Comparative quantitative investigations on brains of wild cavies (Cavia aperea) and guinea pigs (Cavia aperea f. porcellus). A contribution to size changes of CNS structures due to domestication

Intraspecific allometric calculations of the brain to body size relation revealed distinct differences between 127 (67; 60) ancestral wild cavies and 82 (37; 45) guinea pigs, their domesticated relatives. The dependency of both measures from one another remained the same in both animal groups but the brains of guinea pigs were by 14.22% smaller at any net body weight. Consistent with results in other species the domestication of Cavia aperea is also characterized by a decrease of brain size. Fresh tissue sizes of the five brain parts medulla oblongata, cerebellum, mesencephalon, diencephalon and telencephalon were determined for 6 cavies and 6 guinea pigs by the serial section method. Additionally the sizes of 16 endbrain structures and those of the optic tract, the lateral geniculate body and the cochlear nucleus were measured. Different decrease values resulted for all these structures concomitant with domestication as was calculated from the amount of total brain size decrease and average relative structure values in the wild as well as the domesticated brain. The size decrease of the entire telencephalon (−13.7%) was within the range of the mean overall reduction as similarly was the case for the total neocortex (−10.7%) whereas the total allocortex (−20.9%) clearly was more strongly affected. The size decrease of the olfactory bulb (−41.9%) was extreme and clearly higher than found for the secondary olfactory structures (around −11%). The primary nuclei of other sensory systems (vision, audition) were decreased to less extent (lateral geniculate: −18.1%; cochlear nucleus: −12.6%). Mass decreases of pure white matter parts were nearly twice as high in contrast to associated grey matter parts (neocortex white versus grey matter; tractus opticus versus lateral geniculate body). The relatively great decrease values found for the limbic structures hippocampus (−26.9%) and schizocortex (−25.9%) are especially notable since they are in good conformity with domestication effects in other mammalian species. The findings of this study are discussed with regard to results of similar investigations on wild and domesticated gerbils (Meriones unguiculatus), the encephalization of the wild form, the special and species-specific mode and duration of domestication and in connection with certain behavioral changes as resulted from comparative investigations in ethology, socio-biology, endocrinology and general physiology.     

The behavioral endocrinology of domestication: A comparison between the domestic guinea pig (Cavia aperea f. porcellus) and its wild ancestor, the cavy (Cavia aperea)

In this study spontaneous behavior and endocrine parameters were compared between the domestic guinea pig (Cavia aperea f. porcellus) and its wild ancestor, the cavy (Cavia aperea), to elucidate the process of domestication in this species. In 120 h of observation time the behavior of five groups of wild and seven groups of domestic guinea pigs, each consisting of one adult male and two adult females, was analyzed quantitatively. To assess the activities of the pituitary-adrenocortical (PAC), the pituitary-gonadal (PG), and the sympathetic-adrenomedullary (SAM) systems, serum cortisol, testosterone, epinephrine, and norepinephrine concentrations, as well as adrenal tyrosine hydroxylase activities, were determined in males of both forms. The following significant differences between wild cavies and domestic guinea pigs were found: the domesticated animals displayed less aggressive but more sociopositive and more male courtship behavior than their wild ancestors. In addition, they were distinctly less attentive to their physical environment than the wild cavies. The basal activity of the SAM system, as well as the reactivity of the SAM and the PAC systems, was distinctly reduced in the domesticated animals. In contrast, the basal activity of the PAC system did not differ between both forms. The activity of the PG system was significantly higher in males of the domestic guinea pig than in male wild cavies. Thus, in guinea pigs the process of domestication has led to typical behavioral traits-reduced aggressiveness, increased social tolerance-which have also been found in comparisons between wild and domestic forms of other species. The decreased reactivity of the organism's stress axes can be regarded as a physiological mechanism which helps domesticated animals to adjust to man-made housing conditions.     

Changes in size of Baltic field voles over the last 50 years: are they really shrinking?

