Microanatomical and Histological Features in the Long Bones of Mosasaurine Mosasaurs (Reptilia,Squamata) – Implications for Aquatic Adaptation and Growth Rates

Background

During their evolution in the Late Cretaceous, mosasauroids attained a worldwide distribution, accompanied by a marked increase in body size and open ocean adaptations. This transition from land-dwellers to highly marine-adapted forms is readily apparent not only at the gross anatomic level but also in their inner bone architecture, which underwent profound modifications.

Methodology/Principal Findings

The present contribution describes, both qualitatively and quantitatively, the internal organization (microanatomy) and tissue types and characteristics (histology) of propodial and epipodial bones in one lineage of mosasauroids; i.e., the subfamily Mosasaurinae. By using microanatomical and histological data from limb bones in combination with recently acquired knowledge on the inner structure of ribs and vertebrae, and through comparisons with extant squamates and semi-aquatic to fully marine amniotes, we infer possible implications on mosasaurine evolution, aquatic adaptation, growth rates, and basal metabolic rates. Notably, we observe the occurrence of an unusual type of parallel-fibered bone, with large and randomly shaped osteocyte lacunae (otherwise typical of fibrous bone) and particular microanatomical features in Dallasaurus, which displays, rather than a spongious inner organization, bone mass increase in its humeri and a tubular organization in its femora and ribs.

Conclusions/Significance

The dominance of an unusual type of parallel-fibered bone suggests growth rates and, by extension, basal metabolic rates intermediate between that of the extant leatherback turtle, Dermochelys, and those suggested for plesiosaur and ichthyosaur reptiles. Moreover, the microanatomical features of the relatively primitive genus Dallasaurus differ from those of more derived mosasaurines, indicating an intermediate stage of adaptation for a marine existence. The more complete image of the various microanatomical trends observed in mosasaurine skeletal elements supports the evolutionary convergence between this lineage of secondarily aquatically adapted squamates and cetaceans in the ecological transition from a coastal to a pelagic lifestyle.     

The osteohistological variability in the evolution of basal avialans

Basal avialans have been the focus of numerous histological studies in the past decade, from which different osteohistological patterns have been described. In this review, we look at the osteohistology in selected specimens from the four major avian groups: the long-tailed Avialae (Archaeopteryx and Jeholornithiformes), basal Pygostylia, Enantiornithes and Euornithes. Developmental and evolutionary changes in the three major bone layers are observed throughout the bone cortex of the limbs, may it be interspecific or intraspecific. Most noteworthy is the adaptive change from the overall lamellar/parallel-fibered bone tissue to a fibrolamellar complex in the mid-cortex as of the basal Pygostylia, potentially even as of the Jeholornithiformes. This change is generally associated with an increase in the density and complexity of the neurovascular network. Another evolutionary-developmental feature is the progressive loss of post-natal growth marks as of the non-ornithurine Euornithes, indicative of uninterrupted bone growth as observed in extant Neornithes. Our comparisons of the osteohistological patterns allow us to better determine how and when specific features typical observed in the avian crown group developed, associated with external and internal factors, and how they lead to what is commonly observed in extant Neornithes.     

The palaeohistology of the basal ichthyosaur Mixosaurus (Ichthyopterygia,Mixosauridae) from the Middle Triassic: Palaeobiological implications

Here, we provide the first bone histological examination of an ontogenetic series of the basal ichthyosaur Mixosaurus encompassing postnatal to large adult specimens. Growth marks are present in sampled humeri, a femur, a fibula, as well as in other skeletal elements (gastral ribs). Ontogenetic changes are traceable throughout stylo- and zeugopodial development, but interior remodelling and resorption deleted part of the internal growth record in the primary cortex. Mixosaurus humeri started as flat structures consisting of a core of endochondral woven bone and residual calcified cartilage, whereas growth continued by deposition of periosteal fibrolamellar and parallel-fibred bone. Unlike the fast-growing post-Triassic ichthyosaurs that lack growth marks, microstructural and life history data are now becoming available for a basal ichthyosaur. The high growth rate of Mixosaurus may indicate that higher metabolic rates characterised small, non-thunniform ichthyosaurs, as had been suggested already for post-Triassic, cruising forms.     

