“Lucy” (A.L. 288‐1) had five sacral vertebrae

A “long‐backed” scenario of hominin vertebral evolution posits that early hominins possessed six lumbar vertebrae coupled with a high frequency of four sacral vertebrae (7:12‐13:6:4), a configuration acquired from a hominin‐panin last common ancestor (PLCA) having a vertebral formula of 7:13:6‐7:4. One founding line of evidence for this hypothesis is the recent assertion that the “Lucy” sacrum (A.L. 288‐1an, Australopithecus afarensis) consists of four sacral vertebrae and a partially‐fused first coccygeal vertebra (Co1), rather than five sacral vertebrae as in modern humans. This study reassesses the number of sacral vertebrae in Lucy by reexamining the distal end of A.L.288‐1an in the context of a comparative sample of modern human sacra and Co1 vertebrae, and the sacrum of A. sediba (MH2). Results demonstrate that, similar to S5 in modern humans and A. sediba, the last vertebra in A.L. 288‐1an exhibits inferiorly‐projecting (right side) cornua and a kidney‐shaped inferior body articular surface. This morphology is inconsistent with that of fused or isolated Co1 vertebrae in humans, which either lack cornua or possess only superiorly‐projecting cornua, and have more circularly‐shaped inferior body articular surfaces. The level at which the hiatus' apex is located is also more compatible with typical five‐element modern human sacra and A. sediba than if only four sacral vertebrae are present. Our observations suggest that A.L. 288‐1 possessed five sacral vertebrae as in modern humans; thus, sacral number in “Lucy” does not indicate a directional change in vertebral count that can provide information on the PLCA ancestral condition. Am J Phys Anthropol 156:295–303, 2015. © 2014 Wiley Periodicals, Inc.     

A method for estimation of lateral and vertical mobility of platycoelous vertebrae of tetrapods

A new method for the estimation of dorsoventral and lateral mobility of platycoelous vertebrae with V-shaped (radial) articular facets on the zygapophyses is developed. This vertebral pattern is observed in dinosaurs, some other fossil reptiles, and in the cervical and lumbar regions of mammals. Based on theoretical biomechanical analysis of the intervertebral discs and articulations between zygapophyses, the estimation formulas are developed and calibrated, using precise measurements of mobility between cervical vertebrae of domestic sheep. The method is applied to the presacral vertebrae of the horned dinosaur Protoceratops andrewsi. In its cervical, lumbar, and anterior thoracic regions, the differences between the calculated amplitudes of movements and the sought true values are expected to range within ±5°. As compared to the sheep, Protoceratops shows a greater lateral mobility in the presacral region and reduced vertical mobility in the cervical region.     

Vertebrae numbers of the early hominid lumbar spine

General doctrine holds that early hominids possessed a long lumbar spine with six segments. This is mainly based on Robinson's (1972) interpretation of a single partial Australopithecus africanus skeleton, Sts 14, from Sterkfontein, South Africa. As its sixth last presacral vertebra exhibits both thoracic and lumbar characteristics, current definitions of lumbar vertebrae and lumbar ribs are discussed in the present study. A re-analysis of its entire preserved vertebral column and comparison with Stw 431, another partial A. africanus skeleton from Sterkfontein, and the Homo erectus skeleton KNM-WT 15000 from Nariokotome, Kenya, did not provide strong evidence for the presence of six lumbar vertebrae in either of these early hominids. Thus, in Sts 14 the sixth last presacral vertebra has on one side a movable rib. In Stw 431, the corresponding vertebra shows indications for a rib facet. In KNM-WT, 15000 the same element is very fragmentary, but the neighbouring vertebrae do not support the view that it is L1. Although in all three fossils the transitional vertebra at which the articular facets change orientation seems to be at Th11, this is equal to a large percentage of modern humans. Indeed, a modal number of five lumbar vertebrae, as in modern humans, is more compatible with evolutionary principles. For example, six lumbar vertebrae would require repetitive shortening and lengthening not only of the lumbar, but also of the entire precaudal spine. Furthermore, six lumbar vertebrae are claimed to be biomechanically advantageous for early hominid bipedalism, yet an explanation is lacking as to why the lumbar region should have shortened in later humans. All this raises doubts about previous conclusions for the presence of six lumbar vertebrae in early hominids. The most parsimonious explanation is that they did not differ from modern humans in the segmentation of the vertebral column.     