Using museum materials and recently trapped specimens of field voles (Microtus agrestis (Linnaeus, 1761)) from Lithuania and Estonia, we assessed temporal and latitudinal trends in body and skull size, comparing the periods 1980–1996 and 2014–2016. We measured four body and 23 skull characters, size-adjusting them using the geometric mean procedure. A pronounced decrease in the size of M. agrestis was noted in Estonia, where 23 out of 27 adjusted body and skull characters had decreased by up to 21.9%, with only the tail length, hind foot length, maximum height of mandibula excluding coronoid process and coronoid height of mandibula increasing significantly. Decreases were less marked in voles from Lithuania – most pronounced were a 6.1% decrease in adjusted body length, an 11.6% decrease in adjusted length of the braincase, a 3.85% decrease in the breadth of the braincase, measured at the widest part, a 2.9% decrease in condylobasal skull length and a 2.2% decrease in the height of the braincase. The coronoid height of the mandibula of Lithuanian individuals showed an 8.4% size increase. In both countries, the confounding effect of sex on the size changes of M. agrestis from 1980 to 2016 was much smaller than the effect of time period. Concluding, voles in Estonia became significantly smaller, while changes in the measured characters in Lithuania were heterogeneous.     

Miniaturization and its effects on cranial morphology in plethodontid salamanders, genus Thorius (Amphibia, Plethodontidae): II. The fate of the brain and sense organs and their role in skull morphogenesis and evolution

Relative size and arrangement of the brain and paired sense organs are examined in three species of Thorius, a genus of minute, terrestrial salamanders that are among the smallest extant tailed tetrapods. Analogous measurements of representative species of three related genera of larger tropical (Pseudoeurycea, Chiropterotriton) and temperate (Plethodon) salamanders are used to identify changes in gross morphology of the brain and sense organs that have accompanied the evolution of decreased head size in Thorius and their relation to associated changes in skull morphology. In adult Thorius, relative size (area measured in frontal plane, and length) of the eyes, otic capsules, and brain each is greater than in adults of all of the larger genera; relative size of the nasal capsules is unchanged or slightly smaller. Interspecific scaling phenomena--negative allometry of otic capsule, eye and brain size, isometry or slight positive allometry of nasal capsule size, all with respect to skull length--also are characteristic of intraspecific (ontogenetic) comparisons in both T. narisovalis and Pseudoeurycea goebeli. Predominance of the brain and eyes in Thorius results in greater contact and overlap among these structures and the nasal capsules in the anterior portion of the head. This is associated with anterior displacement of both the eyes and nasal capsules, which now protrude anterior to the skull proper; a change in eye shape; and medial deformation of anterior braincase walls. Posteriorly, predominance of the otic capsules has effected a reorientation of the jaw suspensorium to a fully vertical position that is correlated with the novel presence of a posteriorly directed squamosal process and shift in origin of the quadropectoralis muscle. Many of these changes in cranial morphology may be explained simply as results of mechanical (physical) interactions among the skeletal, nervous, and sensory components during head development at reduced size. This provides further evidence of the role of nervous, sensory, and other "soft" tissues in cranial skeletal morphogenesis, and reinforces the need to consider these tissues in analyses of skull evolution.     

The effect of domestication on brain size and composition in the mink (Mustela vison)

The sizes of total brain, the five fundamental brain parts, and certain telencephalic structures were measured in wild mink ( Mustela vison energumenos ) and ranch mink of a Dark Standard strain of the same species. By means of intraspecific allometric methods for analysing the relationship between brain weight and body weight (net carcass weight), the volumes of the brain parts were compared in both groups. In general, total brain, as well as all the parts measured, were smaller in size in ranch mink independent of body size, age, and sex, indicating that domestication has led to a decrease in size. There were differences in the amount of decrease in various brain parts. These are discussed in connection with domestication time, with comparable results obtained in other species, and with regard to the functional importance of the brain parts.     

Allometric comparison of skulls from two closely related weasels, Mustela itatsi and M. sibirica

We conducted an interspecific comparison of skulls from two closely related but differently sized mustelid species, Mustela itatsi and M. sibirica (Mammalia, Carnivora, Mustelidae); a sexual comparison within the latter species showed remarkable size dimorphism. We clarified several differences in skull proportion related to size using allometric analyses and qualitative comparisons. Allometric analysis revealed that the skulls of male M. itatsi (the smaller species) have a relatively long palate; a slender viscerocranium and postorbital constriction; a broad, short, and low neurocranium; small carnassials; and a short mandible with a thin body and small ramus compared to the skulls of male M. sibirica (the larger species). Similar results were obtained when male M. itatsi were compared to female M. sibirica, although the male M. itatsi had a broader viscerocranium than female M. sibirica. A sexual comparison in M. sibirica revealed a larger skull size among the males with a relatively wide viscerocranium; wide postorbital constriction; a slender, long, and high neurocranium; short and wide auditory bullae; short carnassials; and a long and high mandible compared to females. Qualitative comparisons revealed changes in a few characters depending on skull size or with respect to some cranial components in each species. The interspecific differences observed were clearly larger than the intraspecific differences for three qualitative characters. The allometric and qualitative differences detected between these species suggest that each species is not simply the dwarf and/or giant morph of the other, and complicated differences were clarified.     

