Inner ear biomechanics reveals a Late Triassic origin for mammalian endothermy.

Endothermy underpins the ecological dominance of mammals and birds in diverse environmental settings1,2. However, it is unclear when this crucial feature emerged during mammalian evolutionary history, as most of the fossil evidence is ambiguous3-17. Here we show that this key evolutionary transition can be investigated using the morphology of the endolymph-filled semicircular ducts of the inner ear, which monitor head rotations and are essential for motor coordination, navigation and spatial awareness18-22. Increased body temperatures during the ectotherm-endotherm transition of mammal ancestors would decrease endolymph viscosity, negatively affecting semicircular duct biomechanics23,24, while simultaneously increasing behavioural activity25,26 probably required improved performance27. Morphological changes to the membranous ducts and enclosing bony canals would have been necessary to maintain optimal functionality during this transition. To track these morphofunctional changes in 56 extinct synapsid species, we developed the thermo-motility index, a proxy based on bony canal morphology. The results suggest that endothermy evolved abruptly during the Late Triassic period in Mammaliamorpha, correlated with a sharp increase in body temperature (5-9 °C) and an expansion of aerobic and anaerobic capacities. Contrary to previous suggestions3-14, all stem mammaliamorphs were most probably ectotherms. Endothermy, as a crucial physiological characteristic, joins other distinctive mammalian features that arose during this period of climatic instability28.

Evolution of inner ear neuroanatomy of bats and implications for echolocation.

Phylogenomics of bats suggests that their echolocation either evolved separately in the bat suborders Yinpterochiroptera and Yangochiroptera, or had a single origin in bat ancestors and was later lost in some yinpterochiropterans1-6. Hearing for echolocation behaviour depends on the inner ear, of which the spiral ganglion is an essential structure. Here we report the observation of highly derived structures of the spiral ganglion in yangochiropteran bats: a trans-otic ganglion with a wall-less Rosenthal's canal. This neuroanatomical arrangement permits a larger ganglion with more neurons, higher innervation density of neurons and denser clustering of cochlear nerve fascicles7-13. This differs from the plesiomorphic neuroanatomy of Yinpterochiroptera and non-chiropteran mammals. The osteological correlates of these derived ganglion features can now be traced into bat phylogeny, providing direct evidence of how Yangochiroptera differentiated from Yinpterochiroptera in spiral ganglion neuroanatomy. These features are highly variable across major clades and between species of Yangochiroptera, and in morphospace, exhibit much greater disparity in Yangochiroptera than Yinpterochiroptera. These highly variable ganglion features may be a neuroanatomical evolutionary driver for their diverse echolocating strategies4,14-17 and are associated with the explosive diversification of yangochiropterans, which include most bat families, genera and species.

Structure and scaling of the middle ear in domestic dog breeds.

Although domestic dogs vary considerably in both body size and skull morphology, behavioural audiograms have previously been found to be similar in breeds as distinct as a Chihuahua and a St Bernard. In this study, we created micro-CT reconstructions of the middle ears and bony labyrinths from the skulls of 17 dog breeds, including both Chihuahua and St Bernard, plus a mongrel and a wolf. From these reconstructions, we measured middle ear cavity and ossicular volumes, eardrum and stapes footplate areas and bony labyrinth volumes. All of these ear structures scaled with skull size with negative allometry and generally correlated better with condylobasal length than with maximum or interaural skull widths. Larger dogs have larger ear structures in absolute terms: the volume of the St Bernard's middle ear cavity was 14 times that of the Chihuahua. The middle and inner ears are otherwise very similar in morphology, the ossicular structure being particularly well-conserved across breeds. The expectation that larger ear structures in larger dogs would translate into hearing ranges shifted towards lower frequencies is not consistent with the existing audiogram data. Assuming that the audiograms accurately reflect the hearing of the breeds in question, oversimplifications in existing models of middle ear function or limitations imposed by other parts of the auditory system may be responsible for this paradox.

