New insights into the complex structure and ontogeny of the occipito‐vertebral gap in barbeled dragonfishes (Stomiidae,Teleostei)

In all stomiid genera there is an occipito‐vertebral gap between the skull and the first vertebra bridged only by the flexible notochord. Morphological studies from the early 20th century suggested that some stomiid genera have 1–10 of the anteriormost centra reduced or entire vertebrae missing in this region. Our study reviews this previous hypothesis. Using a new approach, we show that only in Chauliodus, Eustomias and Leptostomias gladiator vertebral centra are actually lost, with their respective neural arches and parapophyses persisting. We present results from a comparative analysis of the number and insertion sites of the anteriormost myosepta in 26 of the 28 stomiid genera. Generally in teleosts the first three myosepta are associated with the occiput, and the fourth is the first vertebral myoseptum. The insertion site of the fourth myoseptum plays an important role in this analysis, because it provides a landmark for the first vertebra. Lack of association of the fourth myoseptum with a vertebra is thus evidence that the first vertebra is reduced or absent. By counting the occipital and vertebral myosepta the number of reduced vertebrae in Chauliodus, Eustomias and Leptostomias gladiator can be inferred. Proper identification of the spino‐occipital nerves provides an additional source of information about vertebral reduction. In all other stomiid genera the extensive occipito‐ vertebral gap is not a consequence of the reduction of vertebrae, but of an elongation of the notochord. The complex structure and ontogeny of the anterior part of the vertebral column of stomiids are discussed comparatively. J. Morphol. 271:1006–1022, 2010. © 2010 Wiley‐Liss, Inc.     

Unique occipital articulation with the first vertebra found in pristigasterids, chirocentrids, and clupeids (Teleostei: Clupeiformes: Clupeoidei)

This study recognized a W-shaped occipital articulation associated with the first vertebra in pristigasterids, chirocentrids, and clupeids as a unique character among teleosts, based on an evaluation of 43 species within 40 genera of these three families of the Clupeoidei. This occipital articulation is accompanied by an anterior extension of the neural arch bases, which are autogenous with the first vertebral centrum. In chirocentrids and many of the clupeid species examined, the anterior extension occurs on the second vertebra, and similar occipital articulation is found between the first and second vertebrae. The W-shaped occipital articulation is not found in any other teleosts, including Denticeps (suborder Denticipitoidei), which is thought to be a sister group to the suborder Clupeoidei. The W-shaped occipital articulation is absent in the other family of the Clupeoidei, Engraulidae, based on an evaluation of 11 species in 10 genera. Instead, the convex anterior surface of the first vertebral centrum forms a condyle that articulates with the basioccipital, and the neural arches fuse with the centrum behind this condyle. Therefore, it is unclear whether the anterior extension of the first vertebral neural arch bases, which causes the W-shaped occipital articulation, occurs in engraulids. Based on an evaluation of the osteological development of Konosirus punctatus and Engraulis japonicus, the cartilaginous neural arch bases of the first and second vertebrae extend anteriorly at an early developmental stage in the former, whereas no anterior extension of the first vertebral neural arch bases occurs at any developmental stage in the latter. Therefore, the anterior extension of the neural arch bases, which causes the W-shaped occipital articulation, seems to be a unique character of pristigasterids, chirocentrids, and clupeids among teleosts. Within the recent phylogenetic context, this character may be a synapomorphy of these three families.     

Development of the osteocranium in the suckermouth armored catfish Ancistrus cf. triradiatus (Loricariidae, siluriformes)

The development of the osteocranium of the suckermouth armored catfish Ancistrus cf. triradiatus is described based on specimens ranging from prehatching stages to juvenile stages where the osteocranium is more or less fully formed. The first bony elements that arise are the opercle, jaws, and lateralmost branchiostegal rays, as well as the basioccipital and parasphenoid in the skull floor. The supracleithrum and the membranous and perichondral pterotic components form one large, double-layered skull bone during ontogeny, without clear evidence of the involvement of a supratemporal. The Baudelot's ligament ossifies from two sides, i.e., from the basioccipital medially and the supracleithrum laterally. The lower jaw consists of a dentary, mentomeckelian, and angulo-articular, which all soon fuse. The parurohyal, formed by the fusion of a ventral sesamoid bone and a dorsal cartilage element associated with the first basibranchial, is pierced by a vein, unlike in some other siluriforms. The interhyal cartilage disappears during ontogeny; medially of it, a small sesamoid bone appears in a ligament. The largest, canal-bearing cheek plate is not homologous to the interopercle. The results of the present research, with emphasis on bone formations and homologies, are compared with studies on related catfishes.     

