Structure and growth pattern of the bizarre hemispheric prominence on the rostrum of the fossil beaked whale Globicetus hiberus (Mammalia,Cetacea, Ziphiidae)

The rostrum of most ziphiids (beaked whales) displays bizarre swollen regions, accompanied with extreme hypermineralisation and an alteration of the collagenous mesh of the bone. The functional significance of this specialization remains obscure. With the voluminous and dense hemispheric excrescence protruding from the premaxillae, the recently described fossil ziphiid Globicetus hiberus is the most spectacular case. This study describes the histological structure and interprets the growth pattern of this unique feature. Histologically, the prominence in Globicetus is made up of an atypical fibro‐lamellar complex displaying an irregular laminar organization and extreme compactness (osteosclerosis). Its development is suggested to have resulted from a protraction of periosteal accretion over the premaxillae, long after the end of somatic growth. Complex shifts in the geometry of this tissue are likely to have occurred during its accretion and no indication of Haversian remodeling could be found. X‐ray diffraction and Raman spectroscopy indicate that the bone matrix in the premaxillary prominence of Globicetus closely resembles that of the rostrum of the extant beaked whale Mesoplodon densirostris: apatite crystals are of common size and strongly oriented, but the collagenous meshwork within bone matrix seems to be extremely sparse. These morphological and structural data are discussed in the light of functional interpretations proposed for the highly unusual and diverse ziphiid rostrum. J. Morphol. 277:1292–1308, 2016. © 2016 Wiley Periodicals, Inc.     

On how the periosteal bone of the delphinid humerus becomes cancellous: ontogeny of a histological specialization

In cetaceans, the bones of the flippers lack a free medullary cavity and have a cancellous texture, with compact cortices reduced or absent. The present work discusses the ontogenetic basis of these characters in terms of the ontogeny of the structure and textural bone compactness (TBC) of the humeral diaphysis in a growth series of common dolphins (Delphinus delphis). The texture of the primary periosteal deposits is compact; soon after their accretion, the deposits undergo an extensive erosion that turns them into a cancellous tissue. A diffuse endosteal front of resorption expands in parallel with the growth of the cortex and acts as small units scattered within the cortices. Starting soon after birth and continuing throughout the life of the animals, the compactness of the periosteal cortex decreases at both general and local levels. This trend correlates strongly with the increase in size of the diaphyseal section and reflects the fact that relatively more bone is eroded than deposited during growth in the cancellous parts of the cortex. In the broad sense, this is basically an osteoporotic process, which is not identical, however, to senile or disuse osteoporoses.     

Inner architecture of vertebral centra in terrestrial and aquatic mammals: A two‐dimensional comparative study

Inner vertebral architecture is poorly known, except in human and laboratory animals. In order to document this topic at a broad comparative level, a 2D‐histomorphometric study of vertebral centra was conducted in a sample of 98 therian mammal species, spanning most of the size range and representing the main locomotor adaptations known in therian taxa. Eleven variables relative to the development and geometry of trabecular networks were extracted from CT scan mid‐sagittal sections. Phylogeny‐informed statistical tests were used to reveal the respective influences of phylogeny, size, and locomotion adaptations on mammalian vertebral structure. The use of random taxon reshuffling and squared change parsimony reveals that 9 of the 11 characteristics (the two exceptions are total sectional area and structural polarization) contain a phylogenetic signal. Linear discriminant analyses suggest that the sampled taxa can be arranged into three categories with respect to locomotion mode: a) terrestrial + flying + digging + amphibious forms, b) coastal oscillatory aquatic taxa, and c) pelagic oscillatory aquatic forms represented by oceanic cetaceans. Pairwise comparison tests and linear regressions show that, when specific size increases, the length of trabecular network (Tt.Tb.Le), as well as trabecular proliferation in total sections (Pr.Tb.Tt), increase with positive allometry. This process occurs in all locomotion categories but is particularly pronounced in pelagic oscillators. Conversely, mean trabecular width has a lesser increase with size in pelagic oscillators. Trabecular orientation is not influenced by size. All tests were corrected for multiple testing. By using six structural variables or indices, locomotion mode can be predicted with a 97.4% success rate for terrestrial forms, 66.7% for coastal oscillatory, and 81.3% for pelagic oscillatory. The possible functional meaning of these results and their potential use for paleobiological inference of locomotion in extinct taxa are discussed. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

