Effects of age and size in the ears of gekkotan lizards: auditory sensitivity, its determinants, and new insights into tetrapod middle-ear function

Do related, differently sized species differ in size-related structural or functional traits merely because they mature at different points of a uniform allometric ontogenetic growth curve, or do they evolutionarily diverge? We tested ears of gekkotan lizards through experiments distinguishing the two. Auditory sensitivity was assessed by compound action potential (CAP) thresholds in eight species. The best thresholds characterizing species ranged 22–72 dB sound pressure level at 0.5–1.0 kHz. Direct acoustic stimulation of the columella footplate elevated thresholds by 25–50 dB. Intraspecific CAP sensitivity was primarily affected by body length in Eublepharis macularius, but by tympanic-membrane velocity in Oedura marmorata. The chief factor determining middle-ear function (difference in CAP sensitivity before and after middle-ear ablation) was body length in both species. A secondary factor was the middle-ear hydraulic lever ratio in E. macularius, but the mechanical lever in O. marmorata. When intra- and interspecific data were compared, the relation of CAP thresholds to body size in E. macularius resembled the interspecific regression. The intraspecific regression of auditory sensitivity over tympanic membrane velocity in O. marmorata differed from that calculated interspecifically. Hence, the evolutionary contribution to size effects on CAP sensitivity exceeds the ontogenetic contribution. Putatively, body length affects CAP sensitivity through absolute sizes of tympanic membrane and columella footplate. These newly discovered effects join those of the hydraulic lever and (interspecifically) hair-cell number to improve the hearing of larger species that vocally communicate across wider spaces, apparently throughout the Tetrapoda.     

Finite element analysis of the effects of a floating mass transducer on the performance of a middle ear implant

Experimental evidence has shown that floating mass transducers (FMTs) play a key role in the performance of middle ear implants. However, because of the tiny size and complex structure of the middle ear, systematic experimental study of the influences of FMTs is difficult to carry out. In this paper we develop a FMT-attached middle-ear finite element model to investigate some effects of a FMT on the performance of a middle ear implant. This model was constructed based on a complete set of computerized tomography section images of a healthy volunteer's left ear. The validity of the developed model was verified by comparing the model-predicted motion of the tympanic membrane and stapes footplate with published experimental data. The result shows that the FMT produces a mass loading effect prominently at high frequencies, the force required to drive the incus to the equivalent of 100 dB sound pressure level (SPL) is about 89?μN, and setting the attachment position of the FMT close to the incudostapedial joint can enhance the driving effect.     

Limb bone allometry during postnatal ontogeny in non-avian dinosaurs

Although the interspecific scaling of tetrapods is well understood, remarkably little work has been done on the ontogenetic scaling within tetrapod species, whether fossil or recent. Here the ontogenetic allometry of the femur, humerus, and tibia was determined for 23 species of non‐avian dinosaur by regressing log‐transformed length against log‐transformed circumference for each bone using reduced major axis bivariate regression. The femora of large theropod species became more robust during ontogeny, whereas growth in the femora of sauropodomorphs and most ornithischians was not significantly different from isometry. Hadrosaur hindlimb elements became significantly more gracile during ontogeny. Scaling constants were higher in all theropods than in any non‐theropod taxa. Such clear taxonomically correlated divisions were not evident in the ontogenetic allometry of the tibia and hindlimb bones did not scale uniformly within larger taxonomic groups. For taxa in which the ontogenetic allometry of the humerus was studied, only Riojasaurus incertus exhibited a significant departure from isometry. Using independent contrasts, the regression of femoral allometry against the log of adult body mass was found to have a significant negative correlation but such a relationship could not be established for other limb elements or growth parameters, mainly due to the small sample size. The intraspecific scaling patterns observed in dinosaurs and other amniotes do not support earlier hypotheses that intraspecific scaling differs between endothermic and ectothermic taxa.     

