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The morphology of the temporal region in the tetrapod skull traditionally has been a widely discussed feature of vertebrate anatomy. The evolution of different temporal openings in Amniota (mammals, birds, and reptiles), Lissamphibia (frogs, salamanders, and caecilians), and several extinct tetrapod groups has sparked debates on the phylogenetic, developmental, and functional background of this region in the tetrapod skull. This led most famously to the erection of different amniote taxa based on the number and position of temporal fenestrae in their skulls. However, most of these taxa are no longer recognised to represent natural groupings and the morphology of the temporal region is not necessarily an adequate trait for use in the reconstruction of amniote phylogenies. Yet, new fossil finds, most notably of parareptiles and stem-turtles, as well as modern embryological and biomechanical studies continue to provide new insights into the morphological diversity of the temporal region. Here, we introduce a novel comprehensive classification scheme for the various temporal morphotypes in all Tetrapoda that is independent of phylogeny and previous terminology and may facilitate morphological comparisons in future studies. We then review the history of research on the temporal region in the tetrapod skull. We document how, from the early 19th century with the first recognition of differences in the temporal region to the first proposals of phylogenetic relationships and their assessment over the centuries, the phylogenetic perspective on the temporal region has developed, and we highlight the controversies that still remain. We also compare the different functional and developmental drivers proposed for the observed morphological diversity and how the effects of internal and external factors on the structure of the tetrapod skull have been interpreted., (© 2021 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Comparative osteological analyses of extant organisms provide key insight into major evolutionary transitions and phylogenetic hypotheses. This is especially true for snakes, given their unique morphology relative to other squamates and the persistent controversy regarding their evolutionary origins. However, the osteology of several major snake groups remains undescribed, thus hindering efforts to accurately reconstruct the phylogeny of snakes. One such group is the Atractaspididae, a family of fossorial colubroids. We herein present the first detailed description of the atractaspidid skull, based on fully segmented micro-computed tomography (micro-CT) scans of Atractaspis irregularis. The skull of Atractaspis presents a highly unique morphology influenced by both fossoriality and paedomorphosis. This paedomorphosis is especially evident in the jaws, palate, and suspensorium, the major elements associated with macrostomy (large-gaped feeding in snakes). Comparison to scolecophidians-a group of blind, fossorial, miniaturized snakes-in turn sheds light on current hypotheses of snake phylogeny. Features of both the naso-frontal joint and the morphofunctional system related to macrostomy refute the traditional notion that scolecophidians are fundamentally different from alethinophidians (all other extant snakes). Instead, these features support the controversial hypothesis of scolecophidians as "regressed alethinophidians," in contrast to their traditional placement as the earliest-diverging snake lineage. We propose that Atractaspis and scolecophidians fall along a morphological continuum, characterized by differing degrees of paedomorphosis. Altogether, a combination of heterochrony and miniaturization provides a mechanism for the derivation of the scolecophidian skull from an ancestral fossorial alethinophidian morphotype, exemplified by the nonminiaturized and less extreme paedomorph Atractaspis., (© 2020 Anatomical Society.)

