A nearly complete skeleton of an early juvenile diplodocid (Dinosauria: Sauropoda) from the Lower Morrison Formation (Late Jurassic) of north central Wyoming and its implications for early ontogeny and pneumaticity in sauropods

A nearly complete skeleton of a juvenile sauropod from the Lower Morrison Formation (Late Jurassic, Kimmeridgian) of the Howe Ranch in Bighorn County, Wyoming is described. The specimen consists of articulated mid-cervical to mid-caudal vertebrae and most appendicular bones, but cranial and mandibular elements are missing. The shoulder height is approximately 67 cm, and the total body length is estimated to be less than 200 cm. Besides the body size, the following morphological features indicate that this specimen is an early juvenile; (1) unfused centra and neural arches in presacral, sacral and first to ninth caudal vertebrae, (2) unfused coracoid and scapula, (3) open coracoid foramen, and (4) relatively smooth articular surfaces on the limb, wrist, and ankle bones. A large scapula, short neck and tail and elongate forelimb bones relative to overall body size demonstrate relative growth. A thin-section of the mid-shaft of a femur shows a lack of annual growth lines, indicating an early juvenile individual possibly younger than a few years old. Pneumatic structures in the vertebral column of the specimen SMA 0009 show that pneumatisation of the postcranial skeleton had already started in this individual, giving new insights in the early ontogenetic development of vertebral pneumaticity in sauropods.

The specimen exhibits a number of diplodocid features (e.g., very elongate slender scapular blade with a gradually dorsoventrally expanded distal end, a total of nine dorsal vertebrae, presence of the posterior centroparapophyseal lamina in the posterior dorsal vertebrae). Although a few diplodocid taxa, Diplodocus, cf. Apatosaurus, and cf. Barosaurus, are known from several fossil sites near the Howe Ranch, identification of this specimen, even at a generic level, is difficult due to a large degree of ontogenetic variation.     

ESTIMATING BODY LENGTH OF PYGMY RIGHT WHALES (CAPEREA MARGINATA) FROM MEASUREMENTS OF THE SKELETON AND BALEEN

Various parts of the skeleton and/or the longest baleen plate of 46 specimens of Caperea marginata from Australia and New Zealand were measured and related to body length. Of the 32 skull, postcranial and baleen-plate measurements available, eight were analysed and seven found to be good predictors of body length, by using a curvilinear model describing their relationship with body length. Greatest skull width, supraoccipital length and mandible length had the smallest prediction limits (± 0.28-0.33 m in small animals, ±0.44-0.58 m in large animals) when compared with postcranial measurements (scapula length, vertebra 7 centrum width). Baleen-plate length was also a useful predictor of body length (±0.32-0.77 m). There was a substantial increase in the arch of the skull as body length increased. Bulla length was not a good predictor of body length, because measurements were highly variable and because the bulla grew little during postnatal life. Physical maturity occurred at body lengths of at least 5.9 m, also the shortest length at which both epiphyses of the humerus and proximal epiphyses of the radius and ulna were fused. Weaning appears to occur at about 3-3.5 m. The following approximate relative age/length classes were erected: dependent calves, <3.6 m; subadults, 3.6-5.5 m; adults, >5.5 m. Females were significantly longer than males in the sample of 22 animals greater than 5.9 m, length of the smallest recorded physically mature animal.     

THE POSTCRANIUM OF ARCHEGOSAURUS DECHENI, AND A PHYLOGENETIC ANALYSIS OF TEMNOSPONDYL POSTCRANIA

Abstract:  The gharial-like Archegosaurus decheni from the Permian/Carboniferous boundary of south-west Germany is one of the best known temnospondyls. Based largely on new material, we restudied the postcranial anatomy of this species, including ontogenetic aspects. A. decheni has 24 presacral vertebrae, and the length of the deep tail exceeds the length of the rest of the body. Neural spines are moderately high and slowly become differentiated during ontogeny. The intercentra start to ossify very late. Distal uncinate processes are developed on the anterior ribs in adult specimens. Only the ventral portion of the scapula is ossified. The slender ilium and the ischium are not co-ossified, and the pubis remained cartilaginous. Among stereospondylomorph temnospondyls, the very short and rudimentarily developed humerus exhibits a unique morphology. Carpals and tarsals start to ossify only in the largest specimens. The poorly ossified postcranium indicates that A. decheni was primarily an aquatic temnospondyl. We undertook a phylogenetic study of A. decheni and 16 other temnospondyls, based exclusively on postcranial characters. We analysed 52 characters, obtaining a single most parsimonious tree that agrees in many aspects with cranium-based analyses: Trimerorhachis and Dvinosaurus form a basal clade (Dvinosauria), followed by monophyletic euskelians (dissorophoids plus eryopids) and finally the stereospondylomorphs, within which A. decheni is nested above Sclerocephalus . Among stereospondyls, trematosaurids and metoposaurids form a clade, whereas the chigutisaurid Siderops is nested with capitosauroids. The primitive condition of Temnospondyli is still not adequately understood, especially the degree of terrestriality of the earliest temnospondyls.     

