Comparative neurocranial morphology of two hammerhead species (Chondrichthyes: Carcharhiniformes: Sphyrnidae) with notes on cranial morphology in Sphyrnidae

Recognition of a particularly derived eco-behavioral strategy for the genus Alouatta has been crucial for studying and understanding its equally derived cranial and postcranial morphology. The unique architecture of the skull and mandible has very likely evolved in relation to both masticatory correlates associated with an increasingly folivorous diet as well as the use of vocal communication associated with social behavior and an energy-minimizing strategy. Comparisons of cranial morphology using three-dimensional geometric morphometrics have highlighted significant interspecific shape differences. Alouatta seniculus is the most divergent in both cranial and hyoid morphology and exhibits the most pronounced levels of sexual dimorphism in those areas. Cranial variability is expressed in facial prognathism and airorhynchy, basicranial flexure, and zygomatic height. Inter- and intraspecific differences based on these axes of variation are very likely linked to interspecific variations in diet, behavior, and life history. This is further evident in the dental anatomy of the genus, indicating adaptations to a shift to a more folivorous diet. In addition, recent studies provide further evidence for significant inter- and intraspecific variations in hyoid size and shape. Alouatta seniculus possesses the largest and most inflated hyoid bulla, and the species that occupy distributional extremes (A. palliata, A. caraya) are differentiated by highly distinct hyoid shapes. These data indicate a complex relationship between morphology and behavior, with possible biogeographic implications. In terms of postcranial morphology, the forelimb bears a mixture of features that favor quadrupedalism with restricted abduction and overhead extension, providing stable contact and support along branches. In contrast, the hind limb appears to allow for a wider range of movement in all joints, with an emphasis on thigh extension and abduction, leg rotation, and powerful grasping with a habitually inverted foot. Interspecific variation reveals traits that can be ultimately related to subtle differences in the frequency of use of different positional modes, associated with variable eco-social factors. These results, deriving from different anatomical regions, provide evidence for understanding morphological variation across howler species in terms of morphofunctional adaptations, environmental pressure, and niche partitioning.

Bite force is often used as a predictive indicator of an animal's feeding ecology, although the premise that there is a direct link between diet and cranial morphology can be difficult to test empirically. Studies that have examined this question tend to rely on generalizations of a species’ diet, and age and sex differences are rarely considered. Red foxes (Vulpes vulpes) are introduced predators in Australia, allowing large sample size collection through culling for comparison of skull morphology (size, morphometry, weight), demographics (age, sex) and diet (stomach contents). Over half (57%) of the 540 animals we sampled were juveniles (<1 year old; dispersing from their natal sites) and most variation in skull morphometry was driven by age; adults had significantly more robust skulls than juveniles, with greater estimated bite force. Sexual dimorphism (body mass and body length) was reflected in longer, heavier skulls of males. We also recorded significant sex and age differences in diet. Sheep carrion comprised 47–65% of diet volume; however, adult females ate less sheep but had more mice and invertebrates in their diet than males or juveniles of both sexes. This dietary separation for adult females does not appear to be directly due to estimated bite force constraints, but probably rather prey availability, which may reflect feeding behaviour and space use patterns. Juveniles (both sexes) showed as much consumption of sheep carrion as adult males, despite their lower estimated bite force than adults. This is the first study that directly compares ontogenetic and sex differences in the diet of a carnivore together with their cranial morphology and estimated bite force, and highlights limitations of inferring diet partitioning from skull morphology alone.

Early Holocene human skeletal remains from Sumidouro Cave, Lagoa Santa, Brazil: History of discoveries, geological and chronological context, and comparative cranial morphology