The endoscopic transcanal approach to the internal auditory canal: an anatomic study.

PURPOSE: The internal auditory canal (IAC) plays a key role in lateral skull base surgery. Although several approaches to the IAC have been proposed, endoscope-assisted transcanal corridors to the IAC have rarely been studied. We sought to provide a step-by-step description of the transcanal transpromontorial approach to the IAC and analyze anatomic relationships that might enhance predictability and safety of this approach. METHODS: Ten cadaveric specimens were dissected and the extended transcanal transpromontorial approach to the IAC was established. Various morphometric measurements and anatomic landmarks were reviewed and analyzed. RESULTS: The proposed technique proved feasible and safe in all specimens. There was no inadvertent injury to the jugular bulb or internal carotid artery. The chorda tympani, a key landmark for the mastoid segment of the facial nerve, was identified in all dissections. The spherical recess of the vestibule and middle turn of cochlea are important landmarks for identification of the labyrinthine segment of the facial nerve. Identification of all boundaries of the working area is also essential for safe access. Among various morphometric measurements, the modiolus-IAC angle (≈ 150°) proved particularly consistent; given its ease of use and low variability, we believe it could serve as a landmark for identification and subsequent dissection of the IAC. CONCLUSIONS: The extended transcanal transpromontorial approach to the IAC is feasible and safe. Relying on anatomic landmarks to ensure preservation of the involved neurovascular structures is essential for a successful approach. The modiolus-IAC angle is a consistent, reproducible landmark for IAC identification and dissection.

Changes in paranasal sinus volumes, temporal bone pneumatization, internal acoustic canal and olfactory cleft dimensions over the centuries: a comparison of skulls from different epochs in Anatolia.

OBJECTIVES: Investigating changes in temporal bone pneumatization (TBP) and paranasal sinus volumes (PSV) across different eras may help understanding not only changes in skull anatomy but also pathophysiology of chronic otitis media and sinusitis, respectively, which are common health problems. METHODS: Eight skulls from the second century AD, 20 skulls were from the 10th-11th centuries AD, 20 skulls from the 16th-19th centuries AD, and 60 contemporary skulls were included in this cross-sectional observational study. Using computerized tomography (CT) scans, the PSV were calculated by multiplying the height, width, and antero-posterior distance of the sinuses. TBP was divided into three types. Internal acoustic canal (IAC) length and width, and olfactory cleft (OC) width were measured. RESULTS: No statistically significant differences were found between the paranasal sinus (frontal, maxillary, and sphenoid) volumes between the groups. However, TBP decreased statistically significantly over time on both sides of the skulls (p = 0.001). The contemporary IAC and OC measures were found to be significantly lower on both sides compared to the skulls from the other three eras (p < 0.001 for both). CONCLUSIONS: Although no significant change was observed in PSV, decreases were evident in TBP, OC width and IAC length and width over time. It appears a fair inference that changes in size of OC and IAC might be another indication of the fact that olfaction and hearing were more vital for survival in old eras. Since we do not know incidence of chronic ear problems in old eras, we cannot speculate outcome of increased TBP in terms of developing chronic ear diseases. On the contrary, increased TBP was likely to play a protective role in traumas in old ears. Additionally, the environmental influences may be crucial role in the development of paranasal sinuses.

New infant cranium from the African Miocene sheds light on ape evolution.

The evolutionary history of extant hominoids (humans and apes) remains poorly understood. The African fossil record during the crucial time period, the Miocene epoch, largely comprises isolated jaws and teeth, and little is known about ape cranial evolution. Here we report on the, to our knowledge, most complete fossil ape cranium yet described, recovered from the 13 million-year-old Middle Miocene site of Napudet, Kenya. The infant specimen, KNM-NP 59050, is assigned to a new species of Nyanzapithecus on the basis of its unerupted permanent teeth, visualized by synchrotron imaging. Its ear canal has a fully ossified tubular ectotympanic, a derived feature linking the species with crown catarrhines. Although it resembles some hylobatids in aspects of its morphology and dental development, it possesses no definitive hylobatid synapomorphies. The combined evidence suggests that nyanzapithecines were stem hominoids close to the origin of extant apes, and that hylobatid-like facial features evolved multiple times during catarrhine evolution.