Morphometric variation of the skull during postnatal development in the Lowland European bisonBison bonasus bonasus

We studied the variation of linear measurements and skull capacity in Lowland European bisonBison bonasus bonasus (Linnaeus, 1758) during postnatal development, and the dependencies of the parameters in relation to sex, age, and body mass of the animals. Material consisted of 599 bison skulls (310 males and 289 females), within the age range of 1 month to 21 years (males) and to 27 years (females). In the group of calves to 1 year old, no sex connected differences in skull measurements were observed, whereas the skull capacity in older calves was significantly larger (0.01>p>0.001) in males than in females. From the third year of life, most skull measurements display characteristics of sexual dimorphism. Skull development in both sexes is most intensive during the first three years of life, and slows from the age of 5. In older individuals of both sexes (≥ 6 years), orbital breadth continues growing and, in females, breadth of splanchnocranium continues increasing. Growth in a bison’s skull capacity is most intensive up to the third year of life and slows from the age of 5. During postnatal development, a bison skull grows proportionally except the neurocranium, which grows slightly slower in comparison with basal length and its development finishes earlier than that of splanchnocranium. In ontogenesis, a bison skull grows much slower compared to body mass. In relation to body mass, skull capacity and the height of neurocranium grow most slowly while orbital breadth grows most intensively. The results obtained were compared with data on skull sizes of bison born in 1930–1950 and bred in captivity and with skulls of the American bisonBison bison. Inbreeding is probably responsible for some types of phenotypic abnormalities in the skull which appear in modern European bison.     

A Comparison of Brain Gene Expression Levels in Domesticated and Wild Animals

Domestication has led to similar changes in morphology and behavior in several animal species, raising the question whether similarities between different domestication events also exist at the molecular level. We used mRNA sequencing to analyze genome-wide gene expression patterns in brain frontal cortex in three pairs of domesticated and wild species (dogs and wolves, pigs and wild boars, and domesticated and wild rabbits). We compared the expression differences with those between domesticated guinea pigs and a distant wild relative (Cavia aperea) as well as between two lines of rats selected for tameness or aggression towards humans. There were few gene expression differences between domesticated and wild dogs, pigs, and rabbits (30–75 genes (less than 1%) of expressed genes were differentially expressed), while guinea pigs and C. aperea differed more strongly. Almost no overlap was found between the genes with differential expression in the different domestication events. In addition, joint analyses of all domesticated and wild samples provided only suggestive evidence for the existence of a small group of genes that changed their expression in a similar fashion in different domesticated species. The most extreme of these shared expression changes include up-regulation in domesticates of SOX6 and PROM1, two modulators of brain development. There was almost no overlap between gene expression in domesticated animals and the tame and aggressive rats. However, two of the genes with the strongest expression differences between the rats (DLL3 and DHDH) were located in a genomic region associated with tameness and aggression, suggesting a role in influencing tameness. In summary, the majority of brain gene expression changes in domesticated animals are specific to the given domestication event, suggesting that the causative variants of behavioral domestication traits may likewise be different.     

饶阳地区三种农田啮齿类头骨形态比较及性二型

关于动物头骨形态方面的研究报道曾受到了较多的关注（Cesani et al., 2003）.头骨形态是动物形态特征的一个重要方面,受遗传和环境因素的共同影响,存在着种间、性别和地理差异（张焱等,1999）.头骨的形态特征不仅是动物生理上的一个基本解剖指标,同时也是动物分类（李晓晨和王廷正,1992）和年龄鉴定（张洁,1989）的重要依据,是动物生态研究的重要指标.对头骨形态的研究,不仅对动物的种间、种内关系（Yom et al.,1999）和生态特征（Barlow et al.,1997）等方面的研究具有重要的参考价值,在动物的地理演化、发育生物学以及进化生态学的研究中也具有十分重要的意义.     