Axial elongation in fishes: using morphological approaches to elucidate developmental mechanisms in studying body shape

One of the most notable features in looking across fishes is their diversity of body shape and size. Extant actinopterygian fishes range in shape from nearly spheroidal in pufferfishes to extremely elongate in snipe eels with nearly every shape in-between. One extreme along the body-shape continuum is a highly elongate form, which has evolved multiple times independently in Actinopterygii. Thus, comparison of these separate (independent) radiations provides a unique opportunity for examining the anatomical traits underlying elongation as well as the similarities and differences in the evolutionary pathways followed. Body elongation generally evolves via an increase in region-specific vertebral number, although certain lineages elongate via an increase in vertebral length. In this study, we describe how anatomical characters related to feeding and locomotion are correlated with elongation of the body across Actinopterygii. In addition to modifications of the postcranial axial skeleton, elongation in fishes is often accompanied by an increase in head length, loss of the pelvic fins, reduction of the pectoral fins, and expansion of the median fins. Based on anatomical studies and on recent studies of developmental control of the body axis in different species, we hypothesize how an axial trait might change at the genetic level. Overall, we discuss the evolution of body elongation in fishes in light of an understanding of the underlying anatomical modifications, developmental control, ecology, and locomotion.     

Morphology and Function of the Vertebral Column in Remingtonocetus domandaensis (Mammalia, Cetacea) from the Middle Eocene Domanda Formation of Pakistan

The archaeocete family Remingtonocetidae is a group of early cetaceans known from the Eocene of India and Pakistan. Previous studies of remingtonocetids focused primarily on cranial anatomy due to a paucity of well-preserved postcranial material. Here we describe the morphology of the known vertebral column in Remingtonocetus domandaensis based largely on a single well-preserved partial skeleton recovered from the upper Domanda Formation of Pakistan. The specimen preserves most of the precaudal vertebral column in articulation and includes seven complete cervical vertebrae, ten partial to complete thoracic vertebrae, six complete lumbar vertebrae, and the first three sacral vertebrae. Cervical centra are long and possess robust, imbricating transverse processes that stabilized the head and neck. Lumbar vertebrae allowed for limited flexibility and probably served primarily to stabilize the lumbar column during forceful retraction of the hind limbs. Vertebral evidence, taken together with pelvic and femoral morphology, is most consistent with interpretation of Remingtonocetus domandaensis as an animal that swam primarily by powerful movement of its hind limbs rather than dorsoventral undulation of its body axis.     

A small ichthyosaur from the Clearwater Formation (Alberta,Canada) and a discussion of the taxonomic utility of the pectoral girdle

Albian sedimentary successions of northwestern Canada have yielded a diverse assemblage of Mesozoic marine vertebrates, and ichthyosaurs form an important component of these faunas. Here, we describe a partial postcranial skeleton of a small (estimated at less than 3 m total body length) ichthyosaur from the Wabiskaw Member of the Clearwater Formation (lowermost Albian). The semi-articulated specimen includes much of the presacral vertebral column, dorsal ribs and gastralia. Most significantly, it possesses an articulated pectoral girdle and humerus, and also preserves the pelvic girdle, allowing new insights into girdle evolution in ichthyosaurs. Whereas both sets of girdles are thought to exhibit large amounts of intraspecific variation, the pectoral girdle of ophthalmosaurids appears to evolve very slowly, remaining essentially unchanged from the Middle Jurassic onwards. In contrast, the pelvic girdle shows taxonomically informative changes within Ophthalmosauridae. The variable and poorly known nature of girdle morphology in Cretaceous ichthyosaurs precludes generic referral of the specimen.     

