Evolution of the Dinosauriform Respiratory Apparatus: New Evidence from the Postcranial Axial Skeleton

Examination of the thoracic rib and vertebral anatomy of extant archosaurs indicates a relationship between the postcranial axial skeleton and pulmonary anatomy. Lung ventilation in extant crocodilians is primarily achieved with a hepatic piston pump and costal rotation. The tubercula and capitula of the ribs lie on the horizontal plane, forming a smooth thoracic “ceiling” facilitating movement of the viscera. Although the parietal pleura is anchored to the dorsal thoracic wall, the dorsal visceral pleura exhibits a greater freedom of movement. The air sac system and lungs of birds are associated with bicapitate ribs with a ventrally positioned capitular articulation, generating a rigid and furrowed rib cage that minimizes dorsoventral changes in volume in the dorsal thorax. The thin walled bronchi are kept from collapsing by fusion of the lung to the thorax on all sides. Data from this study suggest a progression from a dorsally rigid, heterogeneously partitioned, multichambered lung in basal dinosauriform archosaurs towards the small entirely rigid avian‐style lung that was likely present in saurischian dinosaurs, consistent with a constant volume cavum pulmonale, thin walled parabronchi, and distinct air sacs. There is no vertebral evidence for a crocodilian hepatic piston pump in any of the taxa reviewed. The evidence for both a rigid lung and unidirectional airflow in dinosauriformes raises the possibility that these animals had a highly efficient lung relative to other Mesozoic vertebrates, which may have contributed to their successful radiation during this time period. Anat Rec, 2011. © 2011 Wiley‐Liss, Inc.     

Vertebral Comparative Anatomy and Morphological Differences in Anguine Lizards With a Special Reference to Pseudopus apodus

The article reports on the first detailed vertebral and rib morphology of anguine taxon Pseudopus apodus using micro-computed tomography. A comparison shows significant morphological differences of vertebrae of Pseudopus relative to those of Anguis and Ophisaurus. Usually, there are 55 presacral vertebrae, two sacral, and 95–97 caudal vertebrae. Pseudopus apodus can be defined by 23 diagnostic features concerning the vertebral column. Although zygapophyseal articulation between atlas and axis is well developed in limbed anguid gerrhonotine lizards like Abronia or Barisia, it is absent in the extant representatives of the clade Anguinae, which are limbless. Thus, our study brings further support to the hypothesis about the complete reduction of this articulation in forms with reduced or absent limbs. Comparison of adult and juvenile morphology of vertebrae of P. apodus was also analyzed. Heterochrony in the evolution of this taxon was previously confirmed by its skull morphology and it can be also documented on the basis of vertebrae. Our data suggest that a peramorphic heterochronic process played a role in the evolution of this largest extant anguine species. Geometric morphometric analyses revealed a pattern of high vertebral disparity among species. We found a clear separation of limbless forms in morphospace. Pseudopus apodus always clusters within Ophisaurus-species confirming molecular and some morphological phylogenies. Only the first tail vertebra shows a distinct difference to those of other anguids, which might be related to altered locomotion associated to the larger body size in this species. Anat Rec, 302:232–257, 2019. © 2018 Wiley Periodicals, Inc.     

Maximum Bony Gape in Primates

Maximum jaw gape has important functional implications for behavior and feeding habits in primates. It has been suggested that gape is correlated to canine height and ingested food size. Extending these correlations to the fossil record would provide insights about the diets and/or social behavior of extinct primates. However, this can be problematic due to uncertainty about size and location of musculature, and it depends on reliability and repeatability of maximum gape estimation using only skeletal elements. In this study, maximum bony gape (MBG) was estimated using reliable landmarks and repeatable methods. The cranium was fixed in position and then the mandible was rotated and translated to the point immediately prior to loss of condyle-glenoid contact. Then it was photographed in a steady position using an adjustable wooden frame. This protocol allowed for photographs and linear measurements to be obtained for many museum specimens in a short time. The sample included 203 individuals, representing 42 species of primates. When scaled for body size, linear MBG correlates with maximum anesthetized gape (Hylander: Am J Phys Anthropol 150 (2013) 247–259), ingested food size (Perry and Hartstone-Rose: Am J Phys Anthropol 142 (2010) 625–635), and canine length but not condylar height. Anat Rec, 302:215–225, 2019. © 2018 Wiley Periodicals, Inc.     

