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Abstract Bones of the skull are traditionally categorized as derived from either endochondral or intramembranous bone. In our previous work, we have observed the interaction of different tissue types in growth of the skull. We find the dichotomy of intramembranous and endochondral bone to be too restrictive, limiting our interpretation of sources of biological variation. Here, we advocate for the use of the termectochondralbone to describe bone that originates from an endochondral model but is directed in its subsequent growth by membranes and other fascial attachments. Growth of the alisphenoid and orbitosphenoid are described as two examples of ectochondral bone, influenced in their shape primarily by the surrounding soft tissues. Ectochondral bone may be an ideal mechanism for rapidly evolving new phenotypes. Instead of evolving novelties by altering morphology of the cartilage template, novel features may be formed by ectochondral ossification, a more direct and rapid mode of osteogenesis than that of the cartilage template. 

ABSTRACT The study of primate auditory morphology is a significant area of interest for comparative anatomists, given the phylogenetic relationships that link primate hearing and the morphology of these auditory structures. Extensive literature addresses the form‐to‐function relationship of the auditory system (outer, middle, and inner ear) in primates and, by extension, provides insight into the auditory system of extinct primates and even modern humans. We add to this literature by describing the ontogenetic trajectory of the middle ear cavity and ossicular chain (malleus, incus, and stapes) due to their critical role in relaying auditory stimuli for interpretation. We examined middle ear morphology in neonatal primates and adult primates using a taxonomically broad sample. We focused primarily on nocturnal primate taxa (Daubentonia,Loris,Galago,Aotus, andTarsier), which are underrepresented in the literature. However, we also included three diurnal taxa (Macaca,Lemur, andSaguinus). Using 3D Slicer, we visualized middle ear structures in three dimensions using conventional micro CT data informed by diffusible iodine‐based contrast‐enhanced CT (diceCT) data. We illustrated how spatial relationships between otic elements, such as the various epitympanic sinuses of the middle ear and the auditory ossicles, vary throughout ontogeny. Our major findings include that the central tympanic cavity scaled with negative allometry in all taxa and that the accessory cavities scaled with isometry or positive allometry in most taxa. Despite these changes in chamber size, the size of the ear ossicles remained relatively consistent through ontogeny in most taxa. We confirmed our expectation that anthropoids exhibit an increase in the complexity of accessory cavities throughout ontogeny, mirroring the exponential pneumatization of the face in anthropoids. These findings provide an ontogenetic perspective and reveal further functional complexities of the middle ear as a conduit for sound proliferation and as a pressure regulator. 

Abstract Tusks are ever‐growing teeth present in mammals of the clade Paenungulata. Unlike the perpetually growing incisors of rodents, tusks are not used in mastication, and in at least some paenungulatans, the tusk is composed of dentin alone in adults. Few studies have provided tissue‐level information on tusks of adult paenungulatans with embedding techniques that identify epithelial and other soft tissues. In order to examine the mineralized tissues as well as the cells that form teeth, we studied a single, subadult rock hyrax (Procavia capensis) using microCT and paraffin histology with traditional staining as well as RUNX2 immunohistochemistry, and compared its teeth to scans of adult hyraxes. Three‐dimensional reconstructions from microCT volumes revealed that the tusk of this specimen is the only fully erupted replacement tooth, the first adult premolar (P1) is starting to erupt, and the first permanent molar (M1) is fully erupted, whereas all other replacement teeth and M2 remain in crypts. The tusk has a thin layer of enamel on its dorsal side; this is confirmed by histology. All deciduous premolars still possess roots that are in the process of resorption. Amelogenesis has progressed to maturation or nearly so in P1–P3. Notable histological characteristics of replacement premolars include the lack of a stellate reticulum in all except P4, and expression of RUNX2 in ameloblasts, a marker which is expressed by ameloblasts at all stages of amelogenesis. Since the pulp chambers of replacement premolars are relatively large compared to adults, a lengthy time in crypts may be important for dentin production. The results confirm that the hyrax has thin enamel on tusks, supporting the hypothesis that enamel is of limited importance for non‐feeding behaviors. 

Abstract Systems of the body develop in a modular manner. For example, neural development in primates is generally rapid, whereas dental development varies much more. In the present study, we examined development of the skull, teeth, and postcrania in a highly specialized leaping primate,Galago moholi. Eighteen specimens ranging from birth to adult were studied. Bones, teeth, and the cranial cavity (i.e., endocast) were reconstructed with Amira software based on microCT cross‐referenced to histology. Amira was also used to compute endocast volume (as a proxy for brain size). Reconstructions of the wrist and ankle show that ossification is complete at 1 month postnatally, consistent with the onset of leaping locomotion in this species. Endocranial volume is less than 50% of adult volume at birth, ~80% by 1 month, and has reached adult volume by 2 months postnatal age. Full deciduous dentition eruption occurs by 2 weeks, and the young are known to begin capturing and consuming arthropods on their own by 4 weeks, contemporaneous with the timing of bone and ankle ossification that accompanies successful hunting. The modular pattern of development of body systems inGalago moholiprovides an interesting view of a “race” to adult morphology for some joints that are critical for specialized leaping and clinging, rapid crown mineralization to begin a transitional diet, but perhaps more prolonged reliance on nursing to support brain growth. 

