Cranial Crest Research Articles

Aerodynamic Characteristics of the Crest with Membrane Attachment on Cretaceous Pterodactyloid Nyctosaurus

Abstract:The Nyctosaurus specimen KJ1 was reconstructed under the hypothesis that there is a membrane attached to the crest; the so‐called headsail crest. The aerodynamic forces and moment acting on the headsail crest were analyzed. It was shown that KJ1 might adjust the angle of the headsail crest relative to the air current as one way to generate thrust (one of the aerodynamic forces, used to overcome body drag in forward flight) and that the magnitude of the thrust and moment could vary with the gesture angle and the relative location between the aerodynamic center of the headsail crest and body's center of gravity. Three scenarios were tested for comparison: the crest with membrane attachment, the crest without membrane attachment and the absence of a cranial crest. It was shown that the aerodynamic characteristics (increasing, maintaining and decreasing thrusts and moment) would have almost disappear in flight for the crest without membrane attachment and was non‐existent without the cranial crest. It is suggested from aerodynamics evidence alone that Nyctosaurus specimen KJ1 had a membrane attached to the crest and used this reconstructed form for auxiliary flight control.

Persistent expression of Pax3 in the neural crest causes cleft palate and defective osteogenesis in mice

Transcription factors regulate tissue patterning and cell fate determination during development; however, expression of early regulators frequently abates upon differentiation, suggesting that they may also play a role in maintaining an undifferentiated phenotype. The transcription factor paired box 3 (Pax3) is expressed by multipotent neural crest precursors and is implicated in neural crest disorders in humans such as Waardenburg syndrome. Pax3 is required for development of multiple neural crest lineages and for activation of lineage-specific programs, yet expression is generally extinguished once neural crest cells migrate from the dorsal neural tube and differentiate. Using a murine Cre-inducible system, we asked whether persistent Pax3 expression in neural crest derivatives would affect development or patterning. We found that persistent expression of Pax3 in cranial neural crest cells resulted in cleft palate, ocular defects, malformation of the sphenoid bone, and perinatal lethality. Furthermore, we demonstrated that Pax3 directly regulates expression of Sostdc1, a soluble inhibitor of bone morphogenetic protein (BMP) signaling. Persistent Pax3 expression renders the cranial crest resistant to BMP-induced osteogenesis. Thus, one mechanism by which Pax3 maintains the undifferentiated state of neural crest mesenchyme may be to block responsiveness to differentiation signals from the environment. These studies provide in vivo evidence for the importance of Pax3 downregulation during differentiation of multipotent neural crest precursors and cranial development.

A new Triassic pterosaur from Switzerland (Central Austroalpine, Grisons), Raeticodactylus filisurensis gen. et sp. nov.

A new basal non-pterodactyloid pterosaur, Raeticodactylus filisurensis gen. et sp. nov., is reported. It has been discovered in shallow marine sediments from the Upper Triassic of the lowest Kössen beds (late Norian/early Rhaetian boundary) in the central Austroalpine of Canton Grisons (Switzerland). The disarticulated specimen is comprised of an almost complete skull and a partial postcranial skeleton. A high and thin bony, sagittal cranial crest characterizes the anterodorsal region of the skull. The large mandible, with an additional keel-like expansion at the front, partly matches the enlarged sagittal cranial crest. A direct and close relationship to Austriadactylus cristatus, the only known Triassic pterosaur with a bony cranial crest so far, cannot be established. The teeth of the premaxilla are monocuspid and exhibit very strongly bowed enamel wrinkles on the lingual side whereas the enamel is smooth on the labial side. These monocuspid teeth are large and fang-like. The numerous smaller teeth of the maxilla show three, four and five cusps. These are very similar to the teeth of the Triassic pterosaur Eudimorphodon ranzii. The humerus shows a thinner construction than that seen in other Triassic pterosaurs. The femur is quite unusual with a caput femoris perpendicular to the shaft. The bones of the extremities are almost twice as long as the ones from the largest Triassic specimen E. ranzii (MCSNB 2888). The newly described pterosaur is an adult, with a wingspan of approximately 135 cm. A morphofunctional analysis suggests that R. filisurensis was a highly specialized piscivore and possibly a skim-feeder.

Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution

Department of Biological Sciences, University of Alberta, CW 405 Biological Sciences Centre, Edmonton, Alberta, CanadaReceived March 2005; accepted for publication March 2007Cryolophosaurus ellioti Hammer and Hickerson, 1994, from the Early Jurassic Hanson Formation of the CentralTransantarctic Mountains, represents a theropod dinosaur from a period of time and geographical area that arepoorly sampled with respect to dinosaur taxa. An in-depth morphological description of Cryolophosaurus is pre-sented here, along with a rigorous phylogenetic analysis of theropod relationships consisting of 347 characters and56 taxa, in an attempt to clarify the relationships of Cryolophosaurus and to provide insight into questions sur-rounding early theropod evolution. Cryolophosaurus is characterized by a unique cranial crest, formed primarily bythe lacrimals, a pronounced constriction of the squamosal and jugal bones across the infratemporal fenestra, andextremely elongate cranial processes on the cervical ribs. Several shared characters, including the presence of a slot-shaped foramen at the base of the nasal process of the premaxilla, nasolacrimal crests and erect tab-like dorsal pro-cesses on the articular, suggest affinities between Cryolophosaurus and a clade of medium-bodied Early Jurassictheropods that includes ‘Dilophosaurus’ sinensis, Dracovenator regenti and Dilophosaurus wetherilli. This clade isrecovered as sister-taxon to a Neoceratosauria + Tetanurae clade, rendering both a traditional Coelophysoidea andCeratosauria non-monophyletic. Cryolophosaurus represents the largest known Early Jurassic theropod, and marksthe beginning of theropod occupation of the dominant predator niche in the Mesozoic. © 2007 The Linnean Societyof London, Zoological Journal of the Linnean Society, 2007, 151, 377–421.

The development of the neural crest in the human.

The first systematic account of the neural crest in the human has been prepared after an investigation of 185 serially sectioned staged embryos, aided by graphic reconstructions. As many as fourteen named topographical subdivisions of the crest were identified and eight of them give origin to ganglia (Table 2). Significant findings in the human include the following. (1) An indication of mesencephalic neural crest is discernible already at stage 9, and trigeminal, facial, and postotic components can be detected at stage 10. (2) Crest was not observed at the level of diencephalon 2. Although pre-otic crest from the neural folds is at first continuous (stage 10), crest-free zones are soon observable (stage 11) in Rh.1, 3, and 5. (3) Emigration of cranial neural crest from the neural folds at the neurosomatic junction begins before closure of the rostral neuropore, and later crest cells do not accumulate above the neural tube. (4) The trigeminal, facial, glossopharyngeal and vagal ganglia, which develop from crest that emigrates before the neural folds have fused, continue to receive contributions from the roof plate of the neural tube after fusion of the folds. (5) The nasal crest and the terminalis-vomeronasal complex are the last components of the cranial crest to appear (at stage 13) and they persist longer. (6) The optic, mesencephalic, isthmic, accessory, and hypoglossal crest do not form ganglia. Cervical ganglion 1 is separated early from the neural crest and is not a Froriep ganglion. (7) The cranial ganglia derived from neural crest show a specific relationship to individual neuromeres, and rhombomeres are better landmarks than the otic primordium, which descends during stages 9-14. (8) Epipharyngeal placodes of the pharyngeal arches contribute to cranial ganglia, although that of arch 1 is not typical. (9) The neural crest from rhombomeres 6 and 7 that migrates to pharyngeal arch 3 and from there rostrad to the truncus arteriosus at stage 12 is identified here, for the first time in the human, as the cardiac crest. (10) The hypoglossal crest provides cells that accompany those of myotomes 1-4 and form the hypoglossal cell cord at stages 13 and 14. (11) The occipital crest, which is related to somites 1-4 in the human, differs from the spinal mainly in that it does not develop ganglia. (12) The occipital and spinal portions of the crest migrate dorsoventrad and appear to traverse the sclerotomes before the differentiation into loose and dense zones in the latter. (13) Embryonic examples of synophthalmia and anencephaly are cited to emphasize the role of the neural crest in the development of cranial ganglia and the skull.