Variation in cross-sectional shape and biomechanical properties of the bat humerus under Wolff's law

Bats use their forelimbs in different ways, but flight is the most notable example of morphological adaptation. Foraging and roosting specializations beyond flight have also been described in several bat lineages. Understanding postcranial evolution during the locomotory and foraging diversification of bats is fundamental to understanding bat evolution. We investigated whether different foraging and roosting behaviors influenced humeral cross-sectional shape and biomechanical variation, following Wolff's law of bone remodeling. The effect of body size and phylogenetic relatedness was also tested, in order to evaluate multiple sources of variation. Our results suggest strong ecological signal and no phylogenetic structuring in shape and biomechanical variation in humeral phenotypes. Decoupled modes of scaling of shape and biomechanical variation were consistently indicated across foraging and roosting behaviors, suggesting divergent allometric trajectories. Terrestrial locomoting and upstand roosting species showed unique patterns of shape and biomechanical variation across all our analyses, suggesting that these rare behaviors among bats place unique functional demands on the humerus, canalizing phenotypes. Our results suggest that complex and multiple adaptive pathways interplay in the postcranium, leading to the decoupling of different features and regions of skeletal elements optimized for different functional demands. Moreover, our results shed further light on the phenotypical diversification of the wing in bats and how adaptations besides flight could have shaped the evolution of the bat postcranium.     

Postsacral Vertebral Morphology in Relation to Tail Length Among Primates and Other Mammals

Tail reduction/loss independently evolved in a number of mammalian lineages, including hominoid primates. One prerequisite to appropriately contextualizing its occurrence and understanding its significance is the ability to track evolutionary changes in tail length throughout the fossil record. However, to date, the bony correlates of tail length variation among living taxa have not been comprehensively examined. This study quantifies postsacral vertebral morphology among living primates and other mammals known to differ in relative tail length (RTL). Linear and angular measurements with known biomechanical significance were collected on the first, mid‐, and transition proximal postsacral vertebrae, and their relationship with RTL was assessed using phylogenetic generalized least‐squares regression methods. Compared to shorter‐tailed primates, longer‐tailed primates possess a greater number of postsacral vertebral features associated with increased proximal tail flexibility (e.g., craniocaudally longer vertebral bodies), increased intervertebral body joint range of motion (e.g., more circularly shaped cranial articular surfaces), and increased leverage of tail musculature (e.g., longer spinous processes). These observations are corroborated by the comparative mammalian sample, which shows that distantly related short‐tailed (e.g., Phascolarctos, Lynx) and long‐tailed (e.g., Dendrolagus, Acinonyx) nonprimate mammals morphologically converge with short‐tailed (e.g., Macaca tonkeana) and long‐tailed (e.g., Macaca fascicularis) primates, respectively. Multivariate models demonstrate that the variables examined account for 70% (all mammals) to 94% (only primates) of the variance in RTL. Results of this study may be used to infer the tail lengths of extinct primates and other mammals, thereby improving our understanding about the evolution of tail reduction/loss. Anat Rec, 298:354–375, 2015. © 2014 Wiley Periodicals, Inc.     

Postcranial skeletal pneumaticity: a case study in the use of quantitative microCT to assess vertebral structure in birds

Limb elements in birds have been characterized as exhibiting a reduction in trabecular bone, thinner cortices and decreased bending strength when pneumatized, yet it is unclear if these characteristics generalize to the axial skeleton. Thin section techniques, the traditional gold standard for bone structure studies, have most commonly been applied to the study of avian bone. This destructive technique, however, makes it subsequently impossible to use the same samples in experimental testing systems that allow researchers to correlate structure with the mechanical properties of the bone. Micro-computed tomography (microCT), a non-destructive X-ray imaging technique, can be used to assess the effect of pneumatization on vertebral cortical and trabecular bone through virtual extraction and structural quantification of each tissue type. We conducted a preliminary investigation of the application of microCT methods to the study of cortical and trabecular bone structure in a small sample of pneumatic and apneumatic thoracic vertebrae. The sample consisted of two similar-sized anatids, Aix sponsa (n = 7) and Oxyura jamaicensis (n = 5). Volumes of interest were created that contoured (outlined) the boundaries of the ventral cortical bone shell, the trabecular compartment and the whole centrum (cortical bone + trabecular bone), and allowed independent structural analysis of each volume of interest. Results indicated that bone volume fraction of the whole centrum was significantly higher in the apneumatic O. jamaicensis than in the pneumatized A. sponsa (A. sponsa = 36%, O. jamaicensis = 48%, P < 0.05). In contrast, trabecular bone volume fraction was similar between the two species. The ventral cortical bone shell was approximately 23% thinner (P < 0.05) in A. sponsa (0.133 mm) compared with apneumatic O. jamaicensis (0.172 mm). This case study demonstrates that microCT is a powerful non-destructive imaging technique that may be applied to the three-dimensional study of avian bone. The preliminary results suggest that pneumatic and apneumatic vertebrae of comparably sized avian species differ in relative bone volume, with the largest difference apparent at the level of the cortex, and not within trabecular bone. The presence of relatively thin cortices in pneumatic vertebrae is consistent with previous studies contrasting diaphyseal cortical bone between pneumatic and apneumatic long bones. Methodological issues related to this and any comparative microCT study of bone structure are discussed.     