Reevaluation of the lumbosacral region of Oreopithecus bambolii

Functional interpretations of the postcranium of the late Miocene ape Oreopithecus bambolii are controversial. The claim that Oreopithecus practiced habitual terrestrial bipedalism is partly based on restored postcranial remains originally recovered from Baccinello, Tuscany ( Köhler and Moyà-Solà, 1997). The lower lumbar vertebrae of BA#72 were cited as evidence that Oreopithecus exhibits features indicative of a lordotic lumbar spine, including dorsal wedging of the vertebral bodies and a caudally progressive increase in postzygapophyseal interfacet distance. Here, we demonstrate why the dorsal wedging index value obtained by Köhler and Moyà-Solà (1997) for the BA#72 last lumbar vertebra is questionable due to distortion in that region, present a more reliable way to measure postzygapophyseal interfacet distance, and include an additional metric (laminar width) with which to examine changes in the transverse dimensions of the neural arches. We also quantify the external morphology of the BA#72 proximal sacrum, which, despite well-documented links between sacral morphology and bipedal locomotion, and excellent preservation of the sacral prezygapophyses, first sacral vertebral body, and right ala, was not evaluated by Köhler and Moyà-Solà (1997). Measures of postzygapophyseal interfacet distance and laminar width on the penultimate and last lumbar vertebrae of BA#72 reveal a pattern encompassed within the range of living nonhuman hominoids and unlike that of modern humans, suggesting that Oreopithecus did not possess a lordotic lumbar spine. Results further show that the BA#72 sacrum exhibits relatively small prezygapophyseal articular facet surface areas and mediolaterally narrow alae compared with modern humans, indicating that the morphology of the Oreopithecus sacrum is incompatible with the functional demands of habitual bipedal stance and locomotion. The Oreopithecus lumbosacral region does not exhibit adaptations for habitual bipedal locomotion.     

Fused mid‐caudal vertebrae in the titanosaur Uberabatitan ribeiroi from the Late Cretaceous of Brazil and other bone lesions

Bone pathology in the titanosaur dinosaur Uberabatitan ribeiroi from the Marília Formation (Bauru Group, Late Cretaceous) of Uberaba city (Minas Gerais State, Brazil) is analysed here. They include two fused procoelous mid‐caudal vertebrae (CPPLIP‐1020) and a haemal arch (CPPLIP‐1006) of the middle section of the tail with a healing fracture callus. The analyses of the caudal vertebrae CPPLIP‐1020 of Uberabatitan permit us to recognize the following signs, based on CT scan and external macroscopic observations: (1) ossified longitudinal tendons; (2) likely ossified intervertebral disc, producing fused vertebral bodies; (3) fused right zygapophyseal process with a laterally developed osteophyte affecting this joint; (4) osteophytes and exostoses at different portions of the vertebrae; (5) cloacae, bone erosion and likely internal infection. According to all the processes observed in these caudals, we could not discard at least two possibilities for the diagnosis of the fused vertebrae. It could be the result of a spondyloarthropathy process (considering most of the observed signs) or possibly have been associated with an infection (e.g. discospondylitis/infections spondylitis or septic arthritis). The bone lesion record in Uberabatitan ribeiroi from the Late Cretaceous of Brazil increases the range of study of titanosaur dinosaurs, which although have a large fossil record, have few pathological studies.     