The New World cichlids Petenia splendida and Caquetaia spp. possess extraordinarily protrusible jaws. We investigated the feeding behavior of extreme (here defined as greater than 30% head length) and modest jaw-protruding Neotropical cichlids by comparing feeding kinematics, cranial morphology, and feeding performance. Digital high-speed video (500 fps) of P. splendida, C. spectabile, and Astronotus ocellatus feeding on live guppy prey was analyzed to generate kinematic and performance variables. All three cichlid taxa utilized cranial elevation, lower jaw depression, and rotation of the suspensorium to protrude the jaws during feeding experiments. Extreme anterior jaw protrusion in P. splendida and C. spectabile resulted from augmented lower jaw depression and anterior rotation of the suspensorium. Morphological comparisons among eight cichlid species revealed novel anterior and posterior points of flexion within the suspensorium of P. splendida and Caquetaia spp. The combination of anterior and posterior loosening within the suspensorium in P. splendida and Caquetaia spp. permitted considerable anterior rotation of the suspensorium and contributed to protrusion of the jaws. Petenia splendida and C. spectabile exhibited greater ram distance and higher ram velocities than did A. ocellatus, resulting primarily from increased jaw protrusion. Petenia splendida and C. spectabile exhibited lower suction feeding performance than A. ocellatus, as indicated by lower suction-induced prey movements and velocities. Thus, extreme jaw protrusion in these cichlids may represent an adaptation for capturing elusive prey by enhancing the ram velocity of the predator but does not enhance suction feeding performance.

Biological variation in the mammalian skull is the product of a series of factors including changes in gene expression, developmental timing, and environmental pressures. When considering the diversity of extant mammalian crania, it is important to understand these mechanisms that contribute to cranial growth and in turn, how differences in cranial morphology have been attained. Various researchers, including Dr. Sue Herring, have proposed a variety of mechanisms to explain the process of cranial growth. This work has set the foundation on which modern analysis of craniofacial morphology happens today. This study focused on the analysis of modularity in three mammalian taxa, all of which exhibit facial reduction. Specifically, we examined facial reduction as a morphological phenomenon through the use of two-module and six-module modularity hypotheses. We recorded three-dimensional coordinate data for 55 cranial landmarks that allowed us to analyze differences in cranial shape in these three taxa (primates n = 88, bats n = 64, dogs n = 81). When assessing modularity within the two-module modularity hypothesis specifically, dogs exhibited the lowest levels of modularity, while bats and primates both showed a slightly more modular covariance structure. We further assessed modularity in the same sample using the Goswami six-module model, where again dogs exhibited a low degree of modularity, with bats and primates being more moderate. We then broke the sample into subsets by analyzing each morphotype separately. We hypothesized that the modularity would be more pronounced in the brachycephalic morphotype. Surprisingly, we found that in brachycephalic dogs, normocephalic dogs, brachycephalic primates, and normocephalic primates, there was a moderate degree of modularity. Brachycephalic bats had a low degree of modularity, while normocephalic bats were the most modular group observed in this study. Based on these results, it is evident that facial reduction is a complex and multifaceted phenomenon with unique morphological changes observed in each of the three taxa studied.

Cranial suture morphology of Lystrosaurus and the generalized dicynodont Oudenodon was investigated to determine the strain environment during mastication, which in turn may indicate a difference in cranial function between the two taxa. Finite element (FE) analysis indicated that less strain accumulated in the cranium of Lystrosaurus during orthal bite simulations than in Oudenodon. Despite the overall difference in strain magnitude, moderate to high FE-predicted strain accumulated in similar areas of the cranium of both taxa. The suture morphology in these cranial regions of Lystrosaurus and Oudenodon was investigated further by examination of histological sections and supplemented by observations of serial sections and computed tomography (CT) scans. The predominant type of strain from selected blocks of finite elements that contain sutures was determined, enabling comparison of suture morphology to strain type. Drawing from strain-suture correlations established in extant taxa, the observed patterns of sutural morphology for both dicynodonts were used to deduce cranial function. The moderate to high compressive and tensile strain experienced by the infraorbital bar, zygomatic arch, and postorbital bar of Oudenodon and Lystrosaurus may have been decreased by small adjustive movements at the scarf sutures in those regions. Disparities in cranial suture morphology between the two taxa may reflect differences in cranial function. For instance, the tongue and groove morphology of the postorbital-parietal suture in Oudenodon could have withstood the higher FE-predicted tensile strain in the posterior skull roof. The scarf premaxilla-nasal suture of Lystrosaurus provided an additional region of sutural mobility in the anterior surface of the snout, suggesting that Lystrosaurus may have employed a different biting regime than Oudenodon. The morphology of several sutures sampled in this study correlated with the FE-predicted strain, although other cranial functional hypotheses remain to be tested.