Inner ear sensory system changes as extinct crocodylomorphs transitioned from land to water.

Major evolutionary transitions, in which animals develop new body plans and adapt to dramatically new habitats and lifestyles, have punctuated the history of life. The origin of cetaceans from land-living mammals is among the most famous of these events. Much earlier, during the Mesozoic Era, many reptile groups also moved from land to water, but these transitions are more poorly understood. We use computed tomography to study changes in the inner ear vestibular system, involved in sensing balance and equilibrium, as one of these groups, extinct crocodile relatives called thalattosuchians, transitioned from terrestrial ancestors into pelagic (open ocean) swimmers. We find that the morphology of the vestibular system corresponds to habitat, with pelagic thalattosuchians exhibiting a more compact labyrinth with wider semicircular canal diameters and an enlarged vestibule, reminiscent of modified and miniaturized labyrinths of other marine reptiles and cetaceans. Pelagic thalattosuchians with modified inner ears were the culmination of an evolutionary trend with a long semiaquatic phase, and their pelagic vestibular systems appeared after the first changes to the postcranial skeleton that enhanced their ability to swim. This is strikingly different from cetaceans, which miniaturized their labyrinths soon after entering the water, without a prolonged semiaquatic stage. Thus, thalattosuchians and cetaceans became secondarily aquatic in different ways and at different paces, showing that there are different routes for the same type of transition.

Fossil basicranium clarifies the origin of the avian central nervous system and inner ear.

Among terrestrial vertebrates, only crown birds (Neornithes) rival mammals in terms of relative brain size and behavioural complexity. Relatedly, the anatomy of the avian central nervous system and associated sensory structures, such as the vestibular system of the inner ear, are highly modified with respect to those of other extant reptile lineages. However, a dearth of three-dimensional Mesozoic fossils has limited our knowledge of the origins of the distinctive endocranial structures of crown birds. Traits such as an expanded, flexed brain, a ventral connection between the brain and spinal column, and a modified vestibular system have been regarded as exclusive to Neornithes. Here, we demonstrate all of these 'advanced' traits in an undistorted braincase from an Upper Cretaceous enantiornithine bonebed in southeastern Brazil. Our discovery suggests that these crown bird-like endocranial traits may have originated prior to the split between Enantiornithes and the more crownward portion of avian phylogeny over 140 Ma, while coexisting with a remarkably plesiomorphic cranial base and posterior palate region. Altogether, our results support the interpretation that the distinctive endocranial morphologies of crown birds and their Mesozoic relatives are affected by complex trade-offs between spatial constraints during development.

Neurosensory anatomy of Varanopidae and its implications for early synapsid evolution.

Varanopids are a group of Palaeozoic terrestrial amniotes which represent one of the earliest-diverging groups of synapsids, but their palaeoneurology has gone largely unstudied and recent analyses have challenged their traditional placement within synapsids. We utilized computed tomography (CT) to study the virtual cranial and otic endocasts of six varanopids, including representative taxa of both mycterosaurines and varanodontines. Our results show that the varanopid brain is largely plesiomorphic, being tubular in shape and showing no expansion of the cerebrum or olfactory bulbs, but is distinct in showing highly expanded floccular fossae. The housing of the varanopid bony labyrinth is also distinct, in that the labyrinth is bounded almost entirely by the supraoccipital-opisthotic complex, with the prootic only bordering the ventral portion of the vestibule. The bony labyrinth is surprisingly well-ossified, clearly preserving the elliptical, sub-orthogonal canals, prominent ampullae, and the short, undifferentiated vestibule; this high degree of ossification is similar to that seen in therapsid synapsids and supports the traditional placement of varanopids within Synapsida. The enlarged anterior canal, together with the elliptical, orthogonal canals and enlarged floccular fossa, lend support for the fast head movements indicated by the inferred predatory feeding mode of varanopids. Reconstructed neurosensory anatomy indicates that varanopids may have a much lower-frequency hearing range compared to more derived synapsids, suggesting that, despite gaining some active predatory features, varanopids retain plesiomorphic hearing capabilities. As a whole, our data reveal that the neuroanatomy of pelycosaur-grade synapsids is far more complex than previously anticipated.