Sex and Laterality Differences in Medullary Humerus Morphology

Percutaneous osseointegrated (OI) prosthetic limb attachment holds promise for transhumeral amputees. Understanding humeral medullary morphology is necessary for informed design of upper extremity OI systems, and is beneficial to the field of megaprosthetic reconstruction of the distal humerus where diaphyseal fixation is desired. The purpose of this study was to quantify the sex and laterality differences in humerus morphology, specifically over the diaphysis. Three-dimensional surface reconstructions of 58 pairs of cadaveric humeri (43 male, 15 female) were generated from CT data. Measures describing periosteal and medullary morphology were collected relative to an anatomic coordinate system. Sex and laterality differences in biomechanical length (BML) were observed (P ≤ 0.001 and 0.022, respectively). Head radius was larger in males than females (P ≤ 0.001). Retroversion was increased in right humeri relative to left (P ≤ 0.001). Canal orientation exhibited a conformational shift from anteversion to retroversion distally at approximately 65% BML. Right humeri exhibited larger medullary diameters than left in the 1st and 2nd principal directions (P ≤ 0.024). Males displayed larger diameter medullary canals proximally (P ≤ 0.029) and an increased rate of divergence of the endosteal cortex in the proximal diaphysis (P ≤ 0.009). Females exhibited higher canal aspect ratios at mid-shaft (P ≤ 0.014) and lower mean cortical thickness (P ≤ 0.001). Human humeral diaphysis morphology exhibits sex and laterality differences, which are dependent on position along the diaphysis. Understanding humeral morphology is necessary to achieve adequate primary stability and bone apposition in design of endoprosthetic stems for percutaneous OI implants, and distal humerus replacement. Anat Rec, 302:1709–1717, 2019. © 2019 The Authors. The Anatomical Record published by Wiley Periodicals, Inc. on behalf of American Association for Anatomy     

Differences in acetabular morphology related to side and sex in a Chinese population

The purpose of the present study was to determine the side and sex differences in acetabular dimension and orientation in normal Chinese adults, which are not known well but are important in hip joint replacement surgery. The acetabular parameters, including anteversion angle (AV.A), abduction angle (AB.A), center edge angle (CE.A), acetabular width (Ac.W) and acetabular depth (Ac.D), were measured on CT images in 100 healthy Chinese adults. The acetabular index of depth to width (Ac.D/Ac.W) was calculated by depth/width × 100. Percentage side difference (PSD) was calculated for each parameter. The absolute numbers of PSD (aPSD) were compared between the groups of right positive (right PSD > left PSD) and left positive (left PSD > right PSD) groups. There was no significant side difference in any of the parameters. Compared to men, a significant increase in AB.A (P = 0.001) and significant decreases in Ac.W (P < 0.001), Ac.D (P < 0.001) and Ac.D/Ac.W (P < 0.05) occurred in women. The differences in Ac.W and Ac.D became insignificant when adjusted for body height. aPSD did not show a significant difference between right and left positive groups in both sexes. In conclusion, the side differences in acetabular parameters in a normal individual are likely to be associated with measurement error. In addition, the larger acetabular dimension in men is attributed to greater body height independent of sex.     