AN ALMOST COMPLETE JUVENILE SPECIMEN OF THE CHELONIID TURTLE CTENOCHELYS STENOPORUS (HAY, 1905) FROM THE UPPER CRETACEOUS NIOBRARA FORMATION OF KANSAS,USA

Abstract: A new, unusually well‐preserved juvenile specimen of Ctenochelys stenoporus from the Niobrara Formation is described. The skull has come apart at its sutures and all bones of the braincase and ear region are preserved three‐dimensionally. This allows a detailed reconstruction of the important brain structures of a basal juvenile cheloniid turtle. It is compared with adult Ctenochelys specimens, and the major ontogenetic changes in the skull and postcranial skeleton are described. Furthermore, the specimen is compared with other fossil and extant cheloniids with well‐known braincases and the differences between basal and advanced cheloniids turtles are specified.     

Osteological development of the vertebral column and of the caudal complex in Dentex dentex

Osteological development of the vertebral column and caudal complex in common dentex was described under extensive larval rearing conditions. Generally, the cartilaginous bones developed prior to the membranous bones. The development of the axial skeleton began with the formation of the hypural 1, the neural arches 2 and 3, as well as the haemal arches 1–8 at 4·8, 4·9 and 5·0 mm total length ( L _T, measured in vivo ), respectively. By 7·5 mm L _T, all the cartilaginous elements were formed, except for the ventral ribs, which formed between the range of 8·4–18·0 mm L _T. The caudal lepidotrichia were the first membranous bones to appear (5·3 mm L _T) and attain their full meristic count (7·4 mm L _T), followed by the vertebral centra, which formed between 6·6 and 9·7 mm L _T. By 25·0 mm L _T, all the elements were fully ossified except for the ventral ribs. The developmental direction and order of all the elements were studied with respect to their formation and ossification. The results were discussed in the contexts of ichthyoplankton, ecology and aquaculture. Compared with other Sparidae species, common dentex followed a pattern of relatively rapid rate of osteological development.     

Differential occupation of axial morphospace

The postcranial system is composed of the axial and appendicular skeletons. The axial skeleton, which consists of serially repeating segments commonly known as vertebrae, protects and provides leverage for movement of the body. Across the vertebral column, much numerical and morphological diversity can be observed, which is associated with axial regionalization. The present article discusses this basic diversity and the early developmental mechanisms that guide vertebral formation and regionalization. An examination of vertebral numbers across the major vertebrate clades finds that actinopterygian and chondrichthyan fishes tend to increase vertebral number in the caudal region whereas Sarcopterygii increase the number of vertebrae in the precaudal region, although exceptions to each trend exist. Given the different regions of axial morphospace that are occupied by these groups, differential developmental processes control the axial patterning of actinopterygian and sarcopterygian species. It is possible that, among a variety of factors, the differential selective regimes for aquatic versus terrestrial locomotion have led to the differential use of axial morphospace in vertebrates.     

FUNCTIONAL AND EVOLUTIONARY ASPECTS OF THE POSTCRANIAL ANATOMY OF DICYNODONTS (SYNAPSIDA, THERAPSIDA)

Abstract:  Restoration of the major skeletal muscles and functional morphological analysis of the postcranium were carried out on two Triassic dicynodont genera, Wadiasaurus and Lystrosaurus . A phylogenetic analysis of 12 selected Permian and Triassic dicynodont taxa was conducted and the postcranial character states were then mapped onto the most parsimonious tree. The analysis revealed changes in pectoral girdle and forelimb morphology, which included reduction of the coracoid plate, increasing robustness of the deltopectoral crest, change in humeral orientation from lateral to caudolateral, increasing prominence of the humeral head, and increasing robustness of the radius. Such changes can be associated with a functional tendency to reduce the lateral component of the propulsive force while still in an abducted mode. On the other hand, changes associated with the pelvic girdle included expansion of the preacetabular iliac process, reduction of the postacetabular iliac process, craniocaudal expansion of the iliac blade, change in the shape of the pubis from flat and plate-like to small and rod-like with a cranial process, and change in acetabular orientation from lateral to caudolateral. The femoral head, starting from a cranioproximal position, progressively became dorsally pronounced and offset from the body. Other features/changes associated with the femur included increasing robustness of the trochanter major, and increasing flattening of the femoral midshaft. Changes in the axial skeleton included increasing stiffening of the trunk to reduce lateral undulations, increasing dorsoventral flexion, and increasing sacral vertebral count, which can be correlated with the preacetabular iliac expansion. These findings suggest that the dicynodont postcranial skeleton evolved towards more upright hindlimb morphology with the body held well off the ground.     