Petrosal and bony labyrinth morphology of the stem paenungulate mammal (Paenungulatomorpha) Ocepeia daouiensis from the Paleocene of Morocco.

Based on high-resolution computed tomography, we describe in detail the petrosal and inner ear anatomy of one of the few known African stem paenungulates (Paenungulatomorpha), Ocepeia daouiensis from the Selandian of the Ouled Abdoun phosphate basin (Morocco). The petrosal of Ocepeia displays some remarkable, probably derived features (among eutherians) such as relatively small pars cochlearis, pars canalicularis labyrinth (including small semicircular canals), a large wing-like pars mastoidea, a large and inflated tegmen tympani, and the dorsoventral orientation of the large canal for the ramus superior. The presence of small semicircular canals in Ocepeia is an interesting shared trait with tenrecoidean afrotherians. Otherwise, and consistent with a general primitive skull morphology, the middle ear and labyrinth of Ocepeia daouiensis is characterised by many plesiomorphic traits close to the eutherian generalised plan. This adds to the rather generalised morphology of the earliest crown paenungulates such as Eritherium, Phosphatherium and Seggeurius to support an ancestral paenungulatomorph morphotype poorly derived from the eutherian pattern. As a result, Ocepeia provides key morphological and fossil data to test phylogenetic relationships of the Afrotheria (including Paenungulatomorpha) at the placental root mostly inferred from molecular studies.

Cochlear partition anatomy and motion in humans differ from the classic view of mammals.

Mammals detect sound through mechanosensitive cells of the cochlear organ of Corti that rest on the basilar membrane (BM). Motions of the BM and organ of Corti have been studied at the cochlear base in various laboratory animals, and the assumption has been that the cochleas of all mammals work similarly. In the classic view, the BM attaches to a stationary osseous spiral lamina (OSL), the tectorial membrane (TM) attaches to the limbus above the stationary OSL, and the BM is the major moving element, with a peak displacement near its center. Here, we measured the motion and studied the anatomy of the human cochlear partition (CP) at the cochlear base of fresh human cadaveric specimens. Unlike the classic view, we identified a soft-tissue structure between the BM and OSL in humans, which we name the CP "bridge." We measured CP transverse motion in humans and found that the OSL moved like a plate hinged near the modiolus, with motion increasing from the modiolus to the bridge. The bridge moved almost as much as the BM, with the maximum CP motion near the bridge-BM connection. BM motion accounts for 100% of CP volume displacement in the classic view, but accounts for only 27 to 43% in the base of humans. In humans, the TM-limbus attachment is above the moving bridge, not above a fixed structure. These results challenge long-held assumptions about cochlear mechanics in humans. In addition, animal apical anatomy (in SI Appendix) doesn't always fit the classic view.

A quick method to expose the structures and relations of the middle ear and inner ear by cadaveric dissection.