Anatomy and Relationships of Lambeosaurus magnicristatus, a crested hadrosaurid dinosaur (Ornithischia) from the Dinosaur Park Formation, Alberta

ABSTRACT The first detailed description of the lambeosaurine Lambeosaurus magnicristatus (Ornithischia: Hadrosauridae) confirms that it is a distinct taxon characterized by its comparatively enormous cranial crest, formed predominantly by the caudodorsal process of the premaxilla, and an acute crest-snout angle. The holotype of L. magnicristatus occurs stratigraphically higher than all other Dinosaur Park Formation lambeosaurines at the Dinosaur Provincial Park locality. The only referred specimen was collected over 170 kilometers southeast of the type locality. Correlation of its host stratum with the well-known Dinosaur Park section reveals that L. magnicristatus has no biostratigraphic overlap with L. lambei and suggests that it replaces L. lambei on a regional scale in southern Alberta at the end of ‘Dinosaur Park time.’ Species-level phylogenetic analysis of Lambeosaurinae corroborates the monophyly of Lambeosaurus. The genus is characterized by five apomorphies, including a procumbent crest, complete enclosure of the ophthalmic canal of the laterosphenoid, the presence of a flange on the caudodorsal process of the premaxilla that overlaps the nasal in the rostral region of the crest, caudal extension of the premaxilla such that it forms the caudal margin of the crest, and a unique joint between the rostral nasal and the caudodorsal process of the premaxilla. Lambeosaurine phylogeny indicates that the development of a hypertrophied cranial crest evolved independently at least three times within the clade, suggesting that the crest enlargement is a recurring evolutionary trend within Lambeosaurinae.

Pax7 identifies neural crest, chromatophore lineages and pigment stem cells during zebrafish development

Using immunostaining during early zebrafish embryogenesis, we report that the cranial and trunk neural crest expresses the paired box protein Pax7, thus revealing a novel neural crest marker in zebrafish. In the head, we show that Pax7 is broadly expressed in the cranial crest cells, which indicates that duplication of the paralogous group Pax3/7 at the origin of vertebrates included the conserved expression of Pax7 in the head neural crest of all of the vertebrates species studied so far. In the trunk, Pax7 recognizes both premigratory and migratory neural crest cells. Notably, we observed the expression of Pax7 during the development of melanophore, xanthophore and iridophore precursor cells. In contrast to the case of melanocyte precursors in birds, Pax7 showed overlapping expression with early melanin pigment. Finally, during the larva to adult transition, we show that pigment stem cells recapitulate the expression of Pax7.

A juvenile Parasaurolophus (Ornithischia: Hadrosauridae) Braincase from Dinosaur Provincial Park, Alberta, with comments on crest ontogeny in the genus

ABSTRACT An incomplete braincase of a juvenile lambeosaurine hadrosaurid (Dinosauria: Ornithischia) is recognized as the second and smallest cranial specimen of Parasaurolophus from the Belly River Group (Campanian), Alberta, Canada. This specimen provides the first information on the ontogeny of the skull roof and autapomorphic tubular crest in this widely recognized yet poorly known taxon. In addition to the distinctive morphology of the cranial crest, Parasaurolophus is characterized by a suite of skull roof characters associated with crest development that manifest at small size, including verticalization of the nasal-frontal joint at the base of the crest. Although the crest is not preserved in the specimen described here, the distinctively thickened and steeply angled frontal platform indicates that the crest and facial profile were significantly different from the equivalent juvenile stages of corythosaurin lambeosaurines (Corythosaurus, Hypacrosaurus, Lambeosaurus), which resemble one another closely. Parasaurolophus therefore appears to deviate from the well-known corythosaurin mode of cranial crest growth early in ontogenetic development.