Sphingosine 1-phosphate differentially regulates proliferation of C2C12 reserve cells and myoblasts

Scoliosis is a condition that involves an abnormal curvature and deformity of the spinal vertebrae. The genetic background and key gene for congenital scoliosis in humans are still poorly understood. Ishibashi rats (ISR) have congenital malformation of the lumbar vertebrae leading to kyphoscoliosis similar to that seen in humans. To understand the pathogenesis of congenital scoliosis, we have studied the abnormality of vertebral formation and the associated gene expression in ISR. Almost all ISR showed kyphosis or scoliosis of the lumbar vertebrae. In ISR with severe kyphosis, some vertebral disks were missing and some vertebral bodies were fused. Of the ISR, 27% showed hemi-lumbarization of lumbar and sacral vertebrae. Homeotic transformation of the first sacral vertebra into the seventh lumbar vertebra and the resultant loss of the fourth sacral vertebra were seen in half of the ISR. We also found unilateral fusions and deformities of primary ossification centers of the lumbar vertebral column in fetal ISR. Moreover, we observed that the expression levels of Hox10 and Hox11 paralogs in lumbo-sacral transitional areas of ISR were extremely low compared with those of normal rats. These results suggest that fusion of primary ossification centers in lumbar vertebrae in the embryonic period causes scoliosis and kyphosis and that Hox genes are involved in the occurrence of homeotic transformation in lumbo-sacral vertebrae of congenital kyphoscoliotic ISR.     

The Developmental and Morphological Studies on the Neural and Skeletal Abnormalities in the T/btm Tailless Mice

The crosses, T /+ or T/t ^w2× btm/btm , give rise to 50% incidence of the tailless mice, development of which was investigated. No difference was seen in external appearance of the embryos at 9 days of gestation. However, some embryos showed fusion of the notochord and the neural tube at the posterior part of the body on the histological examination. The prospective tailless individuals were distinguishable from the normal littermates by the constriction of the root of the tail at 10 days of gestation. Thereafter, they showed several abnormalities such as the poor growth of the posterior part of the body, thinning of the tail and a blood blister at the tail tip or in the lumbosacral region. The abnormal embryos of 11–12 days showed severer abnormalities in the medio-dorsal area, i.e., the notochord was branched or degenerated at several places and the neural tube was distorted, duplicated or fused with the mesenchyme. All the tailless newborn young had blood blisters or red scars on the dorsal skin at the middle of the lumbosacral region.
Histologically, the spinal cord posterior to the lumbosacral level was revealed to be severely distorted or duplicated and completely devoid of the bony vertebrae, and the dorsal blood blister was found to be the meningomyelocele derived from the abnormal development of the spinal cord. Skeletal abnormalities of the tailless young were as follows. The sacral and caudal vertebrae were absent. The cervical vertebrae were mostly normal, but the thoracic and lumbar vertebrae showed several abnormalities such as fusion of the ribs, lack of the vertebral body and vertebral arch.     

Does postcranial palaeoneurology provide insight into pterosaur behaviour and lifestyle? New data from the azhdarchoid Vectidraco and the ornithocheirids Coloborhynchus and Anhanguera

The postcranial palaeoneurology of fossil reptiles is understudied, and those studies that exist focus predominantly on crocodyliforms and dinosaurs. The intervertebral foramina of the spine house nerves that exit to innervate surrounding tissues and the extremities. In the heavily fused (and typically distorted or poorly preserved) pterosaurian sacrum, intervertebral foramina can be difficult to observe and are rarely identified. The Early Cretaceous azhdarchoid Vectidraco from the Isle of Wight, UK, exhibits large, paired foramina on each sacral vertebra, originally identified as pneumatic foramina. Micro‐computed tomography imaging reveals these communicate with the neural canal and are intervertebral foramina for sacral nerves. The sacral vertebrae of Vectidraco are fused, and intervertebral foramina occur dorsolaterally on the centra. We identified these structures in other pterosaur sacra, including those of the ornithocheiroids Anhanguera and Coloborhynchus. The sizes of the sacral and notarial neural canals are compared and considered within interpretations of palaeoecology and locomotion, following previous studies. The relatively large sacral neural canal of Vectidraco implies a sacral enlargement for innervation of the legs and lumbosacral plexus. When compared with Anhanguera, this supports indications that azhdarchoids were more hindlimb‐proficient than ornithocheiroids. Neural canal size in the Coloborhynchus notarium suggests that ornithocheirids spent less time on the ground, their brachial enlargement and small sacral region indicating enhanced innervation of the wings and poor innervation of the sacrum and legs. This is the first study focusing on pterosaur postcranial palaeoneurology; more studies on other taxa are needed to reveal patterns across Pterosauria as a whole.     