Foraging strategies,craniodental traits,and interaction in the bite force of Neotropical frugivorous bats (Phyllostomidae: Stenodermatinae)

1. Bats in the family Phyllostomidae exhibit great diversity in skull size and morphology that reflects the degree of resource division and ecological overlap in the group. In particular, the subfamily Stenodermatinae has high morphological diversification associated with cranial and mandibular traits that are associated with the ability to consume the full range of available fruits (soft and hard).
2. We analyzed craniodental traits and their relationship to the bite force in 343 specimens distributed in seven species of stenodermatine bats with two foraging strategies: nomadic and sedentary frugivory. We evaluated 19 traits related to feeding and bite force in live animals by correcting bite force with body size.
3. We used a generalized linear model (GLM) and post hoc tests to determine possible relationships and differences between cranial traits, species, and sex. We also used Blomberg''s K to measure the phylogenetic signal and phylogenetic generalized least‐squares (PGLS) to ensure the phylogenetic independence of the traits.
4. We found that smaller nomadic species, A. anderseni and A. phaeotis , have a similar bite force to the large species A. planirostris and A. lituratus; furthermore, P. helleri registered a bite force similar to that of the sedentary bat, S. giannae. Our study determined that all the features of the mandible and most of the traits of the skull have a low phylogenetic signal. Through the PGLS, we found that the diet and several cranial features (mandibular toothrow length, dentary length, braincase breadth, mastoid breadth, greatest length of skull, condylo‐incisive length, and condylo‐canine length) determined bite force performance among Stenodermatiane.
5. Our results reinforce that skull size is a determining factor in the bite force, but also emphasize the importance of its relationships with morphology, ecology, and phylogeny of the species, which gives us a better understanding of the evolutionary adaptions of this highly diverse Neotropical bat group.

     

Jaw muscles and the skull in mammals: the biomechanics of mastication.

Among non-mammalian vertebrates, rigid skulls with tight sutural junctions are associated with high levels of cranial loading. The rigid skulls of mammals presumably act to resist the stresses of mastication. The pig, Sus scrofa, is a generalized ungulate with a diet rich in resistant foods. This report synthesizes previous work using strain gages bonded to the bones and sutures of the braincase, zygomatic arch, jaw joint, and mandible with new studies on the maxilla. Strains were recorded during unrestrained mastication and/or in anesthetized pigs during muscle stimulation. Bone strains were 100-1000 micro epsilon, except in the braincase, but sutural strains were higher, regardless of region. Strain regimes were specific to different regions, indicating that theoretical treatment of the skull as a unitary structure is probably incorrect. Muscle contraction, especially the masseter, caused strain patterns by four mechanisms: (1) direct loading of muscle attachment areas; (2) a compressive reaction force at the jaw joint; (3) bite force loading on the snout and mandible; and (4) movement causing new points of contact between mandible and cranium. Some expected patterns of loading were not seen. Most notably, strains did not differ for right and left chewing, perhaps because pigs have bilateral occlusion and masseter activity.     

Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae,Caimaninae)

Melanosuchus niger is a caimanine alligatorid widely distributed in the northern region of South America. This species has been the focus of several ecological, genetic and morphological studies. However, morphological studies have generally been limited to examination of interspecific variation among extant species of South American crocodylians. Here, we present the first study of intraspecific variation in the skull of M. niger using a two‐dimensional geometric morphometric approach. The crania of 52 sexed individuals varying in size were analysed to quantify shape variation and to assign observed shape changes to different types of intraspecific variation, that is, ontogenetic variation and sexual dimorphism. Most of the variation in this species is ontogenetic variation in snout length, skull depth, orbit size and the width of the postorbital region. These changes are correlated with bite force performance and probably dietary changes. However, a comparison with previous functional studies reveals that functional adaptations during ontogeny seem to be primarily restricted to the postrostral region, whereas rostral shape changes are more related to dietary shifts. Furthermore, the skulls of M. niger exhibit a sexual dimorphism, which is primarily size‐related. The presence of non‐size‐related sexual dimorphism has to be tested in future examinations.     