Unusual histology and morphology of the ribs of mosasaurs (Squamata)

We report the presence of two previously unrecognized features in the dorsal ribs of mosasaurs: first, the presence of extremely dense, pervasive extrinsic fibres (anchoring soft tissue to bone, sometimes called Sharpey's fibres); and second, high intraspecific variation in costal bone compactness. Extensive extrinsic fibres are developed in the dorsal ribs of the mosasaurs Tylosaurus proriger and Eonatator sternbergi. The dorsal ribs of these mosasaurs are also characterized by a longitudinally ridged texture that almost completely covers the bone. Pervasive extrinsic fibres and ridged textures are absent in the mosasaur Selmasaurus russelli as well as the dorsal ribs of extant semi‐aquatic reptiles (e.g. crocodyliforms) and mosasaurs' close extant relatives and analogues (e.g. snakes and varanids). Similar ridged textures characterize the dorsal ribs of several other mosasaur taxa but are developed to a lesser extent (e.g. Mosasaurus, Clidastes, Platecarpus and Ectenosaurus), but in no other taxa have pervasive extrinsic fibres been reported. We interpret these osteohistological features in T. proriger and E. sternbergi as evidence of tendinous attachment of extensive and highly differentiated axial musculature capable of producing great stresses, most likely related to stabilization of the trunk relative to contralateral movements of the tail during carangiform locomotion. We also report the compactness indices (percentage of space occupied by bone rather than cavities) for these large mosasaur ribs, which are much higher than previously reported. This suggests high intraspecific variation in bone compactness that complicates its use in reconstructing mosasaur palaeoecology.     

Forearm bone histology of the small theropod Daliansaurus liaoningensis (Paraves: Troodontidae) from the Yixian Formation,Liaoning, China

We present the first histological analysis of forelimb bones in a troodontid dinosaur, Daliansaurus liaoningensis, from the Early Cretaceous of China using osteohistological thin-sectioning and high-resolution synchrotron-based imaging. The thin wall compacta consists of primary bone, and three lines of arrested growth (LAG) in the radius (R) and two in the ulna (U) divide these into successive zones. Results show that the new fossil has four distinct bone depositional rates: (1) fastest deposition in inner zones R1 and U1 (fibro-lamellar bone with a plexiform-like vasculature); (2) slowed deposition in outer zones R1 and U1 + U2 (loss of vascular density and plexiform component); (3) fluctuating rates of deposition in zones R2 + R3 and in the inner zone U3 (alternating bands of circumferentially organised primary osteons and avascular bone); and (4) slowest deposition in zone R4 and the outer zone U3 (lamellar bone constituting the external fundamental system). Collectively, these growth characteristics suggest that the fossil is an individual that passed the exponential growth phase by the first year, and perished three years later. We conclude that the histology is consistent with an interpretation of this specimen as a late maturing individual that had not yet attained maximum somatic size.     

Bone histology of Iberosuchus macrodon (Sebecosuchia,Crocodylomorpha)

Iberosuchus macrodon is a Cenozoic crocodyliform interpreted as a terrestrial, cursorial form. To assess whether this adaptation was accompanied by a high growth rate and an elevated resting metabolic rate (two features commonly attributed to several terrestrial Triassic Crocodylomorpha based on histology), we studied bone histology in the femora of two specimens attributed to I. macrodon. Beyond this question is the broader problem of the possible survival to the Cretaceous‐Palaeogene extinction event of tachymetabolic sauropsids other than birds. At mid‐diaphysis, bone cortices in Iberosuchus are made of a parallel‐fibred tissue that turns locally to true lamellar bone. Cortical vascularization consists of simple longitudinal canals forming a network of medium density. The spacing pattern of conspicuous lines of arrested growth suggests asymptotic growth for Iberosuchus. This general histological structure prevails also in the metaphyseal region of the bones. It is basically similar to that encountered in certain large lizards adapted to active predation, the Varanidae and the Teidae. In one of the two Iberosuchus femora, however, an intracortical meniscus made of a tissue displaying a global radial architecture occurs in the region of the fourth trochanter. Histologically, the latter can be interpreted either as compacted spongiosa or as a fibro‐lamellar complex with a gross radial orientation, a tissue corresponding to fast periosteal apposition. These observations suggest that Iberosuchus basically had a slow, cyclical growth indicative of an ecto‐poikilothermic, lizard‐like, resting metabolic rate. However, it might also have retained a limited capacity for fast periosteal accretion in relation to local morphogenetic requirements as, for instance, the development of crests or trochanters.     