A new candiru of the genus Paracanthopoma (Siluriformes: Trichomycteridae) from the Araguaia River basin,central Brazil

A new species of the candiru genus Paracanthopoma is described from the floodplains of the Bananal Island, a transition area between the Cerrado and Amazon, in the Araguaia River basin, central Brazil. Paracanthopoma cangussu sp. nov. is distinguished from its congeners, Paracanthopoma parva and Paracanthopoma saci, by the presence of seven opercular odontodes, five dentary teeth, five median premaxillary teeth, and first dorsal-fin pterygiophore in a vertical through the centrum of the 23th or 24th vertebra. It is further distinguished from each congener by an exclusive combination of character states, comprising the number of vertebrae, number of precaudal vertebrae, number of dorsal procurrent caudal-fin rays, number of ventral procurrent caudal-fin rays, number of dorsal-fin rays, disposition of pores on the cephalic portion of the latero-sensory system, absence of an anterior process on the anterior margin of parieto-supraoccipital, number of dorsal-fin pteryigiophores and number of interopercular odontodes. Although vandelliines are known for being exclusively hematophagous, with guts gorged with blood, two cleared and stained specimens of P. cangussu sp. nov. had Chironomidae larvae (Insecta) on their guts. Because most specimens of P. cangussu sp. nov. were collected with stomachs filled with blood, it was hypothesized that the species feeds accidentally or occasionally on insects.     

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.     

Development of the Synarcual in the Elephant Sharks (Holocephali; Chondrichthyes): Implications for Vertebral Formation and Fusion

The synarcual is a structure incorporating multiple elements of two or more anterior vertebrae of the axial skeleton, forming immediately posterior to the cranium. It has been convergently acquired in the fossil group ‘Placodermi’, in Chondrichthyes (Holocephali, Batoidea), within the teleost group Syngnathiformes, and to varying degrees in a range of mammalian taxa. In addition, cervical vertebral fusion presents as an abnormal pathology in a variety of human disorders. Vertebrae develop from axially arranged somites, so that fusion could result from a failure of somite segmentation early in development, or from later heterotopic development of intervertebral bone or cartilage. Examination of early developmental stages indicates that in the Batoidea and the ‘Placodermi’, individual vertebrae developed normally and only later become incorporated into the synarcual, implying regular somite segmentation and vertebral development. Here we show that in the holocephalan Callorhinchus milii, uniform and regular vertebral segmentation also occurs, with anterior individual vertebra developing separately with subsequent fusion into a synarcual. Vertebral elements forming directly behind the synarcual continue to be incorporated into the synarcual through growth. This appears to be a common pattern through the Vertebrata. Research into human disorders, presenting as cervical fusion at birth, focuses on gene misexpression studies in humans and other mammals such as the mouse. However, in chondrichthyans, vertebral fusion represents the normal morphology, moreover, taxa such Leucoraja (Batoidea) and Callorhinchus (Holocephali) are increasingly used as laboratory animals, and the Callorhinchus genome has been sequenced and is available for study. Our observations on synarcual development in three major groups of early jawed vertebrates indicate that fusion involves heterotopic cartilage and perichondral bone/mineralised cartilage developing outside the regular skeleton. We suggest that chondrichthyans have potential as ideal extant models for identifying the genes involved in these processes, for application to human skeletal heterotopic disorders.     