Juvenile specimens of the pterosaur Germanodactylus cristatus, with a review of the genus

ABSTRACT Two juvenile specimens of the pterodactyloid pterosaur Germanodactylus cristatus are identified from the Solnhofen Limestone of southern Germany. The specimens had been referred to the nominal species Pterodactylus kochi and P. micronyx. They have edentulous tips on the upper and lower jaws; their skulls are taller and less elongate and have fewer teeth than similarly sized specimens assigned to P. antiquus and P. micronyx; and they have tall neural spines on their cervical vertebrae. In addition, they exhibit a distinctive suite of skeletal proportions that distinguishes them from similarly sized specimens assigned to P. antiquus and P. micronyx. The specimens do not exhibit any trace of a cranial crest, which supports the interpretation that the cranial crest developed late in ontogeny, probably as the individual approached skeletal and/or sexual maturity. The taxonomy of Germanodactylus is reviewed and revised diagnoses are presented.

New Material of Pterosaur Sinopterus (Reptilia: Pterosauria) from the Early Cretaceous Jiufotang Formation, Western Liaoning, China

Abstract: Based on a new nearly complete postcranial skeleton of an adult specimen of Sinopterus from the Early Cretaceous Jiufotang Formation of western Liaoning, China, the diagnosis of Sinopterus is amended. The revised diagnosis of Sinopterus includes skull relatively elongate with weakly developed cranial crest; ratio of the length of Ph2d4 to that of Ph1d4 is approximately 0.73; ratio of the length of wing metacarpal to that of metatarsal 3 is 4.5; ratio of the length of mt3 to that of tibia is approximately 0.21, and wing phalanges 1 and 2 are straight. Comparison between the ratios of the limb bones between non‐adult and adult individuals of Sinopterus indicates that during the ontogenetic process, some ratios between bones are constant, such as the first wing phalanx to the second wing phalanx, the wing metacarpal to the metatarsal 3, metatarsal 3 to the tibia, but others are not, such as humerus to wing metacarpal and femur to tibia, in which the former grows faster than the latter.

Functional analysis of Sox8 during neural crest development inXenopus

Among the families of transcription factors expressed at the neural plate border, Sox proteins have been shown to regulate multiple aspects of neural crest development. Sox8, Sox9 and Sox10, exhibit overlapping expression domains in neural crest progenitors, and studies in mouse suggest that Sox8 functions redundantly with Sox9 and Sox10 during neural crest development. Here, we show that in Xenopus, Sox8 accumulates at the lateral edges of the neural plate at the mid-gastrula stage; in contrast to its mouse and chick orthologs, Sox8 expression precedes that of Sox9 and Sox10 in neural crest progenitors. Later in development, Sox8 expression persists in migrating cranial crest cells as they populate the pharyngeal arches and in trunk neural crest cells, in a pattern that recapitulates both Sox9 and Sox10 expression domains. Although morpholino-mediated knockdown of Sox8 protein did not prevent the formation of neural crest progenitors, the timing of their induction was severely affected. This delay in neural crest specification had dramatic consequences on the development of multiple lineages of the neural crest. We demonstrate that these defects are due to the inability of neural crest cells to migrate into the periphery, rather than to a deficiency in neural crest progenitors specification and survival. These results indicate that the control of Sox8 expression at the neural plate border is a key process in initiating neural crest formation in Xenopus, and highlight species-specific differences in the relative importance of SoxE proteins during neural crest development.

Cranial crest development in the Azhdarchoid pterosaur Tupuxuara, with a review of the genus and tapejarid monophyly