INTRODUCTION: The temporal bone contains structures related to hearing and balance, and is a valuable learning resource for medical students and trainee surgeons. The middle ear and inner ear are difficult to demonstrate by cadaveric dissection as the structures are closely contained in a small space in the dense temporal bone. Consequently, the teaching and learning of the ear are largely relegated to virtual and theoretical images, and models, which has resulted in a knowledge gap in medical students and prospective surgeons. The present study aimed to elucidate a technique that exposes the structures and relations of the middle and inner ear by cadaveric dissection. MATERIALS AND METHODS: Forty-seven adult formalin-fixed cadaveric specimens were dissected by the proposed technique. The method was evaluated based on the extent of the structures exposed and time taken for dissection. RESULTS: The method exposed all the contents and relations of the middle and inner ear, including the course of the facial nerve in the petrous temporal bone, in a few minutes, without use of specialized instruments like saw, drill, endoscope, operating microscope or electric trephine. CONCLUSION: This dissection method combines maximal exposure of the structures and relations of the middle and inner ear with a short dissection time, sans use of specialized tools. It can be incorporated in the gross anatomy curriculum for medical studentsdue to the short dissection time and completeness of structures exposed. The prosected specimen can also be plastinated for use as a teaching-learning resource for medical students and surgeons.

Exploring the ontogenetic development of the inner ear in Aardvarks.

The aardvark is the last living Tubulidentata, an order of afrotherian mammals. Afrotheria is supported strongly by molecular analyses, yet sparingly by morphological characters. Moreover, the biology of the aardvark remains incompletely known. The inner ear, and its ontogeny in particular, has not been studied in details yet, though it bears key ecomorphological characters and phylogenetical signal. The aim of this study is to decipher and discuss the ontogenetic development of the different areas of the inner ear of Orycteropus afer. We focused in particular on their relative size and morphological rates of development. Specimens were scanned with 3D imaging techniques. 3D and 2D geometric morphometrics coupled with qualitative descriptions of the petrosal ossification allowed us to evidence several stages through development. Based on our sample, the cochlea is the first structure of the inner ear to reach adult size, but it is the last one to acquire its adult morphology close to parturition. In contrast, after a delayed growth spurt, the semicircular canals reach their mature morphology before the cochlea, concomitantly with the increase of petrosal ossification. The ontogeny of the aardvark inner ear shows similarities with that of other species, but the apex of the cochlea presents some autapomorphies. This work constitutes a first step in the study of the ontogeny of this sensorial organ in Afrotheria.

Endocranial morphology of the piciformes (Aves, Coraciimorphae): Functional and ecological implications.

We used three-dimensional digital models to investigate the brain and endosseous labyrinth morphology of selected Neotropical Piciformes (Picidae, Ramphastidae, Galbulidae and Bucconidae). Remarkably, the brain morphology of Galbulidae clearly separates from species of other families. The eminentiae sagittales of Galbulidae and Bucconidae (insectivorous with high aerial maneuverability abilities) are smaller than those of the toucans (scansorial frugivores). Galbula showed the proportionally largest cerebellum, and Ramphastidae showed the least foliated one. Optic lobes ratio relative to the telencephalic hemispheres showed a strong phylogenetic signal. Three hypotheses were tested: (a) insectivorous taxa that need precise and fast movements to catch their prey, have well developed eminentiae sagittales compared to fruit eaters, (b) species that require high beak control would show larger cerebellum compared to other brain regions and higher number of visible folia and (c) there are marked differences between the brain shape of the four families studied here that bring valuable information of this interesting bird group. Hypotheses H1 and H2 are rejected, meanwhile H3 is accepted.

Relationship between lateral angle and shape of internal acoustic canal: cautionary note for diagnosis of sex.

Lateral angle (LA) of the internal acoustic canal (IAC) at its opening at the surface of the petrous portion of the temporal bone is known in literature as a dimorphic measurement useful for sex assessment. However, its reliability is still debated. Moreover, no information is available about the possible relationship between LA and shape and size of the IAC. This study aims at assessing breadth, length and lateral angle of IAC on 100 CT scans belonging to subjects aged between 20 and 70 years, equally divided between males and females. Possible differences in the three measurements according to side and sex were assessed through two-way ANCOVA test, using three cranial measurements (distance between anterior and posterior nasal spine, upper facial height, bizygomatic breadth) as covariates (p < 0.05). Possible correlations among IAC measurements and with age were assessed through Pearson's test (p < 0.05). Results showed a significantly greater IAC breadth on the left side than on the right side; moreover, LA was significantly wider in females than in males and was positively correlated with IAC breadth in both males and females. In addition, LA was negatively correlated with age only in the female group. The study first showed that LA is strictly related to IAC morphology; moreover, it is significantly affected by masculinization of skeletal traits in females. Therefore, caution is suggested in using this measurement for sex assessment.

