily or permanently in the water column. Given its relatively high density, skeletal mass is a key determinant of buoyancy. Notothenioids have reduced skeletal ossification, but there is little quantitative data on the phylogenetic distribution of this trait. We obtained dry skeletal masses for 54 specimens representing 20 species from six notothenioid families. Although comparative data are sparse, notothenioid skeletons comprise a smaller percentage of body mass, <3.5%, than those of three non-notothenioid perciforms. With relatively high skeletal mass, the non-Antarctic Bovichtus diacanthus is similar in skeletal mass to some non-notothenioids. Eleginops maclovinus, the non-Antarctic sister group of the Antarctic clade, has a relatively light skeleton (<2% of body mass) similar to many species in the Antarctic clade. Low skeletal mass is therefore a synapomorphy shared by Eleginops plus the Antarctic clade. We provide gross, histological, and micro-CT documentation of the structure and location of bone and cartilage in skulls, pectoral girdles, and vertebrae, with emphasis on the bovichtid B. diacanthus, the eleginopsid E. maclovinus, and the channichthyid Chaenodraco wilsoni. In Eleginops and the Antarctic clade, most bone is spongy and most species have persisting cartilage in the skull and appendicular skeleton. We also measured the relative size of the notochordal canal in adult vertebral centra of 38 species representing all eight families. There is considerable interspecific variation in this pedomorphic trait and all species show an ontogenetic reduction in the relative size of the canal. However, large persisting canals are present in adults of the Antarctic clade, especially in the nototheniids Pleuragramma and Aethotaxis and in a number of bathydraconid and channichthyid genera.

Comparative Biomechanical Modeling of Metatherian and Placental Saber-Tooths: A Different Kind of Bite for an Extreme Pouched Predator.

Questions surrounding the dramatic morphology of saber-tooths, and the presumably deadly purpose to which it was put, have long excited scholarly and popular attention. Among saber-toothed species, the iconic North American placental, Smilodon fatalis, and the bizarre South American sparassodont, Thylacosmilus atrox, represent extreme forms commonly forwarded as examples of convergent evolution. For S. fatalis, some consensus has been reached on the question of killing behaviour, with most researchers accepting the canine-shear bite hypothesis, wherein both head-depressing and jaw closing musculatures played a role in delivery of the fatal bite. However, whether, or to what degree, T. atrox may have applied a similar approach remains an open question. Here we apply a three-dimensional computational approach to examine convergence in mechanical performance between the two species. We find that, in many respects, the placental S. fatalis (a true felid) was more similar to the metatherian T. atrox than to a conical-toothed cat. In modeling of both saber-tooths we found that jaw-adductor-driven bite forces were low, but that simulations invoking neck musculature revealed less cranio-mandibular stress than in a conical-toothed cat. However, our study also revealed differences between the two saber-tooths likely reflected in the modus operandi of the kill. Jaw-adductor-driven bite forces were extremely weak in T. atrox, and its skull was even better-adapted to resist stress induced by head-depressors. Considered together with the fact that the center of the arc described by the canines was closer to the jaw-joint in Smilodon, our results are consistent with both jaw-closing and neck musculature playing a role in prey dispatch for the placental, as has been previously suggested. However, for T. atrox, we conclude that the jaw-adductors probably played no major part in the killing bite. We propose that the metatherian presents a more complete commitment to the already extreme saber-tooth 'lifestyle'.

Neurocranial osteology and neuroanatomy of a late Cretaceous titanosaurian sauropod from Spain (Ampelosaurus sp.).

Titanosaurians were a flourishing group of sauropod dinosaurs during Cretaceous times. Fossils of titanosaurians have been found on all continents and their remains are abundant in a number of Late Cretaceous sites. Nonetheless, the cranial anatomy of titanosaurians is still very poorly known. The Spanish latest Cretaceous locality of "Lo Hueco" yielded a relatively well preserved, titanosaurian braincase, which shares a number of phylogenetically restricted characters with Ampelosaurus atacis from France such as a flat occipital region. However, it appears to differ from A. atacis in some traits such as the greater degree of dorsoventral compression and the presence of proatlas facets. The specimen is, therefore, provisionally identified as Ampelosaurus sp. It was CT scanned, and 3D renderings of the cranial endocast and inner-ear system were generated. Our investigation highlights that, although titanosaurs were derived sauropods with a successful evolutionary history, they present a remarkably modest level of paleoneurological organization. Compared with the condition in the basal titanosauriform Giraffatitan brancai, the labyrinth of Ampelosaurus sp. shows a reduced morphology. The latter feature is possibly related to a restricted range of head-turning movements.