The objectives of the study was to assess and compare the cranial base morphology in long face and short face. The study comprises 64 individuals (32 in each group). Group I consists of individual's with long face and group II comprises individuals with short face. Individuals were classified into groups with the help of lateral cephalometric parameters like Gonial Angle, Anterior facial height, Posterior facial height, Jarabak's Ratio, and Y-axis. Linear and angular measurement in the cranial base was taken. The results showed statistically significant difference in Sella-Nasion, Sella-Basion and Nasion-Sella-Basion between the two groups. Sella-Spinoethmoidal suture, Nasion-Spinoethmoidal suture, Spinoethmoidal suture-Sella-Basion, Frankfurt Horizontal-Sella-Nasion, Frankfurt horizontal-Sella-Basion, Frankfurt horizontal-Sella-Spinoethmoidal suture showed no statistically significant difference among the groups. The study concluded that individuals with long face have a greater Sella-Nasion, Sella-Basion, Nasion-Sella-Basion compared to individuals with short face.

Internal cranial anatomy is a challenging area to study in fossilized skulls because of small sample sizes and varied post-mortem preservational alterations. This difficulty has led to the lack of correspondence between results obtained from direct osteological observation and from more indirect reconstruction methods. This paper presents corroborating evidence from direct osteological observation and from reconstruction based on computed X-ray tomography (CT) on the internal cranial anatomy of the ankylosaurid dinosaur Euoplocephalus tutus. A remarkable specimen of Euoplocephalus preserves rarely observed internal cranial structures such as vascular impressions in the nasal cavity, olfactory turbinates and possible impressions of conchae. Comparison with fossils and CT models of other taxa and other Euoplocephalus specimens adds osteological evidence for the previously reconstructed nasal cavity in this dinosaur and revises the previously described braincase morphology. A new interpretation of the ethmoidal homology identifies a mesethmoid, sphenethmoid and ectethmoid. These ethmoidal ossifications are continuous with the mineralized walls of the nasal cavity. The location of the olfactory fenestra provides further evidence that the olfactory regions of the nasal cavity are pushed to the sides of the main airway. This implies that the function of the vascular impressions in the nasal cavity and the looping of the cavity are not related to olfaction. A byproduct of the elongate, looping airway is a dramatic increase in surface area of the nasal respiratory mucosa, which in extant species has been linked to heat and water balance. A role in vocalization as a resonating chamber is another possible function of the looping and elongation of the nasal cavity. Olfaction remains as a possible function for the enlarged olfactory region, suggesting that multiple functions account for different parts of the ankylosaurid nasal cavity that underwent substantial modification. Cranial endocasts show negligible variation within Euoplocephalus, which lends some confidence to interspecific comparisons of endocranial morphology.

This study looked at the differences in cranial morphology of tree squirrel species (F. leucogenys) from four different locations in Rainforest part of Nigeria. The squirrels were captured through the use of locally fabricated live traps made of wire-mesh and steel. Trapped specimens were immediately transferred to the laboratory in captivity cages, where they were euthanized in a bell-jar containing chloroform-soaked cotton wool. Specimens were then preliminarily identified to the generic level, using an identification key. The skull of 131 tree squirrels were prepared using Long Island Natural History Museum guide and the sex of the specimens was determined by visual inspection of the external genitalia. The skull and other body parameters were measured using digital venier caliper. The results showed that the body parameters (HBL, TL, TBL, EL, HFL and BW) measured were slightly different from one location to another. Also, the cranial measure showed some similarities among some locations (Ile-Ife, Emure-Ekiti and Ado-Ekiti), while measurement on squirrels from Sekona was different from other three locations. In conclusion, the cranial measurements of the tree squirrels shows there was no new species of F. leucogeny from the sampled locations.