LYSTROSAURUS MURRAYI (THERAPSIDA, DICYNODONTIA): BONE HISTOLOGY, GROWTH AND LIFESTYLE ADAPTATIONS

Abstract:  Examination of the bone microstructure of Lystrosaurus murrayi from India and South Africa reveals a predominance of fibrolamellar bone tissue, which suggests rapid periosteal osteogenesis and an overall fast growth. Four distinct ontogenetic stages have been identified based on tissue type, organization of the primary osteons, incidence of growth rings, secondary reconstruction and endosteal bone deposition. An indeterminate growth strategy is proposed for Lystrosaurus . Inter-elemental histovariability suggests differential growth rate of the skeletal elements within the same individual, and among different individuals. The high cortical thickness of the dorsal ribs, an extensive secondary reconstruction in the cortical region of different skeletal elements that resulted in erosionally enlarged channels from the perimedullary to the midcortical region, and trabecular infilling of the medullary region even in the diaphyseal sections of the limb bones suggest at least a semi-aquatic lifestyle for L. murrayi .     

Effects of hatchery rearing on Florida largemouth bass Micropterus floridanus resource allocation and performance under semi‐natural conditions

This study examined the growth, activity, metabolism and post‐release survival of three groups of Florida largemouth bass Micropterus floridanus: wild‐caught fish, hatchery fish reared according to standard practice (hatchery standard) and hatchery fish reared under reduced and unpredictable food provisioning (hatchery manipulated). Hatchery‐standard fish differed from wild‐caught fish in all measured variables, including survival in semi‐natural ponds. Hatchery‐standard and hatchery‐manipulated fish showed higher activity levels, faster growth and lower standard metabolic rates than wild‐caught fish in the hatchery. Fish reared under the manipulated feeding regime showed increased metabolic rates and increased post‐release growth, similar to wild‐caught fish. Their activity levels and post‐release survival, however, remained similar to those of hatchery‐standard fish. Activity was negatively correlated with post‐release survival and failure of the feed manipulation to reduce activity may have contributed to its failure to improve post‐release survival. Activity and post‐release survival may be influenced by characteristics of the rearing environment other than the feeding regime, such as stock density or water flow rates.     

Palaeohistology of the bones of pterosaurs (Reptilia: Archosauria): anatomy, ontogeny, and biomechanical implications

Thin sections from long bones of specimens representing pterosaurs ranging from the Early Jurassic to the latest Cretaceous provide a profile of bone histology across a range of sizes, skeletal elements, growth stages, and phylogenetic positions. Most pterosaur bone is fibro-lamellar, organized in an unusual way that suggests high growth rates through ontogeny. Fibro-lamellar deposits are finished by a relatively abrupt deceleration or cessation of growth represented by lamellar, poorly vascularized subperiosteal bone in what appear to be adults. Pterosaurs had the thinnest bone walls of any tetrapods; they complemented high rates of periosteal deposition with almost equally high rates of endosteal erosion. Pterosaurs show a great variety of histologic features that include articular calcified cartilage, sub-chondral bone plates, trabecular bone struts and related internal supports, and secondary deposition and remodeling of bone. They remodeled their bones internally by (1) depositing endosteal bone coatings on the inner cortex and over struts of pre-existing internal bone, (2) secondarily filling bone spaces, and (3) Haversian reworking. The construction of these struts reflects both developmental patterns of bone construction and biomechanical function. Alternating plywood-like layers of bone, heretofore undescribed in tetrapods, provided strength, as did the obliquely oriented system of reticular blood vessels in the bones. The distribution and ontogenetic features of pterosaur bone tissues, when combined with other evidence, suggest generally high growth rates, high metabolic levels, altricial birth, and extended parental care.     

Synchrotron microtomography-based osteohistology of Gansus yumenensis: new data on the evolution of uninterrupted bone deposition in basal birds

Many studies of the limb bones from birds of the major clades reveal a mosaic evolution in morphological characters. From this, we assume that uninterrupted compact bone evolved independently multiple times outside of the crown group. We hypothesise that there are key intraskeletal changes in the osteohistological features, such as the organisation of the vascular network. To test these hypotheses, we analysed and described the osteohistological features of five different midshaft samples of Gansus yumenensis, a non-ornithurine Euornithes from China, based on virtual models obtained from synchrotron microtomography scans, a less invasive method that the traditional physical cross section. We performed quantitative analyses with volume, surface area and estimated ratios. The osteohistological features of Gansus yumenensis were compared with those of stem and crown birds. From our analyses, we discuss the pros/cons of using synchrotron microtomography scans compared to traditional physical cross section. Our analyses demonstrate that Gansus yumenensis is the fourth described extinct Euornithes to exhibit uninterrupted bone deposition in all bone samples, providing further support for multiple origins of this feature outside of the bird crown group. Finally, our osteohistological investigation of Gansus yumenensis provides future study avenues regarding the evolution and development of bone tissue in fossil birds.     