Evolution of Sirenian Pachyosteosclerosis,a Model-case for the Study of Bone Structure in Aquatic Tetrapods

Osteosclerosis, or inner bone compaction, and pachyostosis, or outer hyperplasy of bone cortices (swollen bones), are typical features of tetrapods secondarily adapted to life in water. These peculiarities are spectacularly exemplified by the ribs of extant and extinct Sirenia. Sea cows are thus the best model for studying this kind of bone structural specializations. In order to document how these features differentiated during sirenian evolution, the ribs of 15 species, from the most basal form (Pezosiren portelli) up to extant taxa, were studied, and compared to those of other mammalian species from both morphometric and histological points of view. Pachyostosis was the first of these two specializations to occur, by the middle of the Eocene, and is a basal feature of the Sirenia. However, it subsequently regressed in some taxa that do not exhibit hyperplasic rib cortices. Osteosclerosis was only incipient in P. portelli. Its full development occurred later, by the end of the Eocene. These two structural specializations of bone are variably pronounced in extinct and extant sirenians, and relatively independent from each other, although frequently associated. They are possibly due to similar heterochronic mechanisms bearing on the timing of osteoblast activity. These results are discussed with respect to the functional constraints of locomotion in water.     

A survey of osteoderm histology and ornamentation among Crocodylomorpha: A new proxy to infer lifestyle?

Osteoderms of eight extant and extinct species of crocodylomorphs are studied histologically and morphologically. Most osteoderms display the typical “crocodilian” structure with a woven-fibered matrix surrounded by an upper and a lower parallel fibered matrix. The dorsal ornamentation of those specimens consists of a pit-and-ridge structure, with corresponding remodeling mechanisms. However, an osteoderm of Iberosuchus, studied here for the first time, differs in being nearly devoid of ornamentation; moreover, it shows strong bundles of straight Sharpey's fibers perpendicular to the surface in its lateral and dorsal walls, along with a rough plywood-like structure in its basal plate. This suggests that this osteoderm was more deeply anchored within the dermis than the other osteoderms studied hitherto. This peculiar structure might have been linked to a terrestrial ecology and a specific thermoregulation strategy. Some other notosuchians in our sample do not exhibit ornamentation on their osteoderms, as opposed to neosuchians. Considering current interpretations of osteoderm function(s) in crocodilians, our observations are discussed in reference to possible ecophysiological peculiarities of Notosuchia in general, and Iberosuchus in particular.     

Ontogenetic development of skeletal weight in a small delphinid,Delphinus delphis (Cetacea,Odontoceti)

Summary Total body weight and skeletal weight, from clean and dry bones, were measured in 61 common dolphins, Delphinus delphis of known ages (age determinations using both growth lines in the jaw, and growth layer groups in dentine). The relative skeletal weight in adults, expressed as a percentage of total body weight, is notably less than that of any cetaceans previously described. Appearently, this ratio is also smaller than that for any other vertebrate. In particular, skeletons of delphinids tend to be proportionally lighter than those of bony fishes. Classical hypotheses concerning the effect of buoyancy on the mass of supportive tissues can only partially explain such a relative reduction of skeletal weight.     

Microanatomical diversity of amniote ribs: an exploratory quantitative study

Bone microanatomical diversity in extant and extinct tetrapods has been studied extensively, using increasingly sophisticated quantitative methods to assess its ecological, biomechanical and phylogenetic significance. Most studies have been conducted on the appendicular skeleton, and a strong relationship was found between limb bone microanatomy and habitat preferences. Few comparative studies have focused on the microanatomy of the axial skeleton and its ecological signal. In the present study, we propose the first exploratory study of the microanatomical diversity of amniote ribs. Our comparative sample comprises 155 species of extant amniotes and encompasses the taxonomic, ecological, and body size diversity of this group. We standardized our sampling location to the midshaft of mid‐dorsal ribs. Transverse sections were obtained from classical petrographic methods, as well as by X‐ray microtomography. Most of the microanatomical and size characters of the ribs display a phylogenetic signal, which is an expected result and is also observed in amniote limb bones and vertebrae. We found a significant relationship between rib cortical thickness, global compactness, and lifestyle. As for the vertebrae, the development of the spongiosa in the medullary region appears to be strongly correlated with size. Even though an ecological signal was found in the inner structure of the ribs, additional work is needed to document the intra‐individual variability of the rib microanatomy along the rib cage and within a single element.     