Abstract Nasal turbinals, delicate and complex bones of the nasal cavity that support respiratory or olfactory mucosa (OM), are now easily studied using high resolution micro‐computed tomography (μ‐CT). Standard μ‐CT currently lacks the capacity to identify OM or other mucosa types without additional radio‐opaque staining techniques. However, even unstained mucosa is more radio‐opaque than air, and thus mucosal thickness can be discerned. Here, we assess mucosal thickness of the nasal fossa using the cranium of a cadaveric adult dog that was μ‐CT scanned with an isotropic resolution of 30 μm, and subsequently histologically sectioned and stained. After co‐alignment of μ‐CT slice planes to that of histology, mucosal thickness was estimated at four locations. Results based on either μ‐CT or histology indicate olfactory mucosa is thicker on average compared with non‐olfactory mucosa (non‐OM). In addition, olfactory mucosa has a lesser degree of variability than the non‐OM. Variability in the latter appears to relate mostly to the varying degree of vascularity of the lamina propria. Because of this, in structures with both specialized vascular respiratory mucosa and OM, such as the first ethmoturbinal (ET I), the range of thickness of OM and non‐OM may overlap. Future work should assess the utility of diffusible iodine‐based contrast enhanced CT techniques, which can differentiate epithelium from the lamina propria, to enhance our ability to differentiate mucosa types on more rostral ethmoturbinals. This is especially critical for structures such as ET I, which have mixed functional roles in many mammals. 

Abstract Living primates show a complex trend in reduction of nasal cavity spaces and structures due to moderate to severe constraint on interorbital breadth. Here we describe the ontogeny of the posterior end of the primate cartilaginous nasal capsule, the thimble shaped posterior nasal cupula (PNC), which surrounds the hind end of the olfactory region. We used a histologically sectioned sample of strepsirrhine primates and two non‐primates (Tupaia belangeri,Rousettus leschenaulti), and histochemical and immunohistochemical methods to study the PNC in a perinatal sample. At birth, most strepsirrhines possess only fragments of PNC, and these lack a perichondrium. Fetal specimens of several species reveal a more complete PNC, but the cartilage exhibits uneven or weak reactivity to type II collagen antibodies. Moreover, there is relatively less matrix than in the septal cartilage, resulting in clustering of chondrocytes, some of which are in direct contact with adjacent connective tissues. In one primate (Vareciaspp.) and both non‐primates, the PNC has a perichondrium at birth. In older, infantVareciaandRousettus, the perichondrium of the PNC is absent, and PNC fragmentation at its posterior pole has occurred in the former. Loss of the perichondrium for the PNC appears to precede resorption of the posterior end of the nasal capsule. These results suggest that the consolidation of the basicranial and facial skeletons happens ontogenetically earlier in primates than other mammals. We hypothesize that early loss of cartilage at the sphenoethmoidal articulation limits chondral mechanisms for nasal complexity, such as interstitial expansion or endochondral ossification. 

Abstract Previous descriptive work on deciduous dentition of primates has focused disproportionately on great apes and humans. To address this bias in the literature, we studied 131 subadult nonhominoid specimens (including 110 newborns) describing deciduous tooth morphology and assessing maximum hydroxyapatite density (MHD). All specimens were CT scanned at 70 kVp and reconstructed at 20.5–39 μm voxels. Grayscale intensity from scans was converted to hydroxyapatite (HA) density (mg HA/cm³) using a linear conversion of grayscale values to calibration standards of known HA density (R²= .99). Using Amira software, mineralized dental tissues were captured by segmenting the tooth cusps first and then capturing the remainder of the teeth at descending thresholds of gray levels. We assessed the relationship of MHD of selected teeth to cranial length using Pearson correlation coefficients. In monkeys, anterior teeth are more mineralized than postcanine teeth. In tarsiers and most lemurs and lorises, postcanine teeth are the most highly mineralized. This suggests that monkeys have a more prolonged process of dental mineralization that begins with incisors and canines, while mineralization of postcanine teeth is delayed. This may in part be a result of relatively late weaning in most anthropoid primates. Results also reveal that in lemurs and lorises, MHD of the mandibular first permanent molar (M₁) negatively correlates with cranial length. In contrast, the MHD of M₁positively correlates with cranial length in monkeys. This supports the hypothesis that natural selection acts independently on dental growth as opposed to mineralization and indicates clear phylogenetic differences among primates. 

Abstract Cranial synchondroses are cartilaginous joints between basicranial bones or between basicranial bones and septal cartilage, and have been implicated as having a potential active role in determining craniofacial form. However, few studies have examined them histologically. Using histological and immunohistochemical methods, we examined all basicranial joints in serial sagittal sections of newborn heads from nine genera of primates (five anthropoids, four strepsirrhines). Each synchondrosis was examined for characteristics of active growth centers, including a zonal distribution of proliferating and hypertrophic chondrocytes, as well as corresponding changes in matrix characteristics (i.e., density and organization of Type II collagen). Results reveal three midline and three bilateral synchondroses possess attributes of active growth centers in all species (sphenooccipital, intrasphenoidal, presphenoseptal). One midline synchondrosis (ethmoseptal) and one bilateral synchondrosis (alibasisphenoidal synchondrosis [ABS]) are active growth centers in some but not all newborn primates. ABS is oriented more anteriorly in monkeys compared to lemurs and bushbabies. The sphenoethmoidal synchondrosis (SES) varies at birth: in monkeys, it is a suture‐like joint (i.e., fibrous tissue between the two bones); however, in strepsirrhines, the jugum sphenoidale is ossified while the mesethmoid remains cartilaginous. No species possesses an SES that has the organization of a growth plate. Overall, our findings demonstrate that only four midline synchondroses have the potential to actively affect basicranial angularity and facial orientation during the perinatal timeframe, while the SES of anthropoids essentially transitions toward a “suture‐like” function, permitting passive growth postnatally. Loss of cartilaginous continuity at SES and reorientation of ABS distinguish monkeys from strepsirrhines.