New insights from bone microanatomy of the Late Triassic Hyperodapedon (Archosauromorpha,Rhynchosauria): implications for archosauromorph growth strategy

Bone microanatomy of multiple postcranial skeletal elements of several individuals of Hyperodapedon collected from India is reported. This reveals that fibrolamellar bone tissue is predominant in the mid‐ and inner cortices, whereas the peripheral region of the cortex is composed of either parallel‐fibred and/or lamellar bone. The pattern of primary osteons mostly ranges between laminar and subplexiform. Such predominance of fibrolamellar bone tissue in the cortex suggests an overall fast growth, which slowed down considerably later in ontogeny. Four distinct ontogenetic stages are identified based on the bone microstructure. During the juvenile stage, growth was fast and continuous, but it became punctuated during the early and late sub‐adult stages. In adult individuals, growth was slow and showed periodic interruption but did not stop completely, suggesting that Hyperodapedon had an indeterminate growth strategy. Interelemental histovariations affecting cortical thickness, organization of the vascular network, incidence of growth rings and extent of secondary reconstruction are noted. Throughout ontogeny, the femora show higher cortical thickness than humeri and tibiae, suggesting differential appositional growth rate between the skeletal elements. Differences in cortical thickness are noted in the ribs, which suggest differential functional constraints based on anatomical site‐specific occurrences. Although fibrolamellar bone tissue became progressively more dominant towards the archosaurs, there are considerable variations in the growth patterns of the archosauromorphs. This is exemplified by the bone microstructure of Hyperodapedon, which deviates from the generalized slow‐growth pattern proposed for all basal archosauromorphs, suggesting that rapid growth was already present in the archosauromorphs. The cortical thickness of various long bones of Hyperodapedon bears similarity with that of several extant terrestrial quadrupeds, suggesting that Hyperodapedon was essentially a terrestrial quadruped.     

Bone microstructure of Vegavis iaai (Aves,Anseriformes) from the Upper Cretaceous of Vega Island,Antarctic Peninsula

Vegavis iaai is a neornithine bird coming from the Late Cretaceous Sandwich Bluff Member of the López de Bertodano Formation (Maastrichtian), Antarctic Peninsula. Vegavis constitutes the only unquestionable Cretaceous neornithine bird, and is known by the holotype and specimen MACN-PV 19.748. The goal of this paper is to present a detailed osteohistological analysis of V. iaai. Vegavis shows a highly vascularized fibrolamellar matrix lacking lines of arrested growths, features widespread among modern birds. This is consistent with previous hypotheses indicating that modern birds were dominant at high latitudes. This is probably related to high-metabolic rates shared by modern birds, whereas archaic taxa as Enantiornithes are absent or form a minority part of High-Latitude bird assemblages. Vegavis was a diver, characterised by a certain degree of limb osteosclerosis, with an increase of bone inner compactness, and inhibition of secondary remodelling, with no effect on the external dimensions of the bone, a combination of characters related to diving lifestyle. Based on Relative Bone Thickness it is possible to infer that Vegavis was a foot-propelled diving bird, similar to some extant anseriforms. Occurrence of osteosclerotic limb bones in Vegavis and Polarornis may constitute a derived shared feature, sustaining the hypothesis that both taxa are phylogenetically related.     