Postcranial Pneumaticity and Bone Structure in Two Clades of Neognath Birds

Most living birds exhibit some degree of postcranial skeletal pneumaticity, aeration of the postcranial skeleton by pulmonary air sacs and/or directly from the lungs. The extent of pneumaticity varies greatly, ranging from taxa that are completely apneumatic to those with air filling most of the postcranial skeleton. This study examined the influence of skeletal pneumatization on bone structural parameters in a sample of two size‐ and foraging‐style diverse (e.g., subsurface diving vs. soaring specialists) clades of neognath birds (charadriiforms and pelecaniforms). Cortical bone thickness and trabecular bone volume fraction were assessed in one cervical and one thoracic vertebra in each of three pelecaniform and four charadriiform species. Results for pelecaniforms indicate that specialized subsurface dive foragers (e.g., the apneumatic anhinga) have thicker cortical bone and a higher trabecular bone volume fraction than their non‐diving clademates. Conversely, the large‐bodied, extremely pneumatic brown pelican (Pelecanus occidentalis) exhibits thinner cortical bone and a lower trabecular bone volume fraction. Such patterns in bone structural parameters are here interpreted to pertain to decreased buoyancy in birds specialized in subsurface dive foraging and decreased skeletal density (at the whole bone level) in birds of larger body size. The potential to differentially pneumatize the postcranial skeleton and alter bone structure may have played a role in relaxing constraints on body size evolution and/or habitat exploitation during the course of avian evolution. Notably, similar patterns were not observed within the equally diverse charadriiforms, suggesting that the relationship between pneumaticity and bone structure is variable among different clades of neognath birds. Anat Rec, 296:867–876, 2013. © 2013 Wiley Periodicals, Inc.     

Development of the pelvis and posterior part of the vertebral column in the Anura

The anuran pelvic girdle is unique among all amphibians in that its acetabular portion is located far posterior to the sacrum, lateral to the postsacral (= caudal) vertebral column, which is reduced to a single rod-like element called the urostyle. This situation in the adult is strikingly different not only from that in ancestral temnospondyls but also in other modern amphibians. Because there is no fossil that would document this evolutionary anatomical modification except for Triadobatrachus, the only data may be inferred from development in modern anurans. We chose seven anuran species (belonging to the genera Discoglossus, Bombina, Pelobates, Bufo, Rana and Xenopus), representing the principal locomotory types (saltation, swimming, crawling and burrowing). Development of the pelvic girdle was studied on cleared and stained whole mounts and partly on serial histological sections. The basic developmental pattern was similar in all species: the pelvis on both sides develops from two centres (puboischiadic and iliac, respectively). The ilium then extends vertically towards the sacral vertebra and later rotates posteriorly so that ultimately the acetabulum is lateral to the tail (= urostyle). Only minor deviations from this pattern were found, mainly associated with differences in water and terrestrial dwelling.     

Serum Leptin and Skeletal Differences between Obese and Non-Obese Patients with Chronic Obstructive Pulmonary Disease

ObjectiveChronic obstructive pulmonary disease (COPD) affects body composition, adipokine secretion, and skeletal integrity. The aim was to determine the association between leptin, body mass (BM) and body composition parameters - fat mass (FM) and fat mass index (FMI), lean tissue mass (LTM), lean tissue mass index (LTMI) and bone mineral density (BMD) in 67 male COPD patients.MethodsBM, body composition and biochemical indicators were measured or calculated using standard methods. Data were analyzed according to groups: non-obese (N = 48, BMI 21.0-29.9 kg/m²) and obese (N = 19, BMI ≥ 30.0 kg/m²).ResultsIn the non-obese group statistically significant correlations were observed: negative ones of age with most BMD T scores, positive ones of BMI with all T scores, FM, FMI, LTMI and leptin, of FMI with leptin and all T scores, and of LTMI with most T scores. In the obese group also statistically significant correlations were found: positive ones of BMI with FMI, LTM, leptin and T scores (trochanter, total hip); of FMI with leptin; and of leptin with total hip T score.ConclusionA positive relationship between FMI and BMD was found only in non-obese but not in obese COPD patients. Leptin concentration was associated positively with the total hip T score only in obese COPD patients, suggesting its protective role on the skeleton of obese COPD patients.Key Words: Body mass, Body mass index, Fat mass, Fat mass index, Lean tissue mass, Lean tissue mass index, 25-OH vitamin D, Leptin, Bone mineral density     

Differences in morphology and force/velocity relationship between Senegalese and Italian sprinters