Choristoderes are extinct neodiapsid reptiles that are well known for their unusual cranial anatomy, possessing an elongated snout and expanded temporal arches. Although choristodere skulls are well described externally, their internal anatomy remains unknown. An internal description was needed to shed light on peculiarities of the choristodere skull, such as paired gaps on the ventral surface of the skull that may pertain to the fenestra ovalis, and a putative neomorphic ossification in the lateral wall of the braincase. Our goals were: (i) to describe the cranial elements of Champsosaurus lindoei in three dimensions; (ii) to describe paired gaps on the ventral surface of the skull to determine if these are indeed the fenestrae ovales; (iii) to illustrate the morphology of the putative neomorphic bone; and (iv) to consider the possible developmental and functional origins of the neomorph. We examined the cranial anatomy of the choristodere Champsosaurus lindoei (CMN 8920) using high-resolution micro-computed tomography scanning. We found that the paired gaps on the ventral surface of the skull do pertain to the fenestrae ovales, an unusual arrangement that may be convergent with some plesiosaurs, some aistopods, and some urodeles. The implications of this morphology in Champsosaurus are unknown and will be the subject of future work. We found that the neomorphic bone is a distinct ossification, but is not part of the wall of the brain cavity or the auditory capsule. Variation in the developmental pathways of cranial bones in living amniotes was surveyed to determine how the neomorphic bone may have developed. We found that the chondrocranium and splanchnocranium show little to no variation across amniotes, and the neomorphic bone is therefore most likely to have developed from the dermatocranium; however, the stapes is a pre-existing cranial element that is undescribed in choristoderes and may be homologous with the neomorphic bone. If the neomorphic bone is not homologous with the stapes, the neomorph likely developed from the dermatocranium through incomplete fusion of ossification centres from a pre-existing bone, most likely the parietal. Based on the apparent morphology of the neomorph in Coeruleodraco, the neomorph was probably too small to play a significant structural role in the skull of early choristoderes and it may have arisen through non-adaptive means. In neochoristoderes, such as Champsosaurus, the neomorph was likely recruited to support the expanded temporal arches., (© 2020 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

Embolomeres have been considered candidate stem-amniotes by many authors, but recent work has shown that anatomical and phylogenetic support for this hypothesis may be ambiguous. Newer phylogenetic approaches focus on the anatomy of the endochondral bones surrounding the brain (neurocranium) and associated dermal investing bones, so accurate description of the neurocranium of tetrapods, and particularly embolomeres, is critical to assess the phylogenetic context of amniote origins. I here amend prior descriptions of the neurocranium of the abundant Early Permian embolomere, Archeria crassidisca , with a description of a previously unreported partial braincase. This specimen reveals features broadly associated with stem-tetrapods of the fin-to-limb transition and recently described in aïstopod stem-tetrapods, including a cranial notochordal remnant, a possible buccohypophyseal canal, and anteroventrally projecting saccular fossae that extend deep into the basioccipital. I present an amended description and reconstruction of the neurocranium and a phylogenetic analysis integrating these codings, which recovers embolomeres more robustly within the tetrapod stem. I review characters previously used to assign embolomeres to the amniote stem and find this evidence is ambiguous or incorrect. I then outline strong character support for a stem-tetrapod position of embolomeres, and implications for the ancestral body plan of crown-group tetrapods. [ABSTRACT FROM AUTHOR]

Feeding efficiency of mammalian herbivores is associated with skull traits known to differ between feeding guilds. As climate is known to affect the architecture and functional traits of plants, skull morphology may change as plant traits change along climate gradients. We tested whether skull traits of more than 70% of the extant bovid and cervid species are related to aridity, and whether these relationships differ across feeding guilds (browser, grazer, mixed feeder). We used muzzle width to represent food ingestion and masseteric fossa length to represent oral processing. For the models without phylogeny, both muzzle width and masseteric fossa length increased towards wetter climates for grazers, but not for mixed feeders or browsers. These patterns suggest that grazer diets may change more dramatically across aridity gradients than the other two groups, possibly linked to changes in both the architecture and toughness of grasses. In models accounting for phylogeny, no changes in either trait across aridity gradient were found, which appeared to be due to subfamily differentiation along the aridity gradient. Our results suggest that deeper insights into the evolution of herbivore skulls may be achieved by using quantified diet data to directly test plant trait effects on skull morphology. [ABSTRACT FROM AUTHOR]