Evidence of Spondyloarthropathy in the Spine of a Phytosaur (Reptilia: Archosauriformes) from the Late Triassic of Halberstadt,Germany

Pathologies in the skeleton of phytosaurs, extinct archosauriform reptiles restricted to the Late Triassic, have only been rarely described. The only known postcranial pathologies of a phytosaur are two pairs of fused vertebrae of “Angistorhinopsis ruetimeyeri” from Halberstadt, Germany, as initially described by the paleontologist Friedrich von Huene. These pathologic vertebrae are redescribed in more detail in this study in the light of modern paleopathologic methods. Four different pathologic observations can be made in the vertebral column of this individual: 1) fusion of two thoracic vertebral bodies by new bone formation within the synovial membrane and articular capsule of the intervertebral joint; 2) fusion and conspicuous antero-posterior shortening of last presacral and first sacral vertebral bodies; 3) destruction and erosion of the anterior articular surface of the last presacral vertebra; and 4) a smooth depression on the ventral surface of the fused last presacral and first sacral vertebral bodies. Observations 1–3 can most plausibly and parsimoniously be attributed to one disease: spondyloarthropathy, an aseptic inflammatory process in which affected vertebrae show typical types of reactive new bone formation and erosion of subchondral bone. The kind of vertebral shortening present in the fused lumbosacral vertebrae suggests that the phytosaur acquired this disease in its early life. Observation 4, the smooth ventral depression in the fused lumbosacral vertebrae, is most probably not connected to the spondyloarthropathy, and can be regarded as a separate abnormality. It remains of uncertain origin, but may be the result of pressure, perhaps caused by a benign mass such as an aneurysm or cyst of unknown type. Reports of spondyloarthropathy in Paleozoic and Mesozoic reptiles are still exceptional, and our report of spondyloarthropathy in fossil material from Halberstadt is the first unequivocal occurrence of this disease in a Triassic tetrapod and in a phytosaur.     

Regionalization of axial skeleton in the lungfish Neoceratodus forsteri (Dipnoi)

Differentiation of the axial skeleton into distinct regions, once thought to be characteristic of the Tetrapoda, also occurs in the actinopterygian Danio rerio. In these taxa, the boundary between the cervical-thoracic regions correlates with Hoxc6 expression and morphological features such as position of the pectoral fin and associated nerves, and the absence of ribs. In the lungfish Neoceratodus, a member of the extant sister taxon to the Tetrapoda, the first vertebral element to chondrify is situated well posterior to the skull, developing from somites 6 and 7 (6/7) and associated with an enlarged cranial rib and nerves innervating the pectoral fin. Two vertebral elements develop later and more anteriorly, associated with somites 4/5 and 5/6. These three elements become incorporated into the occipital region of the skull during Neoceratodus ontogeny, until the cranial rib itself articulates to the rear of the skull. These features of early development indicate a regionalization of the Neoceratodus vertebral column: the cranial rib marks the boundary between the cervical and thoracic regions, the two more anterior vertebrae lacking ribs represent the cervical region, while somites 1-4 (cranial half), lacking any vertebral development, represent the occipital region. However, the cervical region of the vertebral column is effectively lost during ontogeny of Neoceratodus. A recognizable cervical region in the tetrapod vertebral column, as in zebrafish, suggests that cervical vertebrae are not incorporated into the skull but maintained as distinct elements of the column, representing an important shift in relative developmental timing and the influence of heterochrony in this region during the fish-tetrapod transition.     

内蒙古上白垩统二连组一长颈的镰刀龙类(英文)