Developmental Osteology of the Parafrontal Bones of the Sphaerodactylidae

Well‐resolved phylogenetic hypotheses and ontogenetic data are often necessary for investigating the evolution of structural novelty. The Sphaerodactylidae comprises 12 genera of predominantly miniaturized geckos. The genera Aristelliger and Teratoscincus are exceptions, with taxa reaching snout‐to‐vent lengths far exceeding those of other sphaerodactylids. These two genera possess enigmatic, supraorbital ossifications—parafrontal bones—which are encountered nowhere else among squamates. At the time of their discovery, these structures were believed to be the result of evolutionary convergence. Although relationships between other sphaerodactylids remain unresolved, recent molecular and morphological data have supported a close relationship between Aristelliger and Teratoscincus. We investigated the ontogeny of parafrontal bones to better understand relationships between sphaerodactylid body size and the presence of parafrontals, and to evaluate whether ontogenetic data support the homology of parafrontals between Aristelliger and Teratoscincus. We hypothesize that the parafrontals of Aristelliger and Teratoscincus are homologous and that there is a threshold body size in sphaerodactylids below which parafrontals do not develop, thus explaining their absence from the miniaturized taxa. The presence of parafrontals was investigated in all sphaerodactylid genera using cleared‐and‐stained, radiographed, and skeletonized specimens. Total surface area of parafrontals was measured for seven species of Aristelliger and six species of Teratoscincus throughout their ontogeny. Histology was used to investigate the cellular composition of the parafrontals throughout their ontogeny. Our data suggest that parafrontals have evolved in parallel from a homologous, parafrontal precursor and that the onset of parafrontal development is not strictly dependent on a threshold body size. Anat Rec, 301:581–606, 2018. © 2017 Wiley Periodicals, Inc.     

Three-Dimensional Finite Element Modeling of Human Ear for Sound Transmission

An accurate, comprehensive finite element model of the human ear can provide better understanding of sound transmission, and can be used for assessing the influence of diseases on hearing and the treatment of hearing loss. In this study, we proposed a three-dimensional finite element model of the human ear that included the external ear canal, tympanic membrane (eardrum), ossicular bones, middle ear suspensory ligaments/muscles, and middle ear cavity. This model was constructed based on a complete set of histological section images of a left ear temporal bone. The finite element (FE) model of the human ear was validated by comparing model-predicted ossicular movements at the stapes footplate and tympanic membrane with published experimental measurements on human temporal bones. The FE model was employed to predict the effects of eardrum thickness and stiffness, incudostapedial joint material, and cochlear load on acoustic-mechanical transmission through the human ossicular chain. The acoustic-structural coupled FE analysis between the ear canal air column and middle ear ossicles was also conducted and the results revealed that the peak responses of both tympanic membrane and stapes footplate occurred between 3000 and 4000 Hz.     

Allometric relations of cardiovascular function in birds

The avian cardiovascular system is interesting because birds have larger hearts and lower heart rates than mammals of the same size. This study defines how cardiovascular function is related to body size in nine species of birds encompassing a 1,000-fold size range. Heart mass (HM, g) was related to body mass (M, kg) by the following equation: HM = 8.76 M0.92, and the heart rate (HR, beats X min-1) to body mass by HR = 178.5 M-0.282. Cardiac output (Q, ml X min-1) was related to body mass as Q = 290.7 M0.69. Birds also had a greater cardiac output for a given body mass than mammals. Cardiac output was directly proportional to the rate of O2 consumption (VO2, ml X min-1): Q = 17.5 VO2(1.04), with birds having a greater cardiac output for a given VO2 than mammals. (Variables are considered directly proportional if the exponent does not differ significantly from one). As in mammals, stroke volume (SV, ml) was also directly proportional to body mass: SV = 1.72 M0.97, but for a given body size stroke volume was larger in birds than in mammals. However, as bird hearts are disproportionately large in comparison with body size, when stroke volume is expressed per gram of heart (SV = 0.177 HM1.05) birds have a significantly lower stroke volume-to-heart weight ratio than mammals.(ABSTRACT TRUNCATED AT 250 WORDS)     