Human bony labyrinth is an indicator of population history and dispersal from Africa.

The dispersal of modern humans from Africa is now well documented with genetic data that track population history, as well as gene flow between populations. Phenetic skeletal data, such as cranial and pelvic morphologies, also exhibit a dispersal-from-Africa signal, which, however, tends to be blurred by the effects of local adaptation and in vivo phenotypic plasticity, and that is often deteriorated by postmortem damage to skeletal remains. These complexities raise the question of which skeletal structures most effectively track neutral population history. The cavity system of the inner ear (the so-called bony labyrinth) is a good candidate structure for such analyses. It is already fully formed by birth, which minimizes postnatal phenotypic plasticity, and it is generally well preserved in archaeological samples. Here we use morphometric data of the bony labyrinth to show that it is a surprisingly good marker of the global dispersal of modern humans from Africa. Labyrinthine morphology tracks genetic distances and geography in accordance with an isolation-by-distance model with dispersal from Africa. Our data further indicate that the neutral-like pattern of variation is compatible with stabilizing selection on labyrinth morphology. Given the increasingly important role of the petrous bone for ancient DNA recovery from archaeological specimens, we encourage researchers to acquire 3D morphological data of the inner ear structures before any invasive sampling. Such data will constitute an important archive of phenotypic variation in present and past populations, and will permit individual-based genotype-phenotype comparisons.

"Reversed polarization" of Na/K-ATPase-a sign of inverted transport in the human endolymphatic sac: a super-resolution structured illumination microscopy (SR-SIM) study.

The human endolymphatic sac (ES) is believed to regulate inner ear fluid homeostasis and to be associated with Meniere's disease (MD). We analyzed the ion transport protein sodium/potassium-ATPase (Na/K-ATPase) and its isoforms in the human ES using super-resolution structured illumination microscopy (SR-SIM). Human vestibular aqueducts were collected during trans-labyrinthine vestibular schwannoma surgery after obtaining ethical permission. Antibodies against various isoforms of Na/K-ATPase and additional solute-transporting proteins, believed to be essential for ion and fluid transport, were used for immunohistochemistry. A population of epithelial cells of the human ES strongly expressed Na/K-ATPase α1, ß1, and ß3 subunit isoforms in either the lateral/basolateral or apical plasma membrane domains. The ß1 isoform was expressed in the lateral/basolateral plasma membranes in mostly large cylindrical cells, while ß3 and α1 both were expressed with "reversed polarity" in the apical cell membrane in lower epithelial cells. The heterogeneous expression of Na/K-ATPase subunits substantiates earlier notions that the ES is a dynamic structure where epithelial cells show inverted epithelial transport. Dual absorption and secretion processes may regulate and maintain inner ear fluid homeostasis. These findings may shed new light on the etiology of endolymphatic hydrops and MD.

The endocranium and trophic ecology of Velociraptor mongoliensis.

Neuroanatomical reconstructions of extinct animals have long been recognized as powerful proxies for palaeoecology, yet our understanding of the endocranial anatomy of dromaeosaur theropod dinosaurs is still incomplete. Here, we used X-ray computed microtomography (µCT) to reconstruct and describe the endocranial anatomy, including the endosseous labyrinth of the inner ear, of the small-bodied dromaeosaur, Velociraptor mongoliensis. The anatomy of the cranial endocast and ear were compared with non-avian theropods, modern birds, and other extant archosaurs to establish trends in agility, balance, and hearing thresholds in order to reconstruct the trophic ecology of the taxon. Our results indicate that V. mongoliensis could detect a wide and high range of sound frequencies (2,368-3,965 Hz), was agile, and could likely track prey items with ease. When viewed in conjunction with fossils that suggest scavenging-like behaviours in V. mongoliensis, a complex trophic ecology that mirrors modern predators becomes apparent. These data suggest that V. mongoliensis was an active predator that would likely scavenge depending on the age and health of the individual or during prolonged climatic events such as droughts.