Component analysis of craniologic characters of silver foxes (Vulpes fulvus Desm.) and their changes arising under domestication]

The data on cranial measurements performed in silver foxes indicate that there are differences in sizes measured between the farm foxes--bred population and the population selected for domestication. A method of principal components was used to analyse the cranial measurements and their changes under domestication. The first component covers about 50% of cranial diversity, which is interpreted as variation in the total skull size. This component clearly separates the two sexes, but not different populations. The second component presumably reflects the growth rate allometry between the skull length and width. The third and fourth components are measurements of skull width; the fifth one reflects the sizes of brain skull. None of these components clearly separate the foxes from farm--bred and domesticated populations. However, some differences in distribution are observed.     

Cranial ontogenetic variability,sex ratio and age structure of the Red fox

Describing the sex ratio, age structure of the population and ontogenetic variability of Red fox, Vulpes vulpes (Canidae, Carnivora) skull parameters, this study is based on 416 male and 289 female skulls collected in the Czech Republic. The skulls analysed came from feral individuals, that were shot by hunters. The male to female ratio was 1:0.69 regarding the whole population. Individuals younger than one year prevailed in the population (54% males, 48% females were in their first year of life). Four growth patterns of skull dimensions were described. The first group included mainly skull length dimensions (e.g. condylobasal length). They grew rapidly until the sixth month of life, becoming stabilised afterwards. The second group comprised parameters that were stable throughout the life (e.g. cheek tooth rows). Measurements representing the third growth pattern showed continual growth (mainly width dimensions, e.g. zygomatic breadth). Conversely, smaller dimensions of postorbital breadth were observed after the sixth month of life. Postorbital breadth represented the fourth growth pattern. It was concluded, that male and female Red foxes had similar ontogenetic skull development, even though there were some differences, e.g. in jugular breadth, which increased after the age of six months in males unlike in females.     

Ancient and modern steps during the domestication of guinea pigs (Cavia porcellus L.)

To test whether there are differences between living lineages of domestic guinea pigs Cavia porcellus , we studied 118 specimens from six breeds collected along six Andean countries as well as 15 from the wild cavy species ( Cavia tschudii ). The mean weight and body length of 15 adult wild cavies (295±31 g, 242±8.3 mm) were significantly smaller than 25 creole guinea pigs from Bolivia and Chile (639±157 g, 287±23.7 mm, respectively). Eighteen laboratory/pet guinea pigs (including the English Pirbright breed) were also smaller (900±173 g, 308±21 mm) than 25 improved ones from Peru (Tamborada breed, 1241±75.4 g, 317±12 mm) and Ecuador (Auqui breed, 1138±65.5 g, 307±8 mm). Similar size increases appeared in the first axis of a principal component analysis of six skeletal measurements, recovering 84% of total variation. Phylogenetic and haplotype analyses of complete cytochrome b gene sequences consistently joined all 22 domestic individuals (13 shared unambiguous substitutions, 100% bootstrap in 1000 replicates), probably from a single first ancient domestication in the western Andes. Six laboratory/pet sequences were also joined within a common branch (six shared substitutions, 96% bootstrap), probably from a documented European second phase. By contrast, those from improved Auqui joined a northern creole subgroup (one shared substitution, 84% bootstrap), and those from Nativa and improved Tamborada clustered together and with a southern creole subgroup (four shared substitutions, 86% bootstrap); this suggests at least two independent modern events during a more complex third phase, producing two improved guinea pigs selected for size and meat. Cavia tschudii sequences showed some unexpected geographic variation.     

Miniaturization and its effects on cranial morphology in plethodontid salamanders, genus Thorius (Amphibia: Plethodontidae). I. Osteological variation

Cranial skeletal morphology, ontogeny and variation are examined in five species of Thorius , a genus of diminutive plethodontid salamanders that are among the smallest, extant, tailed tetrapods. The skull of adull Thorius is characterized by: (1) limited development or absence of several ossified elements and dentition; (2) increased inter-and intraspecific variability; (3) novel morphological configurations of the braincase and jaw suspensorium. Posthatching cranial mineralization in all species of Thorius is truncated precociously with respect to that typical of larger and more generalized plethodontid genera, such as Pseudoeurycea. These features implicate paedomorphosis as a predominant mechanism responsible for the evolution of decreased size in Thorius from larger plethodontid ancestors. Interspecific differences in cranial morphology are evident; species may be characterized by greater or lesser degrees of truncated development. However, there is no consistent relationship between degree of paedomorphosis and mean adult body size in interspecific comparisons. Adult morphology of several individual elements represent potentially useful taxonomic characters for distinguishing species.
Reduction, increased variability, and morphological novelty are common to many lineages of dwarfed taxa. They represent a null hypothesis for examination of the developmental mechanisms and morphological consequences of miniaturization in other groups.     