Structure and Growth Pattern of Pseudoteeth in Pelagornis mauretanicus (Aves,Odontopterygiformes, Pelagornithidae)

The extinct Odontopterygiformes are the sole birds known to possess strong and sharp bony pseudoteeth, the shape and location of which are closely mimetic of real teeth. The structure of the pseudoteeth is investigated here in a late Pliocene/early Pleistocene species, Pelagornis mauretanicus, using X-ray microtomography and thin sections. The results are interpreted with regard to the pseudotooth mode of growth, and have implications concerning aspects of Pelagornis ecology. The larger pseudoteeth are hollow and approximately cone-shaped, and the smaller ones are rostro-caudally constricted. The walls of pseudoteeth are composed of bone tissue of the fibro-lamellar type, which is intensively remodeled by Haversian substitution. The jaw bones display the same structure as the pseudoteeth, but their vascular canals are oriented parallel to the long axis of the bones, whereas they are perpendicular to this direction in the pseudoteeth. There is no hiatus or evidence of a fusion between the pseudoteeth and the jaw bones. Two possible models for pseudotooth growth are derived from the histological data. The most plausible model is that pseudotooth growth began after the completion of jaw bone growth, as a simple local protraction of periosteal osteogenic activity. Pseudotooth development thus occurred relatively late during ontogeny. The highly vascularized structure and the relative abundance of parallel-fibered bone tissue in the pseudoteeth suggest poor mechanical capabilities. The pseudoteeth were most likely covered and protected by the hardened, keratinized rhamphotheca in the adult during life. The late development of the pseudoteeth would involve a similarly late and/or partial hardening of the rhamphotheca, as displayed by extant Anseriformes, Apterygiformes and some Charadriiformes. This would add support to the hypothesis of a close phylogenetic relationship between Odontopterygiformes and Anseriformes. The late maturation of the Pelagornis feeding apparatus, and hence the delayed capability for efficient prey catching, suggests that Pelagornis was altricial.     

Microanatomy of the amniote femur and inference of lifestyle in limbed vertebrates

The femoral microanatomy of 155 species of extant amniotes (57 species of mammals, 15 species of turtles, 56 species of lepidosaurs, and 27 species of birds) of known lifestyle is studied to demonstrate a possible link between some basic parameters of bone structure and specific lifestyles, as well as phylogenetic relationships between taxa. Squared change parsimony with random taxon reshuffling and pairwise comparisons reveal that most compactness and size parameters exhibit both phylogenetic and ecological signals. A discriminant analysis produces several inference models, including a ternary model (aquatic, amphibious, terrestrial) that yield the correct lifestyle in 88% of the cases. These models are used to infer the lifestyle of three extinct Permian temnospondyls: Eryops megacephalus, Acheloma dunni, and Trimerorhachis insignis. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 644–655.     

The histological structure of glyptosaurine osteoderms (Squamata: Anguidae), and the problem of osteoderm development in squamates

Glyptosaurinae, a fossil clade of anguid lizards, possess robust osteoderms, with granular ornamentation. In this study, the structural and histological features of these osteoderms were described in order to reconstruct their developmental pattern and further document the degree of homology that could exist between vertebrate integumentary skeletons. Glyptosaurine osteoderms have a diploe architecture and display an unusually complex structure that includes four tissue types: a core of woven‐fibered bone intensely remodeled; a peripheral formation of the same tissue containing dense bundles of long Sharpey fibers; a thick basal layer of lamellar bone; and a superficial layer of a non‐osseous material that belongs to the category of hypermineralized tissues such as ganoine, or enameloid and enamel tissues. The growth pattern of glyptosaurine osteoderms involved appositional processes due to osteoblast activity. In early growth stages, osseous metaplasia might have also been involved, but this possibility is not substantiated by histological observations. The superficial layer of the osteoderms must have resulted from epidermal contribution, a conclusion that would support previous hypotheses on the role of epidermal‐dermal interactions in the formation of squamate osteoderms. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.