镰刀龙类 (又称“懒龙”)是一类奇特的植食性兽脚类恐龙 ,化石记录主要局限于亚洲白垩纪地层中。由于镰刀龙类极其特化的形态和化石材料的局限性 ,这类恐龙的系统位置存在较多的争议。最近的发现 (RussellandDong ,1 994;Xuetal.,1 999)表明这类恐龙属于虚骨龙类 ,但其更为具体的系统位置依然存在争议 (Sues ,1 997;MakovickyandSues,1 998;Xuetal.,1 999;Sereno,1 999)。新发现于内蒙古苏尼特左旗赛罕高毕上白垩统二连组的镰刀龙类化石材料代表这类恐龙的一个新属种。杨氏内蒙古龙 (Neimongosaurusyangigen .etsp .nov .)的正型标本为一较为完整的骨架 ,是已知镰刀龙类当中第一件在同一个体中保存了大多数脊椎和几乎所有肢骨的标本。依据以下特征将内蒙古龙归入镰刀龙超科 :U形的下颌联合部、齿骨前端向下弯曲、齿骨前部没有牙齿、牙齿有一个收缩的基部、近圆形的齿根和叶形的齿冠、前部颈椎的神经脊低矮而轴向较长、后部颈椎背视呈X形、肱骨近端角状、肱骨有后转子、肱骨的尺骨髁和挠骨髁位于前部并为一狭窄槽分开、肠骨的耻骨柄细长而坐骨柄短以及跖部短。内蒙古龙的以下特征区别于其他镰刀龙类 :前部尾椎的横突下部有一圆形的窝 ,桡骨二头肌结节非常发育 ,后足趾节近端跟部非常发育 ,胫骨的腓骨嵴长     

Leis' conundrum: homology of the clavus of the ocean sunfishes. 2. Ontogeny of the median fins and axial skeleton of Ranzania laevis (Teleostei, Tetraodontiformes, Molidae)

One of the most conspicuous characters of the ocean sunfishes, family Molidae, is the punctuation of the body by a deep, abbreviated, caudal fin-like structure extending vertically between the posterior ends of the dorsal and anal fins, termed the clavus by Fraser Brunner. Homology of the clavus has been a matter of debate since the first studies on molid anatomy in the early 1800s. Two hypotheses have been proposed: 1) It is a highly modified caudal fin; 2) It is formed by highly modified elements of the dorsal and anal fins. To resolve this homology issue, we studied the ontogeny of the molid vertebral column and median fins and compared it to that of a less morphologically derived gymnodont (see Part 1 of this study), a member of the family Tetraodontidae. We show that in molids the chorda never flexes during development, that the claval rays form from the posterior ends of the dorsal and anal fins toward the middle, thus closing the gap inward, and that elements of the molid clavus have an identical development and composition as the proximal-middle and distal radials of the regular dorsal and anal fins. We thus conclude that the molid clavus is unequivocally formed by modified elements of the dorsal and anal fin and that the caudal fin is lost in molids.     

Variations of cervical vertebrae after expression of a Hox-1.1 transgene in mice

To understand the function of murine homeobox genes, a genetic analysis is mandatory. We generated gain-of-function mutants by introducing genomic sequences of the Hox-1.1 gene under the control of a chicken beta-actin promoter into mice. Our previous data had shown that these transgenic mice are nonviable after birth and are born with craniofacial abnormalities. In a subsequent detailed analysis of severely affected animals, malformations of the basioccipital bone, the atlas, and the axis were observed. Manifestation of an additional vertebra, a proatlas, occurred at the craniocervical transition. The dominant interference of the Hox-1.1 transgene with developmental programs seems to occur around day 9 of gestation, the time of neural crest migration and somite differentiation. We discuss the resulting phenotype with respect to a developmental control function of Hox-1.1.     

Ontogeny of the epicranial portion of the dorsal fin in Solea solea and Scophthalmus maximus (Teleostei, Pleuronectiformes)

A study of laboratory-reared larvae of Solea solea (Soleidae) and Scophthalmus maximus (Scophthalmidae) indicated that the epicranial portion of the dorsal fin results from the anterior displacement of proximal pterygiophores during ontogeny. The adult epicranial formula is attained early during ontogeny, and the anterior displacement is finalized after the passage of the migrating eye. In both species, the first two proximal pterygiophores fuse to form an erisma that is particularly long and well-developed in Solea. Moreover, in Solea, the neural spine of the second abdominal vertebra curves over the otic region, and the neural arch of the first vertebra remains incomplete. Received: August 3, 2000 / Revised: August 10, 2001 / Accepted: October 12, 2001     