This study examined the differences in cranial morphology of tree squirrel species (Heliosciuru srufobranchium) from four different locations in rainforest part of Nigeria. Locally fabricated live traps made of wire-mesh and steel were used to capture the squirrels. Trapped tree squirrels were immediately transferred to the laboratory in cages, where they were euthanized by placement in a bell-jar containing chloroform-soaked cotton wool. Individuals from two locations (33 from Ile Ife and 49 from Emure Ekiti) were then preliminarily identified to the generic level, using an identification key. Eighty-two (82) skulls were prepared from them, using Long Island Natural History Museum Guide. The sexes of the specimens were determined by visual inspection of the external genitalia. Twenty-seven (27) cranial and six (6) external body parameters were measured using digital Vernier caliper. One-way analysis of variance at p<0.05 was used to determine the difference in measured body parameters and Principal Component Analysis was used to differentiate the measured skull parameters. The results showed that the body parameters; head body length, tail length, tail body length, ear length, hind foot length and body weight measured were slightly different from one location to another. Also, the cranial measurements showed some similarities and differences between the two locations (Ile Ife and Emure Ekiti). In conclusion, the cranial measurements of the tree squirrels from the two locations (Ile Ife and Emure Ekiti) showed slight differences despite the fact that the tree squirrels are from the same genus (H. rufobranchium). Keywords: Heliosciurus rufobranchium; rainforest; tree squirrel; cranial; morphometric

The aim of this study was to compare the craniofacial and neurocranial morphology of adults with osteogenesis imperfecta (OI) with controls and to elucidate whether osseous origin impacts on morphological deviations in OI. Fifty-four adults (mean age 45.8) with OI type I, 14 adults (mean age 42.6) with OI types III/IV and 49 adult controls (mean age 41.0) were included. All participants had European ethnicity. Cranial morphology was assessed by 2D-cephalometry. Comparison between groups was made by multiple regression analyses. Comparison between OI groups and controls: (1) Dimension of the maxilla and mandible, respectively was reduced (P < .01), and in relation to the posterior cranial base, the maxilla was retro-positioned (P < .001), and the mandible was prognathic (P < .001). (2) The anterior face height was reduced (P < .04), and in OI types III/IV only, the maxilla was posteriorly inclined (P < .001). (3) Anterior cranial base (P < .001) and the dimension sella-frontale (P < .02) were short. (4) The sagittal dimension of the posterior cranial fossa was increased (P < .01), and the vertical dimension was reduced (P < .01). Adults with OI had a hypoplastic, retro-positioned and posteriorly inclined maxilla, a hypoplastic and forward-positioned mandible, and a reduced anterior face height. Deviations were seen in morphology of the posterior cranial fossa. The impact of OI on cranial morphology was generally more evident in OI type III/IV than in OI type I. OI impacts on osseous cranial structures irrespective of bony origin being intramembranous or endochondral.

Craniosynostosis, a common congenital anomaly, results from premature fusion of the cranial sutures. One of the forms of craniosynostosis is premature fusion of the metopic suture, referred to as trigonocephaly, but the diagnosis of metopic suture synostosis remains controversial. The purpose of this study was to clarify, using geometric morphometric analysis, if a metopic ridge alone observed in cases of mild trigonocephaly represents a pathological phenomenon. Three different cranial morphologies were compared among patients up to 2 years old who were categorized into the true group, the mild group, and the normal group, based on the presence or absence of specific symptoms, history of cranioplasty for trigonocephaly, or lack of any abnormality on computed tomography. Using the obtained computed tomography images, 235 anatomical landmarks and semi-landmarks were plotted on the entire cranial surface for analysis of neurocranial morphology, and the cranial shapes represented by landmarks were analyzed using geometric morphometrics. Principal components of shape variations among specimens were then computed, based on the variance-covariance matrix of the Procrustes residuals of all specimens, and statistically analyzed. The principal component analyses of the variations in endocranial shape, frontal bone shape, and occipital bone shape did not show any significant differences in cranial morphology between mild trigonocephaly and normal skulls; however, true trigonocephaly was found to differ significantly from mild trigonocephaly and normal skulls. These findings suggest that in assessments of cranial morphology, the presence of a ridge alone cannot be diagnosed as fundamentally pathological, and may represent normal morphology.