The unsolved problem in three-dimensional surgical planning for patients with facial deformity, dysgnathia, or asymmetry is the lack of a normative database of "norm skulls" that can be used as treatment objectives. A study was conducted on 90 Eurasian persons (46 male and 44 female adults) for whom cone beam-computed tomography images were available. Inclusion criteria were adult patients with a skeletal Class I pattern, proper interincisal relationship with normal occlusion, the absence of an open bite both in the anterior and posterior region, and a normal and balanced facial appearance; patients with dysgnathia and malformations were excluded. A total of 18 landmarks were digitized and 3D cephalometric measurements were performed and analyzed by means of proportions calculated from the landmarks. Male and female skulls were analyzed, as well as subdivisions revealed by cluster analysis. The data showed that four subtypes of skulls were distinguishable with statistical significance (p < 0.05). A male and a female type subdivided in a brachiocephalic and dolichocephalic phenotype could be identified. For each type, a mean shape was calculated by a Procrustes transformation, which, in turn, was used to create four template skulls from a male and a female skull. This was accomplished by fitting the polygon models of the two skulls to each of the two subtypes based on the landmarks marked on them using a thin plate spline transformation. The normative data of the subtypes can individually serve as a guide for orthodontic surgery in the Eurasian population, which is especially helpful in 3D planning and the execution of craniofacial operations. [ABSTRACT FROM AUTHOR]

In recent decades, the application and adaptation of new technologies have gradually become invaluable in education. When the students are visually impaired, the use of these technologies becomes even more valuable. In this line, a theoretical-practical workshop on human evolution was carried out with 25 students from the Spanish National Organisation for the Blind. Twenty-two realistic replicas of skulls of the main species in human evolution were reproduced with a 3D-FDM printer. Thus, the students were able to learn about the variability of cranial anatomy between species as we could verify with a test to evaluate the usefulness of the technology as a complement to the theoretical explanation. The results were analysed taking into account the sex and age of the students, their educational level, their degree of blindness and previous knowledge of the subject. It showed that the use of 3D models is an excellent support to the teaching-learning process in the case of the visually impaired, since it allows them to form in their imagination a faithful image of the object they hold in their hands, to feel the similarities and morphological differences between the species and to formulate hypotheses about the reasons for them. [ABSTRACT FROM AUTHOR]

Rhynchosaurs were key herbivores over much of the world in the Middle and Late Triassic, often dominating their faunas ecologically. The anatomy of the Middle Triassic forms is debated in certain phylogenetically important details. Among several genera from the Anisian of England, Bentonyx sidensis is represented by a near-complete skull. New CT scanning of the holotype skull reveals previously concealed details of the palate and braincase in particular, and this allows comparison with other Middle Triassic rhynchosaurs and confirmation that Bentonyx is a distinct taxon from others of similar age including Fodonyx, Rhynchosaurus, Langeronyx, and Stenaulorhynchus. These are part of the first wave of rhynchosaur diversification in the Middle Triassic. [ABSTRACT FROM AUTHOR]

The Triassic basal sauropodomorph Plateosaurus trossingensis is well-known from mass accumulations at the German localities of Trossingen and Halberstadt and the Swiss locality of Frick, and is significant especially regarding its taphonomy and proposed developmental plasticity. These implications, however, rely on the assumption that this material derives from a single species, which has been questioned. Here we describe new skull material from Frick including eight complete and six partial skulls, more than doubling the number of known skulls of P. trossingensis. This exceptional sample size allows for gaining a deeper understanding of variability that may occur in a single species. The new material includes the first known juvenile skulls of Plateosaurus, allowing for detecting ontogenetic changes. An attempt is made to distinguish between variability caused by taphonomic plastic deformation and intraspecific variability. Plastic deformation may shorten, but not widen bones. A number of characters commonly included in phylogenetic analyses of basal sauropodomorphs are shown to be variable within P. trossingensis, and possibly require re-evaluation. Although P. trossingensis skulls are highly variable, many of the variable characters include intermediate character states and therefore are continuous. No groupings based on skull features, locality, or stratigraphy are apparent. Consequently, the analyzed skull material from the bonebeds of Frick, Trossingen, and Halberstadt bears no evidence for the presence of more than one species. [ABSTRACT FROM AUTHOR]