Mandibular biomechanics of Crocuta crocuta,Canis lupus,and the late Miocene Dinocrocuta gigantea (Carnivora,Mammalia)

The relative simplicity of the mandible and its functional integration with the upper dentition in carnivorans makes it an ideal subject for functional morphological studies. To compare the mandibular biomechanics of two convergently evolved bone‐cracking ecomorphologies, we used finite element modelling to analyse mandibular corpus stress. The bone‐cracking spotted hyena Crocuta crocuta was used as a living analogue to the late Miocene percrocutid Dinocrocuta gigantea, using the grey wolf Canis lupus as a molar bone‐crushing outgroup. Mandibular stress values during p3, p4, and m1 tooth biting are found to be lowest in Cr. crocuta, and elevated in both Ca. lupus and D. gigantea. However, the stress‐dissipation patterns of the pre‐m1 corpus are similar between Cr. crocuta and D. gigantea. Lastly, D. gigantea has a relatively weaker corpus at the post‐m1 position than either Cr. crocuta or Ca. lupus. These findings suggest that even though stress patterns are similar amongst the bone‐cracking ecomorphs, the extinct D. gigantea had a weaker mandibular structure when performing a comparable bone‐cracking task as in Cr. crocuta because of its slender post‐m1 corpus. Ontogeny could potentially play an important role in strengthening the post‐m1 corpus by growth in the dorsoventral axis, and continuous increase in biting performance through adulthood in living Cr. crocuta suggests the possibility of a relatively more delayed development to full bone‐cracking capability in D. gigantea. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 683–696.     

The utility of osteon shape and circularity for differentiating human and non-human Haversian bone

Distinguishing human from non‐human bone fragments is usually accomplished by observation of gross morphology. When macroscopic analysis is insufficient, histological approaches can be applied. Microscopic features, like plexiform bone or osteon banding, are characteristic of non‐humans. In the absence of such features, distinguishing Haversian bone as either human or non‐human proves problematic. This study proposes a histomorphometric approach for classifying species from Haversian bone. Two variables, osteon area (On.Ar.) and circularity (On.Cr.), are examined. Measurements were collected from three species (deer, dog, human) represented by various skeletal elements; only ribs were available for humans (ribs: deer n = 6, dog n = 6, human n = 26; humeri: deer n = 6, dog n = 6; femora: deer n = 6, dog n = 6). Qualitative analysis comparing human to non‐human On.Ar. demonstrated that human ribs have larger mean On.Ar. (0.036 mm²) than non‐human ribs (deer = 0.017 mm², dog = 0.013 mm²). On.Cr. in the ribs showed minor differences between species (deer = 0.877; dog = 0.885; human = 0.898). Results demonstrated no significant difference across long bone quadrants in long bones. Discriminant analyses run on the means for each sample demonstrated overlap in deer and dog samples, clustering the non‐human and human groups apart from each other. Mean On.Cr. proved a poor criterion (ribs only: 76.3%, pooled elements: 66.1%), while mean On.Ar. proved useful in identifying human from non‐human samples (ribs only: 92.1%, pooled elements: 93.5%). When variables were combined, accuracy increased to 100% correct classification for rib data and 98.4% when considering data from all elements. These results indicate that On.Ar. and On.Cr. are valuable histomorphometric tools for distinguishing human from non‐human Haversian bone. Am J Phys Anthropol, 2012. © 2012 Wiley Periodicals, Inc.     

Physiological components of growth differences between individual oysters (Crassostrea gigas) and a comparison with Saccostrea commercialis

Pacific oysters (Crassostrea gigas) of identical age from two genetically distinct lines, one fast growing and the other slow growing, were held at three levels of ration and analysed for physiological traits to explain differences in their rates of growth. The data supported three hypotheses; faster growth was associated with faster rates of consumption of food, reduced metabolic rate at maintenance (i.e., at zero growth), and reduced metabolic costs of growth. A comparison with the Sydney rock oyster, Saccostrea commercialis, based on similar experiments on the two species, indicated that faster growth of Pacific oysters depended on similar physiological differences; the mean metabolic costs of growth, however, were similar in the two species. It is suggested that a general model for genetically linked differences in the growth rate of bivalve molluscs will need to include the processes of metabolic control rather than relying solely on an analysis of the individual components of the energetics of growth.     

Scaling of growth and life history traits relative to body size, brain size, and metabolic rate in lorises and galagos (Lorisidae, primates)

A broad range of variation in body size, brain size, and metabolic rate occurs within the primate family Lorisidae, thus providing an opportunity to examine the relationship of these three parameters to variation in growth and life history traits. Data on adult body weight, gestation length, lactation length, age at first estrus, litter size, and growth parameters were collected from a captive colony of four lorisid species, Loris tardigradus, Nycticebus coucang, Galago crassicaudatus, and G. senegalensis. The data presented here constitute the most complete life history information available for these poorly understood prosimian species. Correlation and allometric analyses were performed to determine the relationships between variables. Among the lorisids studied, adult body weight, adult cranial capacity, and relative cranial capacity did not predict variation in life history traits. Adult basal metabolic rate predicted most of the variability in gestation length, lactation length, and growth parameters. Lorisines differ from similarly sized galagines in having lower basal metabolic rates, slower growth rates, slower developmental rates, and smaller litter sizes, resulting in reduced reproductive potential. This may be a consequence of lorisine adaptation to a diet of toxic insects. Metabolic rate and diet may be among the most important parameters to examine in any study of life history evolution.     