The ability of cranial morphology to reflect population/phylogenetic history, and the degree to which it might be influenced by environmental factors and selection pressures have been widely discussed. Recent consensus views cranial morphology as largely indicative of population history in humans, with some anatomical cranial regions/measurements being more informative on population history, while others being under selection pressure. We test earlier findings using the largest and most diverse cranial dataset available as yet: 7,423 male specimens from 135 geographic human population samples represented by 33 standard craniometric linear measurements. We calculated Mahalanobis D(2) for three datasets: complete cranial dataset; facial measurement dataset; and neurocranial measurement dataset; these morphological distance matrices were then compared to matrices of geographic distances as well as of several climatic variables. Additionally, we calculated Fst values for our cranial measurements and compared the results to the expected Fst values for neutral genetic loci. Our findings support the hypothesis that cranial, and especially neurocranial morphology, is phylogenetically informative, and that aspects of the face and cranium are subject to selection related to climatic factors. The Fst analysis suggest that selection to climate is largely restricted to groups living in extremely cold environments, including Northeast Asia, North America, and Northern Europe, though each of these regions appears to have arrived at their morphology through distinct adaptive pathways.

The relationship between the Yuanmou hominoid and the Lufeng hominoid (Lufengpithecus lufengensis), both from Yunnan Province, China, and among the most abundantly represented fossil hominoid of Eurasia, has been the subject of much debate. In the past 10 years, comparative studies of cranial and dental morphology of the Lufeng and Yuanmou hominoids indicate that the two hominoids resemble each other more than either of them resemble any other Miocene hominoid. In this paper, we summarize the dental differences seen between the Yuanmou and Lufeng hominoids, and discuss their implications. Our results show that the Lufeng hominoid has relatively smaller front teeth, smaller M1, and higher SQ than the Yuanmou hominoid, indicating a more folivorous or soft diet, such as leaves and berries, in the former. Tooth wear analysis indicates that both upper and lower molars of the Yuanmou hominoid were more heavily worn than those of the Lufeng hominoid, offering additional support to the findings of tooth size proportion and shearing crest development. We propose that different diets might have characterized the two hominoid populations, the differences possibly related to the environment, behavior pattern, and population structure of the Yuanmou and Lufeng hominoids.

Examining species boundaries using data from multiple independent sources is an appropriate and robust method to identify genetically isolated evolutionary units. We used 5 data sets—cytochrome b (Cytb), cytochrome c oxidase (COI), amplified fragment length polymorphisms (AFLPs), karyotypes, and morphology—to estimate phylogenetic relationships and species limits within woolly bats, genus Kerivoula, from Southeast Asia. We genetically analyzed 54 specimens of Kerivoula from Malaysia, assigned to 6 of the 10 species currently reported from the country. Phylogenetic analyses of nuclear AFLPs (33 specimens) and mitochondrial DNA sequences from Cytb (51 specimens) and COI (48 specimens) resulted in similar statistically supported species-level clades with minimal change in branching order. Using comparisons of cranial and dental morphology and original species descriptions, we assigned the resulting phylogenetic clades to K. hardwickii, K. intermedia, K. lenis, K. minuta, K. papillosa, K pellucida, and 1 unidentified species. Karyotypes further documented variability among the 6 clades. Five different karyotypes were identified, with 2 species having indistinguishable karyotypes. We compared our COI gene sequence data to 110 specimens of Kerivoula from Southeast Asia made available by researchers of the Barcode of Life Database. Our Cytb and AFLP species identifications were congruent with those in the COI database. Intraspecific geographic variation of about 5–7% sequence divergence was observed in Cytb and COI genes within both K. hardwickii and K. minuta. Relaxed molecular clock analyses indicated a late Oligocene to early Miocene origin of the Kerivoulinae with intraspecific diversification events coinciding with the late Pliocene and Pleistocene epochs in at least 6 species. Specimens from Sabah (northeastern Borneo) showed relatively high genetic divergence compared to those between Peninsular Malaysia and Sarawak (southwestern Borneo), indicating Pleistocene or Pliocene refugia in Borneo. We conclude that at least 1 distinct lineage of Kerivoula is yet to be described from Borneo and that the intraspecific geographic variation in some species agrees with previous studies on the diversification of flora and fauna in Borneo. DOI: 10.1644/09-MAMM-A-361.1.