In early tetrapods, it is assumed that the tympana were acoustically coupled through the pharynx and therefore inherently directional, acting as pressure difference receivers. The later closure of the middle ear cavity in turtles, archosaurs, and mammals is a derived condition, and would have changed the ear by decoupling the tympana. Isolation of the middle ears would then have led to selection for structural and neural strategies to compute sound source localization in both archosaurs and mammalian ancestors. In the archosaurs (birds and crocodilians) the presence of air spaces in the skull provided connections between the ears that have been exploited to improve directional hearing, while neural circuits mediating sound localization are well developed. In this review, we will focus primarily on directional hearing in crocodilians, where vocalization and sound localization are thought to be ecologically important, and indicate important issues still awaiting resolution. This article is part of a Special Issue entitled < Annual Reviews 2015 > . [ABSTRACT FROM AUTHOR]

Abstract: The Early Permian mesosaurids are the oldest known aquatic amniotes with an exclusively Gondwanan distribution. Although several hundred of complete skeletons have been discovered and intensively studied, the anatomy and taxonomic composition of the group, as well as its phylogenetic relationships remain controversial. Several well-preserved mesosaurid specimens found in Uruguay justify a new anatomical reconstruction of the skull of Mesosaurus tenuidens, differing from earlier ones especially in the presence of a lower temporal fenestra. The significance of this structure for the evolution of temporal fenestration in amniotes is evaluated according to the two most recent phylogenetic hypotheses, in which mesosaurids are basalmost sauropsids or basalmost parareptiles. A synapsid-like fenestration may be the primitive condition for Amniota, and it may be also a basal condition for parareptiles, because recent phylogenies suggest a basal position for mesosaurids and lanthanosuchoids within that group, and both possess a lower temporal fenestra. Our results also give a moderately strengthened support for diapsid affinities of turtles. [Copyright &y& Elsevier]

Abstract: Octodontoidea is the most diverse group of caviomorph rodents. The systematics of most of the fossil representatives has been essentially based upon dental characters. Described here is an almost complete skull with dentition assigned to Prospaniomys Ameghino based upon its dental morphology. The specimen comes from the Sarmiento Formation at Pampa de Gan Gan (central Patagonia, Argentina), assigned to the Colhuehuapian SALMA (early Miocene). The most remarkable features are in the posterior portion of the skull, some of them shared with the modern octodontids and interpreted as specialized by previous authors, which contrast with the generalized dental morphology. These combined features were not previously known in other octodontoids. The comparisons with other fossil and extant members of the superfamily suggest that the characters traditionally used to associate Prospaniomys with the echimyids are very probably plesiomorphies. Prospaniomys would represent an early diverging lineage more closely related to modern octodontids than to echimyids, in which cranial structures evolved more rapidly than dental and mandibular ones. [Copyright &y& Elsevier]

Uropeltid snakes are among the most poorly understood clades within Alethinophidia. Their small size, limited geographic distribution, high incidence of endemism, and fossorial behaviour all contribute to the general paucity of systematic collections of these snakes, especially of adequate skeletal preparations, in most museum collections. Their hypothesized position within the higher-order phylogeny of alethinophidian snakes calls attention to the need for additional morphological work on the group. Hypotheses of uropeltid phylogenetic relationships based on morphological analyses are few, and continue to be hampered by limited taxon-sampling and character matrices that rely predominantly on features that are visible on articulated skulls. We utilized high-resolution X-ray computed tomography (HRCT) to investigate the cranial osteology of Plectrurus aureus Beddome, 1880, a species for which no osteological data were previously available. We provide a detailed description of the skull and mandible, and comment on morphological characters and potential phylogenetic relationships. Clarity in character descriptions is of paramount importance, and additional morphological characters are desirable. HRCT provides a nondestructive way to identify new systematically informative morphological characters from digitally disarticulated specimens. The small size of many uropeltid species, including P. aureus, will help to frame a greater appreciation of the limitations of traditional HRCT protocols for revealing detailed anatomical features of small vertebrates. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 118–138. [ABSTRACT FROM AUTHOR]