The braincase of the basal sauropod dinosaur Spinophorosaurus and 3D reconstructions of the cranial endocast and inner ear.

BACKGROUND: Sauropod dinosaurs were the largest animals ever to walk on land, and, as a result, the evolution of their remarkable adaptations has been of great interest. The braincase is of particular interest because it houses the brain and inner ear. However, only a few studies of these structures in sauropods are available to date. Because of the phylogenetic position of Spinophorosaurus nigerensis as a basal eusauropod, the braincase has the potential to provide key evidence on the evolutionary transition relative to other dinosaurs. METHODOLOGY/PRINCIPAL FINDINGS: The only known braincase of Spinophorosaurus ('Argiles de l'Irhazer', Irhazer Group; Agadez region, Niger) differs significantly from those of the Jurassic sauropods examined, except potentially for Atlasaurus imelakei (Tilougguit Formation, Morocco). The basisphenoids of Spinophorosaurus and Atlasaurus bear basipterygoid processes that are comparable in being directed strongly caudally. The Spinophorosaurus specimen was CT scanned, and 3D renderings of the cranial endocast and inner-ear system were generated. The endocast resembles that of most other sauropods in having well-marked pontine and cerebral flexures, a large and oblong pituitary fossa, and in having the brain structure obscured by the former existence of relatively thick meninges and dural venous sinuses. The labyrinth is characterized by long and proportionally slender semicircular canals. This condition recalls, in particular, that of the basal non-sauropod sauropodomorph Massospondylus and the basal titanosauriform Giraffatitan. CONCLUSIONS/SIGNIFICANCE: Spinophorosaurus has a moderately derived paleoneuroanatomical pattern. In contrast to what might be expected early within a lineage leading to plant-eating graviportal quadrupeds, Spinophorosaurus and other (but not all) sauropodomorphs show no reduction of the vestibular apparatus of the inner ear. This character-state is possibly a primitive retention in Spinophorosaurus, but due the scarcity of data it remains unclear whether it is also the case in the various later sauropods in which it is present or whether it has developed homoplastically in these taxa. Any interpretations remain tentative pending the more comprehensive quantitative analysis underway, but the size and morphology of the labyrinth of sauropodomorphs may be related to neck length and mobility, among other factors.

Evolution of olfaction in non-avian theropod dinosaurs and birds.

Little is known about the olfactory capabilities of extinct basal (non-neornithine) birds or the evolutionary changes in olfaction that occurred from non-avian theropods through modern birds. Although modern birds are known to have diverse olfactory capabilities, olfaction is generally considered to have declined during avian evolution as visual and vestibular sensory enhancements occurred in association with flight. To test the hypothesis that olfaction diminished through avian evolution, we assessed relative olfactory bulb size, here used as a neuroanatomical proxy for olfactory capabilities, in 157 species of non-avian theropods, fossil birds and living birds. We show that relative olfactory bulb size increased during non-avian maniraptoriform evolution, remained stable across the non-avian theropod/bird transition, and increased during basal bird and early neornithine evolution. From early neornithines through a major part of neornithine evolution, the relative size of the olfactory bulbs remained stable before decreasing in derived neoavian clades. Our results show that, rather than decreasing, the importance of olfaction actually increased during early bird evolution, representing a previously unrecognized sensory enhancement. The relatively larger olfactory bulbs of earliest neornithines, compared with those of basal birds, may have endowed neornithines with improved olfaction for more effective foraging or navigation skills, which in turn may have been a factor allowing them to survive the end-Cretaceous mass extinction.

Cartilaginous epiphyses in extant archosaurs and their implications for reconstructing limb function in dinosaurs.