Age estimation from stages of epiphyseal union in the presacral vertebrae

The presacral vertebrae have various secondary centers of ossification, whose timing of fusion can be used for age estimation of human skeletal remains up to the middle to the latter third decade. However, detailed information about the age at which these secondary centers of ossification fuse has been lacking. In this study, the timing of epiphyseal union in presacral vertebrae was studied in a sample of modern Portuguese skeletons (57 females and 47 males) between the ages of 9 and 30, taken from the Lisbon documented skeletal collection. A detailed photographic record of these epiphyses and the age ranges for the different stages of epiphyseal union are provided. Partial union of epiphyses was observed from 11 to 27 years of age. In general, centers of ossification begin to fuse first in the cervical and lumbar vertebrae, followed by centers of ossification in the thoracic region. The first center of ossification to complete fusion is usually that of the mammillary process in lumbar vertebrae. This is usually followed by that of the transverse process, spinous transverse process, and annular ring, regardless of vertebra type. There were no statistically significant sex differences in timing of fusion, but there was a trend toward early maturation in females for some vertebra or epiphyses. Bilateral epiphyses did not show statistically significant differences in timing of fusion. This study offers information on timing of fusion of diverse epiphyseal locations useful for age estimation of complete or fragmented human skeletal remains. Am J Phys Anthropol, 2011. © 2010 Wiley‐Liss, Inc.     

Congenital malformation of the skeleton in Weiser-Maples guinea pigs]

Some abnormalities were observed in the occipital bone, cervical vertebrae and thoracic vertebrae of Weiser-Maples guinea pigs. In the occipital bone, the medial basilar impression was suggested to occur in 40 (32.8%) out of 122 animals. The basilar impression was classified into right, left and both side types and observed in 24, 11 and 5 animals, respectively. The basilar impression was known to be accompanied in human with some anomalies such as platybasia, Klippel-Feil syndrome, deformation of foramen magnum and so-on. These anomalies were also observed in guinea pigs. The fusion of the axis with the 3rd cervical vertebra was observed in 12 (10.5%) out of 114 animals. The deformation was sometimes observed in the temporal, interparietal, atlas and axis as well as the occipital bone. The fusion of the 7th cervical vertebra with the 1st thoracic vertebra was found in 46 (51.7%) out of 89 animals. This fusion was thought to have no relation with the basilar impression. Weiser-Maples guinea pigs are now in 19 generations of sibmating. Because these abnormalities as mentioned above are all thought to be inherited, the selective breeding will make Weiser-Maples guinea pigs suitable for the study of the basilar impression.     

Demineralization of the vertebral skeleton in Atlantic salmon Salmo salar L. during spawning migration

In Atlantic salmon, Salmo salar, the mineral rate of vertebrae in a given fish varies according to the position of the vertebra along the rachidian axis. Indeed, the mean rate goes from 49% in the anterior vertebrae and raises to 51% in post-truncal vertebrae. Although no significant difference in the mineral rate was noticed between males and females either in the lower river basin or after spawning, the mineral rate of vertebral bone decreased significantly (1-2%) during spawning migration. Vertebrae, like scales, are an important reservoir of calcium from which fasting salmon draws the minerals and organic materials necessary for the substantial remodeling of cranial bones in males and for sexual maturation. We hypothesize that mineral decrease in vertebrae may be the result of a halastasic demineralization of the vertebral tissues.     

Characterization of a novel insertional mouse mutation, kkt: A closely linked modifier of Pax1