Abstract: A portion of pterosaur skull from the Romualdo Member of the Santana Formation (?Albian–?Turonian, Cretaceous) of north‐east Brazil provides new data on the morphology and ontogeny of azhdarchoid pterosaur cranial crests. The specimen consists of parts of the cranial bones posterodorsal to the nasoantorbital fenestra, including partial nasals, lacrimals, frontals and possibly the parietals. A posterodorsally directed premaxillary crest with a concave posterior border is located dorsal to the posterior border of the nasoantorbital fenestra. A well‐defined suture indicates overlapping, posterodorsally directed growth of the premaxilla over the skull roof, suggesting that the generation of the premaxillary crest is a late ontogenetic feature and thus probably related to sexual display. The systematics of Tupuxuara and its relationship to other azhdarchoids is reviewed and a cladistic analysis of the group is presented. Tupuxuara is found to be the sister‐taxon to Azhdarchidae. Tupuxuara longicristatus Kellner and Campos, 1988 is argued to be the only valid named species in this genus and Thalassodromeus Kellner and Campos, 2002 is considered a junior subjective synonym of this taxon. As originally conceived, Tapejaridae is paraphyletic: a new, more restrictive version of Tapejaridae (including Tapejara and Sinopterus dongi) might exist, but its monophyly is weakly supported. Furthermore, Tapejara was found to be paraphyletic in all trees.

A basal tyrannosauroid dinosaur from the Late Jurassic of China

The tyrannosauroid fossil record is mainly restricted to Cretaceous sediments of Laurasia, although some very fragmentary Jurassic specimens have been referred to this group. Here we report a new basal tyrannosauroid, Guanlong wucaii gen. et sp. nov., from the lower Upper Jurassic of the Junggar Basin, northwestern China. G. wucaii is the oldest known tyrannosauroid and shows several unexpectedly primitive pelvic features. Nevertheless, the limbs of G. wucaii share several features with derived coelurosaurs, and it possesses features shared by other coelurosaurian clades. This unusual combination of character states provides an insight into the poorly known early radiation of the Coelurosauria. Notably, the presumed predatory Guanlong has a large, fragile and highly pneumatic cranial crest that is among the most elaborate known in any non-avian dinosaur and could be comparable to some classical exaggerated ornamental traits among vertebrates.

Nasal cavity homologies and cranial crest function in lambeosaurine dinosaurs

Abstract A neurological method for assessment of nasal cavity homologies in extant archosaurs is extended to lambeosaurine hadrosaurids (Dinosauria: Ornithischia) to test functional hypotheses associated with their hypertrophied nasal passages and highly derived cranial crests. The olfactory system and associated cranial nerve pathways that have consistent relationships to soft tissue divisions of the nasal cavity are reconstructed in lambeosaurines on the basis of new paleoneurological data and a comparative phylogenetic approach. The new model of the lambeosaurine olfactory system and nasal cavity shows that a significant portion of nasal cavity proper was located outside the crest cavities and that the primary olfactory region was located rostromedial to the orbits. All available data indicate that the evolutionary hypertrophy of the nasal cavity occurred predominantly within the non-olfactory nasal vestibule, and that crest development was not causally associated with olfaction. The high level of inte...

Nasal cavity homologies and cranial crest function in lambeosaurine dinosaurs

Nasal cavity homologies and cranial crest function in lambeosaurine dinosaurs

The ligamentum olecrani of the Elbow Joint in Dogs and Cats

The olecranon ligament (ligamentum olecrani) is described as an elastic ligament of the elbow joint in carnivores that tenses the caudomedial part of the joint capsule. The aim of the study was to compare the course and the microscopic structure of the ligament in dogs and cats. The elbow regions of 25 dogs and 15 cats were dissected to examine the topography of the ligament in extension and flexion. Furthermore, the olecranon ligaments of five dogs and five cats were studied using routine histological methods. Additional sections were stained with Resorcin–Fuchsin and Orcein to detect elastic fibres. In both species the olecranon ligament originates at the lateral surface of the epicondylus medialis humeri and inserts at the cranial crest of the olecranon extending distally to the roof of the processus anconeus. Tension of the ligament only occurs when the elbow joint is flexed maximally. This tension is increased by a slight supination of the forearm, which takes place automatically in this joint position. In dogs the ligament is long (30–40 mm in medium sized breeds) and relatively slim (approx. 2–4 mm). In cats the ligament is short (10–12 mm) and relatively strong (5–8 mm). The histological examination of the olecranon ligament shows all signs of a tight collagenous ligament with a negligible amount of elastic fibres. The olecranon ligament helps to limit the maximal flexion of the elbow joint. In addition, it controls a slight lateral movement of the processus anconeus during the automatic supination of the antebrachial bones in extreme flexion of the elbow joint.