Virtual endocranial and inner ear endocasts of the Paleocene 'condylarth' Chriacus: new insight into the neurosensory system and evolution of early placental mammals.

The end-Cretaceous mass extinction allowed placental mammals to diversify ecologically and taxonomically as they filled ecological niches once occupied by non-avian dinosaurs and more basal mammals. Little is known, however, about how the neurosensory systems of mammals changed after the extinction, and what role these systems played in mammalian diversification. We here use high-resolution computed tomography (CT) scanning to describe the endocranial and inner ear endocasts of two species, Chriacus pelvidens and Chriacus baldwini, which belong to a cluster of 'archaic' placental mammals called 'arctocyonid condylarths' that thrived during the ca. 10 million years after the extinction (the Paleocene Epoch), but whose relationships to extant placentals are poorly understood. The endocasts provide new insight into the paleobiology of the long-mysterious 'arctocyonids', and suggest that Chriacus was an animal with an encephalization quotient (EQ) range of 0.12-0.41, which probably relied more on its sense of smell than vision, because the olfactory bulbs are proportionally large but the neocortex and petrosal lobules are less developed. Agility scores, estimated from the dimensions of the semicircular canals of the inner ear, indicate that Chriacus was slow to moderately agile, and its hearing capabilities, estimated from cochlear dimensions, suggest similarities with the extant aardvark. Chriacus shares many brain features with other Paleocene mammals, such as a small lissencephalic brain, large olfactory bulbs and small petrosal lobules, which are likely plesiomorphic for Placentalia. The inner ear of Chriacus also shares derived characteristics of the elliptical and spherical recesses with extinct species that belong to Euungulata, the extant placental group that includes artiodactyls and perissodactyls. This lends key evidence to the hypothesized close relationship between Chriacus and the extant ungulate groups, and demonstrates that neurosensory features can provide important insight into both the paleobiology and relationships of early placental mammals.

Functional anatomy of the middle and inner ears of the red fox, in comparison to domestic dogs and cats.

Anatomical middle and inner ear parameters are often used to predict hearing sensitivities of mammalian species. Given that ear morphology is substantially affected both by phylogeny and body size, it is interesting to consider whether the relatively small anatomical differences expected in related species of similar size have a noticeable impact on hearing. We present a detailed anatomical description of the middle and inner ears of the red fox Vulpes vulpes, a widespread, wild carnivore for which a behavioural audiogram is available. We compare fox ears to those of the well-studied and similarly sized domestic dog and cat, taking data for dogs and cats from the literature as well as providing new measurements of basilar membrane (BM) length and hair cell numbers and densities in these animals. Our results show that the middle ear of the red fox is very similar to that of dogs. The most obvious difference from that of the cat is the lack of a fully formed bony septum in the bulla tympanica of the fox. The cochlear structures of the fox, however, are very like those of the cat, whereas dogs have a broader BM in the basal cochlea. We further report that the mass of the middle ear ossicles and the bulla volume increase with age in foxes. Overall, the ear structures of foxes, dogs and cats are anatomically very similar, and their behavioural audiograms overlap. However, the results of several published models and correlations that use middle and inner ear measurements to predict aspects of hearing were not always found to match well with audiogram data, especially when it came to the sharper tuning in the fox audiogram. This highlights that, although there is evidently a broad correspondence between structure and function, it is not always possible to draw direct links when considering more subtle differences between related species.