We describe a novel transgene insertional mouse mutant with skeletal abnormalities characterized by a kinked tail and severe curvature of the spine. The disrupted locus is designated kkt for "kyphoscoliosis kinked tail." Malformed vertebrae including bilateral ossification centers and premature fusion of the vertebral body to the pedicles are observed along the vertebral column, and the lower thoracic and lumbar vertebrae are the most affected. Some of the homozygous kkt neonates displayed two backward-pointing transverse processes in the sixth lumbar vertebra (L6) that resembled the first sacral vertebra, and some displayed one forward- and one backward-pointing transverse process in L6. The fourth and fifth sternebrae were also fused, and the acromion process of the scapula was missing in kkt mice. The skeletal abnormalities are similar to those observed in the mouse mutant undulated (un). The transgene is integrated at the distal end of chromosome 2 close to the Pax1 gene, as revealed by FISH analysis. However, mutation of the Pax1 gene is responsible for the un phenotype, but the Pax1 gene in the kkt mice is not rearranged or deleted. Pax1 is expressed normally in kkt embryos and in the thymus of mature animals, and there is no mutation in its coding sequence. Thus, the skeletal abnormalities observed in the kkt mutant are not due to a lack of functional Pax1. Mouse genomic sequences flanking the transgene and PAC clones spanning the wild-type kkt locus have been isolated, and reverse Northern analysis showed that the PACs contain transcribed sequence. Compound heterozygotes between un and kkt (un(+/-)/kkt(+/-)) display skeletal abnormalities similar to those of un or kkt homozygotes, but they have multiple lumbar vertebrae with a split vertebral body that is more severe than in homozygous un or kkt neonates. Furthermore, the sternebrae are not fused and no backward-pointing transverse processes are detected in L6. It is therefore apparent that these two mutations do not fully complement each other, and we propose that a gene in the kkt locus possesses a unique role that functions in concert with Pax1 during skeletal development.     

Demineralization of the vertebral skeleton in Atlantic salmon Salmo salar L. during spawning migration

In Atlantic salmon, Salmo salar, the mineral rate of vertebrae in a given fish varies according to the position of the vertebra along the rachidian axis. Indeed, the mean rate goes from 49% in the anterior vertebrae and raises to 51% in post-truncal vertebrae. Although no significant difference in the mineral rate was noticed between males and females either in the lower river basin or after spawning, the mineral rate of vertebral bone decreased significantly (1–2%) during spawning migration. Vertebrae, like scales, are an important reservoir of calcium from which fasting salmon draws the minerals and organic materials necessary for the substantial remodeling of cranial bones in males and for sexual maturation. We hypothesize that mineral decrease in vertebrae may be the result of a halastasic demineralization of the vertebral tissues.     

Ontogeny and homology of the claustra in otophysan Ostariophysi (Teleostei)

We studied the ontogeny of the claustrum comparatively in representatives of all otophysan subgroups. The claustrum of cypriniforms has a cartilaginous precursor, the claustral cartilage, which subsequently ossifies perichondrally at its anterior face and develops an extensive lamina of membrane bone. The membrane bone component of the claustrum and its close association with the atrium sinus imparis, a perilymphatic space of the Weberian apparatus, are both synapomorphies of cypriniforms. The characiform claustrum is not preformed in cartilage and originates as a membrane bone ossification, a putative synapomorphy of that taxon. Among siluriforms, the claustrum is present only in more basal groups and originates as an elongate cartilage that ossifies in a characteristic ventrodorsal direction, possibly a synapomorphy of catfishes. Gymnotiforms lack the claustral cartilage and claustrum. We review all previous hypothesis of claustrum homology in light of the above findings and conclude that the most plausible hypothesis is the one originally proposed by Bloch ([1900] Jen Z Naturw 34:1-64) that claustra are homologs of the supradorsals of the first vertebra.     

The Rutland Cetiosaurus: the anatomy and relationships of a Middle Jurassic British sauropod dinosaur

A relatively well–preserved specimen of Cetiosaurus oxoniensis, from the Middle Jurassic (Bajocian) of Rutland, United Kingdom, is described in detail. The material includes a nearly complete cervical series, representative dorsal vertebrae, a fragment of sacrum, anterior caudals, the right femur, and numerous rib and limb fragments. Contrary to previous suggestions that this specimen possesses 14 cervical and ten dorsal vertebrae, it seems more probable that there were at most 13 cervicals and at least 12 dorsals. The vertebral column displays several autapomorphic features which supplement the generic diagnosis of Cetiosaurus, including: (1) a stout, anteriorly directed process located at the top of the neural spine of the twelfth (?) cervical vertebra; and (2) the presence of lateral pits, separated by a thin midline septum, below the transverse processes of middle dorsal vertebrae. Cladistic analysis indicates that Cetiosaurus is probably the sister–taxon to the advanced neosauropod clade. This relationship affects the distribution of particular character states that have played an important role in determining sauropod phylogeny.