Redefining the head–trunk interface for the neural crest

The head–trunk interface lies at the occipito-cervical boundary, which corresponds to the somite 5/6 level. Previous studies have demonstrated that neural crest cells also behave differently either side of this boundary and that this may be due to intrinsic differences between cranial and trunk crest. However, it is also possible that some of the observed differences between cranial and trunk crest are assigned by environmental cues. We have therefore scrutinised the behaviour of the neural crest cells generated either side of the occipito-cervical boundary in chick and, interestingly, find that both behave in a truncal fashion by traversing the anterior half of their adjacent somites. Furthermore, although not previously described, we find that transient DRGs form opposite somites 4 and 5. Crest cells produced anterior of the somite 3/4 boundary avoid the somites and behave in a non-truncal fashion; these cells populate the pharyngeal arches, and thus contribute to the developing head. We have further shown, via somite transplantations, that differential behaviour of the posterior versus anterior occipital crest is assigned by the somites. If somites 1 to 3 are replaced by trunk somites, then the anterior occipital crest will behave in a truncal fashion by invading the somites. Correspondingly, if these anterior occipital somites are transplanted in place of trunk somites, they perturb the migration of trunk crest. Thus, for the neural crest, the head–trunk interface does not lie at the occipito-cervical boundary, but rather lies at the somite 3/4 level and is defined by the somites. The fact that this boundary lies at the somite 3/4 level in chick is significant as it reflects the more ancient posterior occipital boundary; in fish, only the first three somites contribute to the occipital bone.

Significance of the cranial neural crest.

The cranial neural crest has long been viewed as being of particular significance. First, it has been held that the cranial neural crest has a morphogenetic role, acting to coordinate the development of the pharyngeal arches. By contrast, the trunk crest seems to play a more subservient role in terms of embryonic patterning. Second, the cranial crest not only generates neurons, glia, and melanocytes, but additionally forms skeletal derivatives (bones, cartilage, and teeth, as well as smooth muscle and connective tissue), and this potential was thought to be a unique feature of the cranial crest. Recently, however, several studies have suggested that the cranial neural crest may not be so influential in terms of patterning, nor so exceptional in the derivatives that it makes. It is now becoming clear that the morphogenesis of the pharyngeal arches is largely driven by the pharyngeal endoderm. Furthermore, it is now apparent that trunk neural crest cells have skeletal potential. However, it has now been demonstrated that a key role for the cranial neural crest streams is to organise the innervation of the hindbrain by the cranial sensory ganglia. Thus, in the past few years, our views of the significance of the cranial neural crest for head development have been altered. Developmental Dynamics 229:5-13, 2004.

New crested specimens of the Late Cretaceous pterosaurNyctosaurus

Two recently collected specimens of the Late Cretaceous pterosaurNyctosaurus differ from all previously known specimens in the possession of a large branching cranial crest. The crest extends upward and backward from the posterior skull roof and is nearly three times the length of the skull proper. Despite the large crest, the specimens do not differ significantly in morphology from previously known specimens ofNyctosaurus, and do not seem to represent a new species ofNyctosaurus. The specimens suggest that the cranial crest was developed late in ontogeny, which is consistent with the interpretation of pterosaur cranial crests as intraspecific display structures.

A new species of tapejarid pterosaur with soft-tissue head crest

Abstract Two specimens of a tapejarid pterosaur (Pterodactyloidea, Tapejaridae) are described as representing a new species. Both specimens show evidence for soft tissues preserved in association with a sagittal bony cranial crest. Both specimens are from the Nova Olinda Member Konservat Lagerstätte of the Crato Formation of the Araripe Basin, northeastern Brazil. They represent the second tapejarid species from this formation. Comparisons are made with other crested pterosaurs and comments on the utility and aerodynamics of pterosaurian head crests are made.