The present study measured the position of the greater palatine foramen relative to adjacent anatomical landmarks in Brazilian skulls. The perpendicular distance of the greater palatine foramen to the midline maxillary suture in Brazilian skulls was about 14 mm and the distance of greater palatine foramen to the incisive foramen was approximately 36 mm. The distance of greater palatine foramen to the posterior border of the hard palate was approximately 3 mm, and the mean angle between the midline maxillary suture and the line from the incisive foramen and the greater palatine foramen was 22.71°. In almost 70% of the cases, the greater palatine foramen opened in an anterior direction. The mean palatine length was approximately 52 mm. In the greater majority of the skulls (93.81%), the greater palatine foramina were opposite or distal to the maxillary third molar. These data will be helpful in comparing these skulls to those from various other regions as well as comparing skulls of different races. It can also provide professionals with anatomical references, in order to block the maxillary division of the trigeminal nerve through the greater palatine foramen. Our results would help clinicians locate the greater palatine foramen in patients with and without upper molars. [ABSTRACT FROM AUTHOR]

Background Context: The halo skeletal fixator provides the most rigid type of immobilization of all the orthoses that stabilize the cervical spine. Sometimes with older patients (>70 years old), the pin penetrates the cortical and cancellous bone of the skull and enters the intracranial space, which can result in serious complications such as brain injury, infection, hematoma, and loss of cerebrospinal fluid from the subarachnoid space. Currently, there is a lack of relevant literature that examines these concerns. Purpose: To evaluate the thickness of the outer table, diploe, and inner table at the anterolateral and posterolateral pin insertion areas of the skull in elderly cadavers by using computed tomography (CT) scans. In addition, insertion torques at the four standard pin insertion areas was determined by applying halo pins at incremental torque in an effort to suggest safe torque levels for the anterolateral and posterolateral pins. Study Design/setting: A human cadaveric anatomical and biomechanical study relating to thickness and insertion torques at standard pin areas in the elderly. Patient Sample: Twenty-one elderly cadaveric skull specimens. Outcome Measures: Thickness of cortices (tables) and diploe of skull and insertion torque at halo pin insertion areas. Methods: Aquarius Image software at the CT scanner's TeraRecon Aquarius Workstation was used to make the necessary skull thickness measurements at the pin insertion areas. Six, 8, 12, 18, and 36 inch lb of torque were used to determine penetration of the pins through the inner table at each of the four locations (two anterolateral and two posterolateral). Results: The mean anterolateral thickness was 7.36+/-1.57 mm. The average thickness of the outer table, diploe, and inner table were 2.24+/-0.44 mm, 1.52+/-0.41 mm, and 3.59+/-1.70 mm, respectively. The mean posterolateral thickness was 9.47+/-1.12 mm. The average thickness of the outer table, diploe, and inner table were 4.32+/-0.92 mm, 1.88+/-0.35 mm, and 3.27+/-1.21 mm, respectively. No pin penetration was seen at the traditional 8 inch lb of insertion torque in both the anterolateral and posterolateral pin insertion areas. Eighteen inch lb of torque resulted in penetration in 90.48% (19/21) and in 85.71% (18/21) of specimens in the left anterolateral and right anterolateral pin insertion areas, respectively. No penetration was seen even at 36 inch lb of torque in 80.95% (17/21) of the cadavers in both the left and right posterolateral pin insertion areas. Conclusions: The current study supported previous research that 8 inch lb of torque is safe for application of halo pins in the elderly. The posterolateral skull is thicker and stronger than the anterolateral skull. The safe maximum torque is 8 inch lb for anterolateral pin insertion area and 18 inch lb for the posterolateral pin insertion area. [ABSTRACT FROM AUTHOR]