Extinct archosaurs, including many non-avian dinosaurs, exhibit relatively simply shaped condylar regions in their appendicular bones, suggesting potentially large amounts of unpreserved epiphyseal (articular) cartilage. This "lost anatomy" is often underappreciated such that the ends of bones are typically considered to be the joint surfaces, potentially having a major impact on functional interpretation. Extant alligators and birds were used to establish an objective basis for inferences about cartilaginous articular structures in such extinct archosaur clades as non-avian dinosaurs. Limb elements of alligators, ostriches, and other birds were dissected, disarticulated, and defleshed. Lengths and condylar shapes of elements with intact epiphyses were measured. Limbs were subsequently completely skeletonized and the measurements repeated. Removal of cartilaginous condylar regions resulted in statistically significant changes in element length and condylar breadth. Moreover, there was marked loss of those cartilaginous structures responsible for joint architecture and congruence. Compared to alligators, birds showed less dramatic, but still significant changes. Condylar morphologies of dinosaur limb bones suggest that most non-coelurosaurian clades possessed large cartilaginous epiphyses that relied on the maintenance of vascular channels that are otherwise eliminated early in ontogeny in smaller-bodied tetrapods. A sensitivity analysis using cartilage correction factors (CCFs) obtained from extant taxa indicates that whereas the presence of cartilaginous epiphyses only moderately increases estimates of dinosaur height and speed, it has important implications for our ability to infer joint morphology, posture, and the complicated functional movements in the limbs of many extinct archosaurs. Evidence suggests that the sizes of sauropod epiphyseal cartilages surpassed those of alligators, which account for at least 10% of hindlimb length. These data suggest that large cartilaginous epiphyses were widely distributed among non-avian archosaurs and must be considered when making inferences about locomotor functional morphology in fossil taxa.

The evolution of mammal-like crocodyliforms in the Cretaceous Period of Gondwana.

Fossil crocodyliforms discovered in recent years have revealed a level of morphological and ecological diversity not exhibited by extant members of the group. This diversity is particularly notable among taxa of the Cretaceous Period (144-65 million years ago) recovered from former Gondwanan landmasses. Here we report the discovery of a new species of Cretaceous notosuchian crocodyliform from the Rukwa Rift Basin of southwestern Tanzania. This small-bodied form deviates significantly from more typical crocodyliform craniodental morphologies, having a short, broad skull, robust lower jaw, and a dentition with relatively few teeth that nonetheless show marked heterodonty. The presence of morphologically complex, complementary upper and lower molariform teeth suggests a degree of crown-crown contact during jaw adduction that is unmatched among known crocodyliforms, paralleling the level of occlusal complexity seen in mammals and their extinct relatives. The presence of another small-bodied mammal-like crocodyliform in the Cretaceous of Gondwana indicates that notosuchians probably filled niches and inhabited ecomorphospace that were otherwise occupied by mammals on northern continents.

New insights into the brain, braincase, and ear region of tyrannosaurs (Dinosauria, Theropoda), with implications for sensory organization and behavior.

The braincase region of tyrannosaurs was investigated to provide insights on anatomical attributes relevant to inferences of sensory biology and behavior. CT scanning focused on three specimens of Tyrannosaurus rex, a juvenile Gorgosaurus, and the controversial Cleveland skull (CMNH 7541). Analysis shows that the cerebral hemispheres were enlarged, but conflicting information on the optic lobes suggests that brain conformation was not fully avian. Previous estimates of olfactory bulb size for T. rex were much too large, but even the corrected sizes are relatively larger than other theropods, suggesting that odor detection was indeed of particular importance to tyrannosaurs. The inner ears show a number of coelurosaurian traits, such as elongate and rounded and rostral, lateral semicircular canals, and incipient twisting of the common crus, which we interpret to be related to enhanced reflexes coordinating rapid eye and head movements. The cochlea is elongate, which, coupled with the finding of extensive tympanic pneumaticity, supports the inference of behavioral emphasis of low-frequency sounds. Three main groups of sinuses pneumatized the braincase, and there are a number of perhaps systematically relevant differences. Orientation of the endosseous labyrinth reveals that alert head postures of T. rex and Gorgosaurus were somewhat depressed below the horizontal, but the Cleveland skull had a very strongly down-turned posture. It is concluded that tyrannosaur sensory biology is consistent with their predatory coelurosaurian heritage, with emphasis on relatively quick, coordinated eye and head movements, and probably sensitive low-frequency hearing; tyrannosaurs apomorphically enhanced their olfactory apparatus. The taxonomic status of the Cleveland skull remains unresolved.