Aquatic adaptations in the nose of carnivorans: evidence from the turbinates

Inside the mammalian nose lies a labyrinth of bony plates covered in epithelium collectively known as turbinates. Respiratory turbinates lie anteriorly and aid in heat and water conservation, while more posterior olfactory turbinates function in olfaction. Previous observations on a few carnivorans revealed that aquatic species have relatively large, complex respiratory turbinates and greatly reduced olfactory turbinates compared with terrestrial species. Body heat is lost more quickly in water than air and increased respiratory surface area likely evolved to minimize heat loss. At the same time, olfactory surface area probably diminished due to a decreased reliance on olfaction when foraging under water. To explore how widespread these adaptations are, we documented scaling of respiratory and olfactory turbinate surface area with body size in a variety of terrestrial, freshwater, and marine carnivorans, including pinnipeds, mustelids, ursids, and procyonids. Surface areas were estimated from high-resolution CT scans of dry skulls, a novel approach that enabled a greater sampling of taxa than is practical with fresh heads. Total turbinate, respiratory, and olfactory surface areas correlate well with body size (r(2) ≥0.7), and are relatively smaller in larger species. Relative to body mass or skull length, aquatic species have significantly less olfactory surface area than terrestrial species. Furthermore, the ratio of olfactory to respiratory surface area is associated with habitat. Using phylogenetic comparative methods, we found strong support for convergence on 1:3 proportions in aquatic taxa and near the inverse in terrestrial taxa, indicating that aquatic mustelids and pinnipeds independently acquired similar proportions of olfactory to respiratory turbinates. Constraints on turbinate surface area in the nasal chamber may result in a trade-off between respiratory and olfactory function in aquatic mammals.     

Comparative analysis of the shape and size of the middle ear cavity of turtles reveals no correlation with habitat ecology

The middle ear of turtles differs from other reptiles in being separated into two distinct compartments. Several ideas have been proposed as to why the middle ear is compartmentalized in turtles, most suggesting a relationship with underwater hearing. Extant turtle species span fully marine to strictly terrestrial habitats, and ecomorphological hypotheses of turtle hearing predict that this should correlate with variation in the structure of the middle ear due to differences in the fluid properties of water and air. We investigate the shape and size of the air‐filled middle ear cavity of 56 extant turtles using 3D data and phylogenetic comparative analysis to test for correlations between habitat preferences and the shape and size of the middle ear cavity. Only weak correlations are found between middle ear cavity size and ecology, with aquatic taxa having proportionally smaller cavity volumes. The middle ear cavity of turtles exhibits high shape diversity among species, but we found no relationship between this shape variation and ecology. Surprisingly, the estimated acoustic transformer ratio, a key functional parameter of impedance‐matching ears in vertebrates, also shows no relation to habitat preferences (aquatic/terrestrial) in turtles. We suggest that middle ear cavity shape may be controlled by factors unrelated to hearing, such as the spatial demands of surrounding cranial structures. A review of the fossil record suggests that the modern turtle ear evolved during the Early to Middle Jurassic in stem turtles broadly adapted to freshwater and terrestrial settings. This, combined with our finding that evolutionary transitions between habitats caused only weak evolutionary changes in middle ear structure, suggests that tympanic hearing in turtles evolved as a compromise between subaerial and underwater hearing.     

人耳鼓膜病变数值分析

目的研究鼓膜厚度和硬度对人耳传声的影响。方法利用CT获取志愿者耳部结构临床资料,使用Matlab软件提取相关结构的边界,将边界文件导入ANSYS建立人耳结构数值有限元模型。结果利用本文人耳数值模型,在外耳道口施加105dB声压,进行200～8000Hz频率范围的谐响应分析。以此研究在鼓膜病变情况下,鼓膜和镫骨底板位移幅值的变化规律。结论用数值方法解释了鼓膜病变对传声的影响,为鼓膜修补提供了力学参考。     

Ontogeny of Orbit Orientation in Primates

Orbit orientation in primates has been linked to adaptive factors related to activity pattern and size-related variation in structural influences on orbit position. Although differences in circumorbital form between anthropoids and strepsirrhines appear to be related to interspecific disparities in levels of orbital convergence and orbital frontation, there is considerable overlap in convergence between suborders. Unfortunately, putative links between convergence and frontation across primates, and consequent arguments about primate and anthropoid origins, are likely to be influenced by allometry, the size range of a respective sample, and adaptive influences on encephalization and activity patterns. Such a multifarious system is less amenable to interspecific treatment across higher-level clades. An ontogenetic perspective is one way to evaluate transformations from one character state to another, especially as they pertain to allometric effects on phenotypic variation. We characterized the ontogeny of orbital convergence and frontation in 13 anthropoid and strepsirrhine species. In each suborder, correlation and regression analyses were used to test hypotheses regarding the allometric bases of variation in orbital orientation. Growth trajectories were analyzed intra- and inter-specifically. Frontation decreased postnatally in all taxa due to the negative scaling of brain vs. skull size. Further, interspecific variation in relative levels of frontation was linked to corresponding ontogenetic transpositions in encephalization that differed within both suborders. In strepsirrhines, postnatal increases in convergence were largely due to the negative allometry of orbit vs. skull size. In contrast, convergence in anthropoids varied little during growth, being unrelated to ontogenetic variation in either relative orbit or interorbit size. Anat Rec, 302:2093–2104, 2019. © 2019 American Association for Anatomy     