Correlation coefficients in human skulls: significant sexual differences

This study of 500 male and female skulls proves: significant differences of correlation coefficients (p less than or equal to 0.01) occur between the two sexes. The combined variables are: 1. bizygomatic breadth (45)/skull base length (5) 2. bizygomatic breadth (45)/foramen magnum length (7) 3. bizygomatic breadth (45)/foramen magnum breadth (16) 4. bizygomatic breadth (45)/basion-bregma height (17) 5. foramen magnum breadth (16)/skull base length (5) Female skulls have generally higher correlation coefficients values than males which is interpreted as an indication of homogeneous growth.     

Skull morphometry of Pygoscelis (Sphenisciformes): inter and intraspecific variations

Morphological diversity of 47 pygoscelid skulls was tested empirically through geometric morphometrics approach. Using 14 landmarks, shape is analyzed independently of other aspects of the body form. The shape disparity within and between the three living species of Pygoscelis is explored as well as how the structure of the skull is related to food preferences. Comparison of the three mean configurations of the species suggests that differences among groups are small relative to the variability within each group. However, some differences at the posterior portion of the braincase are indicated. Sexual dimorphism within each species is not noticeable. Comparison with a piscivorous species (Spheniscus humboldti) shows two cranial patterns: pygoscelid type with wide nasal gland depression limited by well-marked edges, shallow temporal fossae, and a poorly developed temporal nuchal crest; the second type represented by Spheniscus humboldti with laterally open nasal gland depression, deep temporal fossae, and a well-developed temporal nuchal crest.     

Morphometric variation in the Hooded seal (Cystophora cristata)

Intra- and intersexual variation in 16 skull dimensions and total body length of 233 aged Hooded seals caught in the north-west Atlantic were investigated. Absolute growth was described by asymptotic growth curves applied to single dimensions, as well as to scores on the first principal component of logarithmically transformed cranial data, which are believed to reflect the multivariate nature of growth. All dimensions were found to be fully developed later in males than in females. The growth of the male skulls was found to continue for more than 20 years, while the females approach final size about–7 years earlier than males, according to the scores on the first principal component. Female seals were found to reach 86 % of final total body length at the time of maturation, which is a generalized pinniped pattern. In both sexes, a yearling skull was characterized by a large brain-case, which decreased relatively with growth. The male skulls were further characterized by an increase in zygomatic width and orbital width in relation to basal length, a pattern which was not found in females.
All the asymptotic values were significantly larger in males than in females. The dimorphism develops mainly as a result of prolonged growth of males after the attainment of sexual maturity.     

The influence of feeding on the evolution of sensory signals: a comparative test of an evolutionary trade‐off between masticatory and sensory functions of skulls in southern African Horseshoe bats (Rhinolophidae)

The skulls of animals have to perform many functions. Optimization for one function may mean another function is less optimized, resulting in evolutionary trade‐offs. Here, we investigate whether a trade‐off exists between the masticatory and sensory functions of animal skulls using echolocating bats as model species. Several species of rhinolophid bats deviate from the allometric relationship between body size and echolocation frequency. Such deviation may be the result of selection for increased bite force, resulting in a decrease in snout length which could in turn lead to higher echolocation frequencies. If so, there should be a positive relationship between bite force and echolocation frequency. We investigated this relationship in several species of southern African rhinolophids using phylogenetically informed analyses of the allometry of their bite force and echolocation frequency and of the three‐dimensional shape of their skulls. As predicted, echolocation frequency was positively correlated with bite force, suggesting that its evolution is influenced by a trade‐off between the masticatory and sensory functions of the skull. In support of this, variation in skull shape was explained by both echolocation frequency (80%) and bite force (20%). Furthermore, it appears that selection has acted on the nasal capsules, which have a frequency‐specific impedance matching function during vocalization. There was a negative correlation between echolocation frequency and capsule volume across species. Optimization of the masticatory function of the skull may have been achieved through changes in the shape of the mandible and associated musculature, elements not considered in this study.