Dual origin and segmental organisation of the avian scapula

Bones of the postcranial skeleton of higher vertebrates originate from either somitic mesoderm or somatopleural layer of the lateral plate mesoderm. Controversy surrounds the origin of the scapula, a major component of the shoulder girdle, with both somitic and lateral plate origins being proposed. Abnormal scapular development has been described in the naturally occurring undulated series of mouse mutants, which has implicated Pax1 in the formation of this bone. Here we addressed the development of the scapula, firstly, by analysing the relationship between Pax1 expression and chondrogenesis and, secondly, by determining the developmental origin of the scapula using chick quail chimeric analysis. We show the following. (1) The scapula develops in a rostral-to-caudal direction and overt chondrification is preceded by an accumulation of Pax1-expressing cells. (2) The scapular head and neck are of lateral plate mesodermal origin. (3) In contrast, the scapular blade is composed of somitic cells. (4) Unlike the Pax1-positive cells of the vertebral column, which are of sclerotomal origin, the Pax1-positive cells of the scapular blade originate from the dermomyotome. (5) Finally, we show that cells of the scapular blade are organised into spatially restricted domains along its rostrocaudal axis in the same order as the somites from which they originated. Our results imply that the scapular blade is an ossifying muscular insertion rather than an original skeletal element, and that the scapular head and neck are homologous to the 'true coracoid' of higher vertebrates.     

Evolution of the hominoid vertebral column: The long and the short of it

The postcranial axial skeleton exhibits considerable morphological and functional diversity among living primates. Particularly striking are the derived features in hominoids that distinguish them from most other primates and mammals. In contrast to the primitive catarrhine morphotype, which presumably possessed an external (protruding) tail and emphasized more pronograde trunk posture, all living hominoids are characterized by the absence of an external tail and adaptations to orthograde trunk posture. Moreover, modern humans evolved unique vertebral features that satisfy the demands of balancing an upright torso over the hind limbs during habitual terrestrial bipedalism. Our ability to identify the evolutionary timing and understand the functional and phylogenetic significance of these fundamental changes in postcranial axial skeletal anatomy in the hominoid fossil record is key to reconstructing ancestral hominoid patterns and retracing the evolutionary pathways that led to living apes and modern humans. Here, we provide an overview of what is known about evolution of the hominoid vertebral column, focusing on the currently available anatomical evidence of three major transitions: tail loss and adaptations to orthograde posture and bipedal locomotion.     

Cranial variables as predictors of hominine body mass

Body mass is a key variable in investigating the evolutionary biology of the hominines (Australopithecus, Paranthropus, and Homo). It is not only closely related to life-history parameters but also provides a necessary baseline for studies of encephalization or megadonty. Body mass estimates are normally based on the postcranial skeleton. However, the majority of hominid fossils are cranio-dental remains that are unassociated with postcranial material. Only rarely can postcranial material be linked with craniodentally defined hominid taxa. This study responds to this problem by evaluating body mass estimates based on 15 cranial variables to determine whether they compare in reliability with estimates determined from postcranial variables. Results establish that some cranial variables, and particularly orbital area, orbital height, and biporionic breadth, are nearly as good mass predictors for hominoids as are some of the best postcranial predictors. For the hominines in particular, orbital height is the cranial variable which produces body mass estimates that are most in line with postcranially generated estimates. Both orbital area and biporionic breadth scale differently in the hominines than they do in the other hominoids. This difference in scaling results in unusually large estimates of body mass based on these variables for the larger-sized hominines, although the three cranial variables produce equivalent predicted masses for the smaller-bodied hominines. © 1994 Wiley-Liss, Inc.     

Scapular position in primates

Scapular position affects shoulder mobility, which plays an important role in the upper limb adaptations in primates. However, currently available data on scapular position are unsatisfactory because of the failure to simultaneously consider the relative dimensions of all the three skeletal elements of the shoulder girdle, i.e. the clavicle, the scapula and the thorax. In the present study, the clavicular length and the scapular spine length were measured on preserved cadavers, and the dorsoventral thoracic diameter was measured on scaled radiographs of a wide range of primates, permitting a quantitative comparison of scapular position among primates. It was found that arboreal monkeys have a more dorsally situated scapula than terrestrial ones, but the same difference was not found between terrestrial and arboreal prosimians. Hominoids were found to have the most dorsally situated scapula. Contrary to the slow climbing theory of hominoid evolution, which tries to explain most postcranial specializations of hominoids as adaptations for slow climbing, the scapulae of slow-climbing lorines and Alouatta are much less dorsal than those of the hominoids.     