Digestive Efficiency Mediated by Serum Calcium Predicts Bone Mineral Density in the Common Marmoset (Callithrix jacchus)

Two health problems have plagued captive common marmoset (Callithrix jacchus) colonies for nearly as long as those colonies have existed: marmoset wasting syndrome and metabolic bone disease. While marmoset wasting syndrome is explicitly linked to nutrient malabsorption, we propose metabolic bone disease is also linked to nutrient malabsorption, although indirectly. If animals experience negative nutrient balance chronically, critical nutrients may be taken from mineral stores such as the skeleton, thus leaving those stores depleted. We indirectly tested this prediction through an initial investigation of digestive efficiency, as measured by apparent energy digestibility, and serum parameters known to play a part in metabolic bone mineral density of captive common marmoset monkeys. In our initial study on 12 clinically healthy animals, we found a wide range of digestive efficiencies, and subjects with lower digestive efficiency had lower serum vitamin D despite having higher food intakes. A second experiment on 23 subjects including several with suspected bone disease was undertaken to measure digestive and serum parameters, with the addition of a measure of bone mineral density by dual‐energy X‐ray absorptiometry (DEXA). Bone mineral density was positively associated with apparent digestibility of energy, vitamin D, and serum calcium. Further, digestive efficiency was found to predict bone mineral density when mediated by serum calcium. These data indicate that a poor ability to digest and absorb nutrients leads to calcium and vitamin D insufficiency. Vitamin D absorption may be particularly critical for indoor‐housed animals, as opposed to animals in a more natural setting, because vitamin D that would otherwise be synthesized via exposure to sunlight must be absorbed from their diet. If malabsorption persists, metabolic bone disease is a possible consequence in common marmosets. These findings support our hypothesis that both wasting syndrome and metabolic bone disease in captive common marmosets are consequences of inefficient nutrient absorption. Am. J. Primatol. 75:153‐160, 2013. © 2012 Wiley Periodicals, Inc.     

Chondrogenesis of the limbs and mesopodial ossification of Podocnemis expansa Schweigger, 1812 (Testudines: Podocnemidae)

We address the chondrogenic formation of the limbs and the mesopodial ossification pattern of the Pleurodira Podocnemis expansa, to resolve the homology of these elements as well as the pattern of connection of the autopodial elements and the origin of the digital arch. Embryos and juveniles of P. expansa were cleared and stained for cartilage and bone. The fore‐ and hind‐limbs were also studied histologically. We describe the development of the stylopodium and zeugopodium originating from a Y‐shaped cartilaginous condensation, and the differentiation of the primary axis and the digital arch in the initial stages of limb development. The most pronounced changes were observed in the chondrogenic pattern and ossification of the mesopodium, although development of the digits is similar and we found no ontogenetic reduction such as that described for other Testudines. In this study, as in previous research involving several groups of reptilian sauropsids, we found an inconsistent pattern between the chondrogenic formation and mesopodial ossification of the limbs, indicating that these developmental events are dissociated. In summary, the chondrogenic and ossification sequences of these elements do not follow the same pattern. In addition, the differences found between P. expansa and other species to which it was compared clearly indicate that these events follow more than one pattern in Testudines. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

Studies on skeleton formation in reptiles. I. The postembryonic development of the skeleton in Cyrtodactylus pubisulcus (Reptilia: Gekkonidae)

The study of ossification during postembryonic development of the lizard Cyrtodactylus pubisulcus reveals consistent patterns in the skeleton of the body axis and of the limbs. The vertebral column shows a distinct antero-posterior gradient in ossification; the serial homology of sacral ribs and caudal transverse processes with dorsal ribs requires further scrutiny. The sequence of ossification of carpal and tarsal elements is constant, yet different from the pattern of chondrification as described in the literature. The homology of a separate 'intermedium' in the ossified lizard carpus requires further discussion. The development of the lizard astragalus is discussed in detail, as is the ossification of epiphyses in the limbs.