Allometric growth in mass by the brain of mammals

I re-examined published data for ontogenetic change in relative mass of the brain in six species of mammal (i.e., sheep, pig, cow, horse, rat, cat) to illustrate an insidious problem with conventional analyses of brain–body allometry. Graphical displays of logarithmic transformations of the original data for each species give the appearance of two discrete mathematical distributions, but untransformed observations nonetheless conform to a single distribution that is well described by a single, nonlinear equation. The concept of biphasic, allometric growth by the brain consequently is an artifact of transformation. The notion of Rapid and Slow phases in relative growth by the brain also is an artifact, because the notion is based explicitly on the concept of biphasic growth allometry. Relative growth by the brain in sheep, pigs, cows, and horses follows the path of a power curve with an exponent less than 1, so relative growth declines progressively as animals grow to their maximum size, at which point growth effectively ends for both brain and body. Relative growth by the brain in rats and cats follows the path of an exponential curve and consequently is more like relative growth by the brain of odontocoete cetaceans and primates, with the brain growing rapidly relative to the body early in ontogeny and attaining maximum (cats) or near-maximum (rats) mass well before the body reaches its maximum. An exponential pattern of relative growth by the brain appears to have evolved independently in rodents, carnivores, odontocoetes, and primates.     

Petrosal anatomy and inner ear structures of the Late Jurassic Henkelotherium (Mammalia, Cladotheria, Dryolestoidea): insight into the early evolution of the ear region in cladotherian mammals

The petrosal anatomy and inner ear structure of Jurassic cladotherian mammals represent the ancestral morphological conditions (groundplan) from which modern therian mammals (marsupials and placentals) have evolved. We present the reconstruction of the petrosal and inner ear features of the Late Jurassic dryolestoid mammal Henkelotherium guimarotae from high-resolution computed tomography and three-dimensional imaging analysis. This study of Henkelotherium revealed a combination of derived and primitive features, including: cladotherian apomorphies, such as the promontorial sulcus for the internal carotid artery and reduced lateral trough; trechnotherian characters, such as an enclosed cochlear canaliculus for the perilymphatic duct, post-promontorial tympanic sinus and caudal tympanic process; in addition to plesiomorphic mammalian features, such as the cavum supracochleare and prootic canal. The inner ear of Henkelotherium shows a division between the utricle and saccule, a cochlear canal coiled through at least 270°, a distinctive primary bony lamina for the basilar membrane, and a secondary bony lamina. The development of the primary and secondary bony laminae in the cochlear canal is suggested here to be correlated with the concurrent coiling of the bony canal and membranous duct of the inner ear cochlea, apomorphies of the more inclusive cladotherian clade that also represent the ancestral morphotype of modern therian mammals. Because these features are crucial for high-frequency hearing in extant therian mammals, their early appearance in Late Jurassic cladotherians suggests a more ancient origination for high-frequency hearing in mammalian history than previously thought.     

Regeneration of rat auditory ossicles using recombinant human BMP-2/collagen composites

Tympanoplasty operations to improve hearing impairment require a wide middle ear cavity and reconstruction of columellar formations. There is no specific material for use in the reconstruction of columellar formations. Tissue response to BMP has been employed as regenerative material. To our knowledge, however, there are no reports of the reconstruction of columellar formations using recombinant human bone morphogenetic protein-2/bovine collagen composites in the middle ear. The purpose of this study is to investigate whether recombinant human bone morphogenetic protein-2/bovine collagen composites (rhBMP-2 composites) are appropriate for use as regenerative material for tympanoplasty. In the form of pellets, rhBMP-2 composites were implanted as columellae into the tympanic cavity. At 2, 4, 6, 8, and 12 weeks after surgery, the middle ear of the animals (n = 4 at each week) was stained with hematoxylin-eosin for light microscopic observation. All composites were in the process of ossification or had ossified according to their developmental stages and were covered with a single layer of squamous or cuboidal epithelium. The new bone formed in these composites was persistently stable and displayed some columellar conditions assessed by histological examination. This study led to the conclusion that rhBMP-2 composites make excellent regenerative material for auditory ossicles.     