Spinal arthritis and physical stress at Bronze Age Harappa

This study examines joint changes in the vertebral skeleton in human remains excavated in 1987 and 1988 at Bronze Age Harappa, an urban center of the Indus Valley civilization. The sample consists of 23 complete skeletons from primary burial context, the partial remains of more than 69 other individuals, and hundreds of skeletal elements from secondary context, totalling 3,084 vertebral joint margins and articular surfaces. Marginal bone proliferation, pitting of articular surfaces, eburnation, and ankylosis were scored macroscopically for vertebral body margins and surfaces and posterior apophyseal facet joints. Marginal lipping is far more prevalent on the vertebral bodies than on the apophyseal facets and surface pitting is also more frequent on vertebral bodies although its expression is relatively low overall. Cervical vertebrae in this sample exhibit the same amount of marginal new bone and much more surface pitting of the vertebral bodies than do either thoracic or lumbar vertebrae; the cervical segment also exhibits the most severe expressions of both types of lesions. In addition, although the frequencies of cervical and lumbar posterior facet involvement are similar, the cervical facets exhibit much more severe lipping as well as the only cases of eburnation and ankylosis. Pitting of the posterior facets is most common in the lumbar segment, but the cervical examples are the only severe cases. It is proposed that the severe joint changes in the cervical spine result from trauma, perhaps accumulated microtrauma from activity stresses. There are no age or sex associated patterns in the frequency of arthritis although this result may be influenced by the small proportions of the total sample for which age and sex could be determined. © 1994 Wiley-Liss, Inc.     

Remodelling of the vertebral axis during metamorphic shrinkage in the pearlfish

Body shortening was observed in the pearlfish Carapus homei during metamorphosis. The tenuis larva at first possessed a suite of osseous vertebral bodies of similar length. The reduction in both the number and size of vertebrae followed increasing decalcification, degeneration of organic tissue and shortening. This involved a complete degradation and disappearance of the caudal tip vertebrae, and there was a reduction in the size of most of the remaining vertebrae. The further development of the vertebrae began with ossification of the neural and haemal arches before that of the vertebral body. This second part of the development followed a gradient: a gradual decreases towards the caudal tip in the size of the vertebrae and their completeness.     

Numerical variation of the presacral vertebral column in three population groups in North America

A total of 1,239 skeletons from among Mongoloid, Caucasoid and Negroid population groups in North America was examined for variations in the number of presacral vertebrae. The overall incidence of variation was 11%; 6% with 23 and 5% with 25 presacral vertebrae. Differences in total variation among the three groups were not significant, but differences in the incidence of 23 and 25 presacral vertebrae among the groups were highly significant. Numerical vertebral variation occurred in 11% of both sexes, but with males having a higher frequency of 25 presacral vertebrae and females having a higher frequency of 23 presacral vertebrae. The incidence of the specific variation of 23 or 25 presacral vertebrae was not significantly different among the males of the three groups studied, whereas 23 presacral vertebrae were found significantly more often in the Negroid females. Numerical variation of vertebrae was not associated with age. The data strongly support the conclusion that the total frequency of variation in the number of presacral vertebrae is a specific characteristic of any particular population group and that there is a tendency in all population groups toward an increase in number in males and a decrease in number in females.     