In order to investigate whether the supremacy of African sprinters is related to the leg extensor force/velocity relationship or to leg morphology, two groups of elite sprinters originating respectively from Senegal (S) and Italy (I) were compared in this respect. The groups included 13 S and 15 I male sprinters. Their mean best performances over 100 m during the preceding track and field season were 10.66 (0.3) and 10.61 (0.3) s (NS), respectively. Age, height and mass were similar in the two groups. The force/velocity relationship of the leg extensors was assessed during maximal half-squats on a guided horizontal barbell with masses of 20–140 kg added on the shoulders. Leg morphology was assessed by relating the sub-ischial length to the standing height (L/H) and by measuring the inertia in the vertical (I _Z in kg.cm²), antero-posterior (I _Y, kg.cm²) and medio-lateral (I _X, kg.m²) planes. The two groups developed non-different force and power when lifting the heaviest loads. Inversely, the lighter the load, the lower the force and power developed by S, as compared to I (P<0.001). S demonstrated greater L/H (P<0.001), and 26% lower I _Z (P<0.01), 15% lower I _Y (P=0.09), and 14% lower I _X (P=0.10). These results suggest that S and I sprinters were similar as regards the muscle abilities involved in slow maximal contractions. However, S demonstrated lower values in muscle abilities related to high-speed contractions, suggesting that S sprinters had a lower percentage of fast twitch fibres. This is likely to be compensated for by the lower level of internal work due to longer and lighter legs.     

Skeletal morphology and development of the olfactory region of Spea (Anura: Scaphiopodidae)

The nasal capsules of anurans are formed by an intricate set of sac-like cavities that house the olfactory organ and constitute the beginning of the respiratory system. In tadpoles, nasal capsules do not have a respiratory function, but each is composed of a single soft tissue cavity lined with olfactory epithelium. Our study has revealed that in Spea the nasal cartilages and septomaxillae are de novo adult structures that form dorsal to the larval skeleton of the ethmoid region. The only element of the adult nasal capsule that is partially derived from the larval skeleton is the solum nasi. Development of the nasal skeleton begins at about Gosner Stage 31, with chondrification of the septum nasi and lamina orbitonasalis. The alary cartilage and superior prenasal cartilage are the first of the anterior nasal cartilages to chondrify at Gosner Stage 37. By Gosner Stages 40/41, the ethmoid region is composed of the larval structures ventrally and the adult structures dorsally. By Stage 44, the larval structures have eroded. The adult nasal capsule is characterized by: (1) a septum nasi that projects ventrally beyond the plane of the nasal floor; (2) a paranasal commissure that forms the ventral margin of the fenestra nasolateralis; and (3) a large skeletal support for the eminentia olfactoria formed by the nasal floor and vomer. The timing of chondrification of the anterior nasal cartilages and the development of the postnasal wall, inferior prenasal cartilage, fenestra nasolateralis, and paranasal commissure are discussed and compared with those of other anuran species. This study also includes a discussion of the morphology of the skeletal support for the eminentia olfactoria, a structure best developed in distinctly ground-dwelling frogs such as spadefoot toads. Finally, we propose a more precise restriction of the terminology that is used to designate the posterior structures of the olfactory region of anurans.     

Localization of Papillofoveal Bundles in Primates

Axons in the fovea are precisely organized to ensure accurate vision. We investigated the morphologic characteristics and localization of nerve bundles in the optic nerve in primates. Macaque eyes were studied for conventional and immunostaining, and also marmoset eyes for carbocyanine dye tracing. Locally confined lesions associated with similar findings to human age‐related macular degeneration (ARMD) were also evaluated. Axons of retinal ganglion cells formed fasciculi near their origin, and these fasciculi formed bundles thereafter. In the retinal nerve fiber layer, ascending bundles assembled stratification adding proximal bundle underneath successively. Bundles in the arcuate zone displayed a characteristic fine, parallel arrangement, whereas those in the outside zone intermingled with undefined reticular bundles as they approached the optic nerve head. Macular bundles remained in groups and were distributed in the temporal wedge of the optic nerve head. Orthograde and retrograde tracing revealed that these bundles formed confined groups of various sizes and, ultimately, a specific group of small bundles located in the innermost row, near the central vessels. In addition, these bundles showed evidence of focal degenerative deterioration in eyes with ARMD. Papillomacular bundles have a characteristic alignment and configuration. Foveal bundles that compose the confined group closest to the optic trunk (which we term papillofoveal bundles) appear to have functional significance with respect to the isolated lesions that accompany central vision loss or preservation. Anat Rec, 2012. © 2011 Wiley Periodicals, Inc.