Scaling and Accommodation of Jaw Adductor Muscles in Canidae

The masticatory apparatus amongst closely related carnivoran species raises intriguing questions about the interplay between allometry, function, and phylogeny in defining interspecific variations of cranial morphology. Here we describe the gross structure of the jaw adductor muscles of several species of canid, and then examine how the muscles are scaled across the range of body sizes, phylogenies, and trophic groups. We also consider how the muscles are accommodated on the skull, and how this is influenced by differences of endocranial size. Data were collected for a suite of morphological metrics, including body mass, endocranial volume, and muscle masses and we used geometric morphometric shape analysis to reveal associated form changes. We find that all jaw adductor muscles scale isometrically against body mass, regardless of phylogeny or trophic group, but that endocranial volume scales with negative allometry against body mass. These findings suggest that head shape is partly influenced by the need to house isometrically scaling muscles on a neurocranium scaling with negative allometry. Principal component analysis suggests that skull shape changes, such as the relatively wide zygomatic arches and large sagittal crests seen in species with higher body masses, allow the skull to accommodate a relative enlargement of the jaw adductors compared with the endocranium. Anat Rec, 299:951–966, 2016. © 2016 The Authors The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology Published by Wiley Periodicals, Inc.     

Comparative analysis of vestibular ecomorphology in birds

The bony labyrinth of vertebrates houses the semicircular canals. These sense rotational accelerations of the head and play an essential role in gaze stabilisation during locomotion. The sizes and shapes of the semicircular canals have hypothesised relationships to agility and locomotory modes in many groups, including birds, and a burgeoning palaeontological literature seeks to make ecological interpretations from the morphology of the labyrinth in extinct species. Rigorous tests of form–function relationships for the vestibular system are required to support these interpretations. We test the hypothesis that the lengths, streamlines and angles between the semicircular canals are related to body size, wing kinematics and flying style in birds. To do this, we applied geometric morphometrics and multivariate phylogenetic comparative methods to a dataset of 64 three‐dimensional reconstructions of the endosseous labyrinth obtained using micro‐computed tomography scanning of bird crania. A strong relationship between centroid size of the semicircular canals and body size indicates that larger birds have longer semicircular canals compared with their evolutionary relatives. Wing kinematics related to manoeuvrability (and quantified using the brachial index) explain a small additional portion of the variance in labyrinth size. We also find strong evidence for allometric shape change in the semicircular canals of birds, indicating that major aspects of the shape of the avian labyrinth are determined by spatial constraints. The avian braincase accommodates a large brain, a large eye and large semicircular canals compared with other tetrapods. Negative allometry of these structures means that the restriction of space within the braincase is intense in small birds. This may explain our observation that the angles between planes of the semicircular canals of birds deviate more strongly from orthogonality than those of mammals, and especially from agile, gliding and flying mammals. Furthermore, we find little support for relationships between labyrinth shape and flying style or wing kinematics. Overall, our results suggest that the topological problem of fitting long semicircular canals into a spatially constrained braincase is more important in determining the shape of the avian labyrinth than the specifics of locomotory style or agility. Our results tentatively indicate a link between visual acuity and proportional size of the labyrinth among birds. This suggests that the large labyrinths of birds compared with other tetrapods may result from their generally high visual acuities, and not directly from their ability to fly. The endosseous labyrinths of extinct birds and their close dinosaurian relatives may allow broad inferences about flight or vision, but so far provide few specific insights into detailed aspects of locomotion.     