Short-snouted toothless ichthyosaur from China suggests Late Triassic diversification of suction feeding ichthyosaurs

Background

Ichthyosaurs were an important group of Mesozoic marine reptiles and existed from the Early Triassic to the early Late Cretaceous. Despite a great diversity in body shapes and feeding adaptations, all share greatly enlarged eyes, an elongated rostrum with numerous conical teeth, and a streamlined body.

Methodology/Principal Findings

Based on new material from China and the restudy of Shastasaurus pacificus, we here reinterpret the classical large-bodied Late Triassic ichthyosaur genus Shastasaurus to differ greatly from the standard ichthyosaurian body plan, indicating much greater morphological diversity and range of feeding adaptations in ichthyosaurs than previously recognized. Phylogenetic analysis indicates a monophyletic clade consisting of the giant Shonisaurus sikanniensis, Guanlingsaurus liangae, and Shastasaurus pacificus to which the genus name Shastasaurus is applied. Shastasaurus liangae comb. nov. is from the Late Triassic (Carnian) Xiaowa Formation of Guizhou Province, southwestern China. The species combines a diminutive head with an entirely toothless and greatly reduced snout. The species also has by far the highest vertebral count among ichthyosaurs (86 presacral vertebrae and >110 caudal vertebrae), a count that is also very high for tetrapods in general. A reduced toothless snout and a diminutive head is also apparently present in the giant S. sikanniensis and presumably in S. pacificus.

Conclusions/Significance

In analogy to many modern odontocetes, Shastasaurus is interpreted as a specialized suction feeder on unshelled cephalopods and fish, suggesting a unique but widespread Late Triassic diversification of toothless, suction-feeding ichthyosaurs. Suction feeding has not been hypothesized for any of the other diverse marine reptiles of the Mesozoic before, but in Shastasaurus may be linked to the Late Triassic minimum in atmospheric oxygen.     

Dietary constraints of phytosaurian reptiles revealed by dental microwear textural analysis

Phytosaurs are a group of large, semi‐aquatic archosaurian reptiles from the Middle–Late Triassic. They have often been interpreted as carnivorous or piscivorous due to their large size, morphological similarity to extant crocodilians and preservation in fluvial, lacustrine and coastal deposits. However, these dietary hypotheses are difficult to test, meaning that phytosaur ecologies and their roles in Triassic food webs remain incompletely constrained. Here, we apply dental microwear textural analysis to the three‐dimensional sub‐micrometre scale tooth surface textures that form during food consumption to provide the first quantitative dietary constraints for five species of phytosaur. We furthermore explore the impacts of tooth position and cranial robustness on phytosaur microwear textures. We find subtle systematic texture differences between teeth from different positions along phytosaur tooth rows, which we interpret to be the result of different loading pressures experienced during food consumption, rather than functional partitioning of food processing along tooth rows. We find rougher microwear textures in morphologically robust taxa. This may be the result of seizing and processing larger prey items compared to those captured by gracile taxa, rather than dietary differences per se. We reveal relatively low dietary diversity between our study phytosaurs and that individual species show a lack of dietary specialization. Species are predominantly carnivorous and/or piscivorous, with two taxa exhibiting slight preferences for ‘harder’ invertebrates. Our results provide strong evidence for higher degrees of ecological convergence between phytosaurs and extant crocodilians than previously appreciated, furthering our understanding of the functioning and evolution of Triassic ecosystems.     