Endocranial anatomy of lambeosaurine hadrosaurids (Dinosauria: Ornithischia): a sensorineural perspective on cranial crest function.

Brain and nasal cavity endocasts of four corythosaurian lambeosaurines (Dinosauria: Ornithischia) were investigated to test hypotheses of cranial crest function related to sensorineural systems. Endocasts were generated through computed tomography and three-dimensional rendering and visualization software. The sample comprises a range of ontogenetic stages from the taxa Lambeosaurus, Corythosaurus, and Hypacrosaurus. Results show that the morphology of brain endocasts differs little from that of hadrosaurines. The strikingly convoluted nasal vestibule of Hypacrosaurus altispinus, when interpreted in the context of lambeosaurine phylogeny, suggests selective pressure for nasal cavity function independent from changes in the external shape of the crest and associated visual display function. The plesiomorphically small olfactory bulbs and apparently small olfactory region of the nasal cavity argues against the hypothesis that increased olfactory acuity played a causal role in crest evolution. The elongate cochlea of the inner ear reveals that hearing in lambeosaurines emphasized low frequencies consistent with the hypothesized low-frequency calls made by the crests under the resonation model of crest function. The brain is relatively large in lambeosaurines compared with many other large dinosaurs, and the cerebrum is relatively larger than that of all non-hadrosaurian ornithischians and large theropods, but compares favorably with hadrosaurine hadrosaurids as well as some maniraptoran theropods. It is concluded that the large brains of lambeosaurines are consistent with the range of social behaviors inferred when the crest is interpreted as an intraspecific signaling structure.

The paranasal air sinuses of predatory and armored dinosaurs (archosauria: theropoda and ankylosauria) and their contribution to cephalic structure.

The paranasal air sinuses and nasal cavities were studied along with other cephalic spaces (brain cavity, paratympanic sinuses) in certain dinosaurs via CT scanning and 3D visualization to document the anatomy and examine the contribution of the sinuses to the morphological organization of the head as a whole. Two representatives each of two dinosaur clades are compared: the theropod saurischians Majungasaurus and Tyrannosaurus and the ankylosaurian ornithischians Panoplosaurus and Euoplocephalus. Their extant archosaurian outgroups, birds and crocodilians (exemplified by ostrich and alligator), display a diversity of paranasal sinuses, yet they share only a single homologous antorbital sinus, which in birds has an important subsidiary diverticulum, the suborbital sinus. Both of the theropods had a large antorbital sinus that pneumatized many of the facial and palatal bones as well as a birdlike suborbital sinus. Given that the suborbital sinus interleaves with jaw muscles, the paranasal sinuses of at least some theropods (including birds) were actively ventilated rather than being dead-air spaces. Although many ankylosaurians have been thought to have had extensive paranasal sinuses, most of the snout is instead (and surprisingly) often occupied by a highly convoluted airway. Digital segmentation, coupled with 3D visualization and analysis, allows the positions of the sinuses to be viewed in place within both the skull and the head and then measured volumetrically. These quantitative data allow the first reliable estimates of dinosaur head mass and an assessment of the potential savings in mass afforded by the sinuses.

Cephalic vascular anatomy in flamingos (Phoenicopterus ruber) based on novel vascular injection and computed tomographic imaging analyses.

Head vascular anatomy of the greater (or Caribbean) flamingo (Phoenicopterus ruber) is investigated and illustrated through the use of a differential contrast, dual vascular injection technique, and high-resolution X-ray computed tomography (CT), allowing arteries and veins to be differentiated radiographically. Vessels were digitally isolated with segmentation tools and reconstructed in 3D to facilitate topographical visualization of the cephalic vascular tree. Major vessels of the temporal, orbital, pharyngeal, and encephalic regions are described and illustrated, which confirm that the general pattern of avian cephalic vasculature is evolutionarily conservative. In addition to numerous arteriovenous vascular devices, a previously undescribed, large, bilateral, paralingual cavernous sinus that excavates a large bony fossa on the medial surface of the mandible was identified. Despite the otherwise conservative vascular pattern, this paralingual sinus was found only in species of flamingo and is not known otherwise in birds. The paralingual sinus remains functionally enigmatic, but a mechanical role in association with the peculiar lingual-pumping mode of feeding in flamingos is perhaps the most likely hypothesis.