人工听骨不同接入方式对耳结构动力响应的影响

目的 研究人工听骨接入方式对听力恢复的影响。方法 通过CT扫描技术,结合自编C＋＋程序读取CT数据中体单元建立人耳结构几何模型,将几何模型导入PATRAN中建立有限元数值模型。采用频率响应方法对耳结构模型进行流固耦合计算,同时分析人工听骨不同接入方式及接入位置对耳结构声音传导的影响。结果 通过对正常人耳的动力响应分析,得到数值模型中计算出的鼓膜凸与镫骨底板振幅与试验数据吻合,验证本文模型的正确性。结论接在鼓膜凸的位置其动力响应最好,镫骨振幅高于其他连接方式。即人工听骨接在鼓膜凸的位置比较吻合人耳的生理功能,其重建听力效果更好。     

A Novel Understanding of Phocidae Hearing Adaptations Through a Study of Northern Elephant Seal (Mirounga angustirostris) Ear Anatomy and Histology

The most conspicuous aural adaptation in northern elephant seals (NES) is complete absence of an auricle and a tortuous collapsed external acoustic meatus. The NES epitympanic recess contains massive ossicles immersed in the middle ear cavernous sinuses. Engorgement of the cavernous sinuses would make ossicles fully buoyant during deep diving. NES have a comparatively larger cochlear nerve, which carries a significantly larger number of axons than in terrestrial mammals, which would give them auditory ability similar to the obligate marine mammals such as cetaceans. Our calculations show that the traditional “air-dependent” impedance matching mechanism in NES functions to just half of the capacity compared with the one described in terrestrial mammals. Impedance matching would be further hindered in NES while diving due to fully collapsed external acoustic meatus. Thanks to similarities of acoustic impedance between the sea water, soft tissues, and blood sinuses, very little sound energy would be reflected and lost. When sound is generated underwater, the large ossicles, buoyant in the cavernous sinus, would not move due to oscillation of tympanic membrane. Rather, they would be oscillating due to their inertia and process of acoustic streaming. Our mathematical simulation shows that an increase in sound frequency would cause increased displacement of the stapedial footplate and thus transmit the sound energy to the inner ear. We contend that during diving, impedance matching and sound signal amplification in the middle ear courses through the cavernous sinuses and oscillates the enlarged ossicles, thus enabling a high-frequency ultrasonic hearing range in Phocidae. Anat Rec, 302:1605–1614, 2019. © 2018 American Association for Anatomy     

Effect of anterior tympanomeatal angle blunting on the middle ear transfer function using a finite element ear model

The anterior tympanomeatal angle (ATA) blunting is clinically defined as a certain degree of the ATA obliteration due to excessive fibrous tissue formation, which is a relatively common complication of external auditory canal (EAC) related operations. The aim of this study was to examine the effect of ATA blunting on the middle ear transfer function using a finite element (FE) model. Results showed that the displacements at the tympanic membrane (TM), at the manubrium and at the stapes footplate, and also the ratio of stapes footplate velocity to the sound pressure in the EAC were decreased to various degrees from ATA blunting of Grades 1–4. This was more significant with TM thickening at the frequencies below 3.2 kHz, particularly in Grades 3 and 4 when analyzing the anterior region of the TM. The phase differences of TM and stapes footplate increased with the ATA blunting from Grades 1 to 4 in relation to normal ATA. It is noteworthy that the vibration mode of the malleus does not show obvious change, compared to the displacement reduction at the TM with ATA blunting Grades 1–4. These results suggest that FE analysis of ATA blunting effect appears to be effective.     

The morphological diversity of the quadrate bone in squamate reptiles as revealed by high-resolution computed tomography and geometric morphometrics

We examined the morphological diversity of the quadrate bone in squamate reptiles (i.e. lizards, snakes, amphisbaenians). The quadrate is the principal splanchnocranial element involved in suspending the lower jaw from the skull, and its shape is of particular interest because it is potentially affected by several factors, such as phylogenetic history, allometry, ecology, skull kinesis and hearing capabilities (e.g. presence or absence of a tympanic ear). Due to its complexity, the quadrate bone is also considered one of the most diagnostic elements in fragmentary fossil taxa. We describe quadrates from 38 species spread across all major squamate clades, using qualitative and quantitative (e.g. geometric morphometrics) methods. We test for possible correlations between shape variation and factors such as phylogeny, size, ecology and presence/absence of a tympanum. Our results show that the shape of the quadrate is highly evolutionarily plastic, with very little of the diversity explained by phylogenetic history. Size variation (allometric scaling) is similarly unable to explain much shape diversity in the squamate quadrate. Ecology (terrestrial/fossorial/aquatic) and presence of a tympanic ear are more significant, but together explain only about 20% of the diversity observed. Other unexplored and more analytically complex factors, such as skull biomechanics, likely play additional major roles in shaping the quadrates of lizards and snakes.