Functional morphology of the primate head and neck

The vertebral column plays a key role in maintaining posture, locomotion, and transmitting loads between body components. Cervical vertebrae act as a bridge between the torso and head and play a crucial role in the maintenance of head position and the visual field. Despite its importance in positional behaviors, the functional morphology of the cervical region remains poorly understood, particularly in comparison to the thoracic and lumbar sections of the spinal column. This study tests whether morphological variation in the primate cervical vertebrae correlates with differences in postural behavior. Phylogenetic generalized least-squares analyses were performed on a taxonomically broad sample of 26 extant primate taxa to test the link between vertebral morphology and posture. Kinematic data on primate head and neck postures were used instead of behavioral categories in an effort to provide a more direct analysis of our functional hypothesis. Results provide evidence for a function-form link between cervical vertebral shape and postural behaviors. Specifically, taxa with more pronograde heads and necks and less kyphotic orbits exhibit cervical vertebrae with longer spinous processes, indicating increased mechanical advantage for deep nuchal musculature, and craniocaudally longer vertebral bodies and more coronally oriented zygapophyseal articular facets, suggesting an emphasis on curve formation and maintenance within the cervical lordosis, coupled with a greater resistance to translation and ventral displacement. These results not only document support for functional relationships in cervical vertebrae features across a wide range of primate taxa, but highlight the utility of quantitative behavioral data in functional investigations. Am J Phys Anthropol 156:531–542, 2015. © 2015 Wiley Periodicals, Inc.     

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.     

Evaluation of "unique" aspects of human vertebral bodies and pedicles with a consideration of Australopithecus africanus

Recent functional studies of human vertebrae have revealed that loads borne by the axial skeleton during bipedal postures and locomotion pass through the pedicles and posterior elements as well as through the bodies and discs. Accordingly, particular morphological attributes of these vertebral elements have been linked exclusively with bipedalism. In order to test the validity of current form-function associations in human vertebral anatomy, this study considers the morphology of human thoracolumbar vertebral bodies and pedicles in the context of a wide comparative primate sample. The last lumbar vertebra of STS 14 (Australopithecus africanus) is also included in the analysis. Results indicate that certain features of human vertebrae previously thought to reflect bipedalism are characteristic of several nonhuman primates, including those whose posture is habitually pronograde. These features include the decrease in vertebral body surface area and the increase in cross-sectional area of the pedicle between the penultimate and last lumbar vertebra. In addition, although humans have relatively large and wide last lumbar pedicles, the enlargement and widening of the pedicle between the penultimate and last lumbar vertebra is not unique to humans. On the other hand, human vertebrae do exhibit several unique adaptations to bipedal posture and locomotion: (1) the vertebral body surface areas of the lower lumbar vertebrae and the cross-sectional areas of the last lumbar pedicles are large relative to body size, and (2) the last lumbar pedicles are wider relative to length and to body size than are those of nonhuman primates. The last lumbar vertebra of STS 14 does not exhibit any of these human-like vertebral features—its pedicles and body surface areas are relatively small, and its pedicles are not relatively wide, but relatively short.     

The first evidence of osteomyelitis in a sauropod dinosaur

Osteomyelitis is reported for the first time in a sauropod dinosaur. The material (MCS‐PV 183) comes from the Anacleto Formation (Campanian, Late Cretaceous), at the Cinco Saltos locality, Río Negro Province, Argentina. The specimen consists of 16 mid and mid‐distal caudal vertebrae of a titanosaur sauropod. Evidence of bacterial infection is preserved in all of these vertebrae. The main anomalies are as follows: irregular ‘microbubbly’ texture of bone surfaces produced by periosteal reactive bone, abscesses on the rims of the anterior articular surfaces of two centra, numerous pits on centra anterior articulation surfaces, erosions on the anterior articulation of the vertebral centra, a vertical groove in posterior articular face of all the centra and disruption of the prezygapophysis and postzygapophysis (mainly the articular face) from the vertebra 19 and beyond. The last anomaly is increasingly pronounced in more distal elements of the series. Thin sections reveal that the anomalous cortical tissue is composed of avascular and highly fibrous bone matrix. The fibres of the bone matrix are organized into thick bundles oriented in different directions. Both morphological and histological abnormalities in the MCS‐PV 183 specimen are pathognomonic for osteomyelitis.