The axial skeleton of the labyrinthodont Eogyrinus attheyi

An articulated length of vertebral column is used as a basis for the reconstruction of the salient features of the axial skeleton of the embolomerous anthracosaur Eogyrinus attheyi Watson, together with other material, including the holotype, in the Hancock Museum, Newcastle upon Tyne.
The trunk vertebrae are typically emboloinerous, with disoshaped notochordal pleuro-centra, firmly attached by broad facets to this neural arches, and much thinner intercentra. Regional variation is chiefly concerned with the span of the transverse processes, which diminishes posteriorly, and the associated separation of the two heads of each rib. A longitudinal series of trunk ribs, of diminishing length from the mid-trunk backwards, is reconstructed.
Eogyrinus has a normal tetrapod sacrum with one characteristic sacral rib. The first few caudal vertebrae bear ribs of unusual form, (of which four are preserved in sequence in the articulated specimen. The fifth caudal intercentrum bears the first and largest haemal arch and the pleurocentrum of the seventh caudal is distinguished by marked muscle origins presumably for the caudifemoral muscles.
The probability that Eogyrinus, like the few other embolomeres known, had an unusually long vertebral column for a labyrinthodont, is supported by an orthometric comparison using Romer's data on the American form Archeria.     

A NEW RECONSTRUCTION OF ONYCHOSELACHE TRAQUAIRI, COMMENTS ON EARLY CHONDRICHTHYAN PECTORAL GIRDLES AND HYBODONTIFORM PHYLOGENY

Abstract:  A new, third, specimen of Onychoselache traquairi from the Viséan (Holkerian) of Scotland allows a significant revision of the anatomy of this stem-group elasmobranch. This first report of material from the Mumbie Quarry exposure of the Glencartholm fish beds presents a new reconstruction of Onychoselache showing broad-based cephalic and nuchal spines, and exceptionally large pectoral fins. Details of the jaws, braincase and postcranial skeleton demonstrate that Onychoselache is a well-characterized member of the Hybodontiformes. Comparisons of the pectoral skeleton with other early chondrichthyan examples, including new material of Tristychius arcuatus and Plesioselachus macracanthus , highlight a range of early chondrichthyan conditions that are incorporated into a revised hybodontiform phylogeny. Close resemblance between Onychoselache and Mesozoic and late Palaeozoic hybodonts implies that these clades diverged within the Carboniferous and Permian. Major differences between Onychoselache and the coeval Tristychius (a modified reconstruction of which is included) indicate that the Neoselachii-Hybodontiformes split is probably Late Devonian, consistent with records of isolated teeth. The pectoral fins of Onychoselache , while unique among Palaeozoic forms, resemble those of Recent bamboo and epaulette sharks (Orectolobiformes). The functional corollary of this convergence is that Onychoselache represents an instance of a non-tetrapod early vertebrate with a near-walking gait.     

Functional aspects of strepsirrhine lumbar vertebral bodies and spinous processes

The relationship between form and function in the lumbar vertebral column has been well documented among platyrrhines and especially catarrhines, while functional studies of postcranial morphology among strepsirrhines have concentrated predominantly on the limbs. This morphometric study investigates biomechanically relevant attributes of the lumbar vertebral morphology of 20 species of extant strepsirrhines. With this extensive sample, our goal is to address the influence of positional behavior on lumbar vertebral form while also assessing the effects of body size and phylogenetic history. The results reveal distinctions in lumbar vertebral morphology among strepsirrhines in functional association with their habitual postures and primary locomotor behaviors. In general, strepsirrhines that emphasize pronograde posture and quadrupedal locomotion combined with leaping (from a pronograde position) have the relatively longest lumbar regions and lumbar vertebral bodies, features promoting sagittal spinal flexibility. Indrids and galagonids that rely primarily on vertical clinging and leaping with orthograde posture share a relatively short (i.e., stable and resistant to bending) lumbar region, although the length of individual lumbar vertebral bodies varies phylogenetically and possibly allometrically. The other two vertical clingers and leapers, Hapalemur and Lepilemur, more closely resemble the pronograde, quadrupedal taxa. The specialized, suspensory lorids have relatively short lumbar regions as well, but the lengths of their lumbar regions are influenced by body size, and Arctocebus has dramatically longer vertebral bodies than do the other lorids. Lumbar morphology among galagonids appears to reflect a strong phylogenetic signal superimposed on a functional one. In general, relative length of the spinous processes follows a positively allometric trend, although lorids (especially the larger-bodied forms) have relatively short spinous processes for their body size, in accordance with their positional repertoire. The results of the study broaden our understanding of postcranial adaptation in primates, while providing an extensive comparative database for interpreting vertebral morphology in fossil primates.