Cranial kinesis is the term for significant movement of skull bones relative to each other in addition to movement at the joint between the upper and lower jaw. It is usually taken to mean relative movement between the upper jaw and the braincase.
Most vertebrates have some form of kinetic skull. Cranial kinesis, or lack thereof, is usually linked to feeding. Animals which must exert powerful bite forces, such as crocodiles, often have rigid skulls with little or no kinesis, for maximum strength. Animals which swallow large prey whole (snakes), which grip awkwardly shaped food items (parrots eating nuts), or, most often, which feed in the water via suction feeding often have very kinetic skulls, frequently with numerous mobile joints. In the case of mammals, which have akinetic skulls (except for perhaps hares), the lack of kinesis is most likely to be related to the secondary palate, which prevents relative movement. This in turn is a consequence of the need to be able to create a suction during suckling.
Ancestry also plays a role in limiting or enabling cranial kinesis. Significant cranial kinesis is rare in mammals (the human skull shows no cranial kinesis at all). Birds have varying degrees of cranial kinesis, with parrots exhibiting the greatest degree. Among reptiles, crocodilians and turtles lack cranial kinesis, while lizards possess some, often minor, degree of kinesis and snakes possessing the most exceptional cranial kinesis of any tetrapod. In amphibians, cranial kinesis varies, but is unknown in frogs and rare in salamanders. Almost all fish have highly kinetic skulls, and teleost fish have developed the most kinetic skulls of any living organism.
Versluys (1910, 1912, 1936) classified types of cranial kinesis based on the location of the joint in the dorsal part of the skull.
Hofer (1949) further partitioned mesokinesis into
Streptostyly is the fore-aft movement of the quadrate about the otic joint (quadratosquamosal joint), although transverse movements may also be possible. Many hypothesized types of kinesis require basal joint kinesis (neurokinesis of Iordansky, 1990), that is, movement between the braincase and palate at the basipterygoid joint.
The first example of cranial kinesis was in the chondrichthyans, such as sharks. There is no attachment between the hyomandibular and the quadrate, and instead the hyoid arch suspends the two sets of jaws like pendulums. This allows sharks to swing their jaws outwards and forwards over the prey, allowing the synchronous meeting of the jaws and avoiding deflecting the prey when it comes close.
Actinopterygii (ray finned fish) possess a huge range of kinetic mechanisms. As a general trend through phylogenetic trees, there is a tendency to liberate more and more bony elements to allow greater skull motility. Most actinopts use kinesis to rapidly expand their buccal cavity, to create suction for suction feeding.
Early Dipnoi (lungfishes) had upper jaws fused to their braincase, which implies feeding on hard substrates. Many crossopterygian fishes had kinesis also.
Reptiles exhibit an extraordinary range of kinetic mechanisms, the most spectacular of which is snakes, who use highly kinetic joints to allow a huge gap; it is these highly kinetic joints that allow the wide gape and not the "unhinging" of joints, as many believe. Kinesis also prevents the "scissor effect", whereby the food item is pushed out of the mouth as the jaw occludes posteriorly to anteriorly. Typically, most modern reptile skulls are dikinetic, having both meta- and meso-kinetic joints. The mandibular bone is connected to the neurocranium via the quadrate and squamosal. The mandibulo-quadrate joint also articulates with the (palatine-pterygoid) bar which then connects to the maxilla, when the quadrate is pulled towards the skull by muscle x then the bar pushes on the base of the maxilla and causes the upper jaw to open.
The three principle types of kinesis found in Dinosaurs are,
Some show a combination of the two, such as streptostyly and prokinesis (Shuvuuia). Many, on the other hand, have at various points been thought to show akinesis, such as sauropods, ankylosaurs, and ceratopsians. It can be very difficult to prove that skulls were akinetic, and many of the above examples are contentious.
Pleurokinesis refers to the complex multiple jointing thought to occur in ornithopods, such as hadrosaurs. Ornithopod jaws are isognathic (meet simultaneously), working like a guillotine to slice plant material which can be manipulated with their teeth. However, because of the wedge shape of their teeth, the occlusional plane is tilted away from the centre of the head, causing the jaws to lock together and, due to the lack of a secondary palate, the force of this would not be braced. Because of this, Norman and Weishampel proposed a pleurokinetic skull. Here, there are four (or perhaps even more) kinetic parts of the skull,
As the lower jaw closes, the maxillojugal units move laterally producing a power stroke. These motions were later proved by a microwear analysis on an Edmontosaurus jaw.
Birds show a vast range of cranial kinetic hinges in their skulls. Zusi recognised three basic forms of cranial kinesis in birds,
Rhynchokinesis is further subdivided into double, distal, proximal, central and extensive. The older terms "schizorhynal" and "holorhynal" are generally synonymous with rhynchokinesis. In schizorhinal birds and most rhynchokinetic birds, the presence of two hinge axes at the base of the upper jaw imposes a requirement of bending within the jaw during kinesis. Bending takes different forms according to the number of hinges and their geometric configuration within the upper jaw. Proximal rhynchokinesis and distal rhynchokinesis apparently evolved from double rhynchokinesis by loss of different hinges. Extensive rhynchokinesis is an unusual and probably specialized variant. Kinesis in hummingbirds is still little understood.
Rhynchokinesis is an ability possessed by some birds to flex their upper beak or rhinotheca. Rhynchokinesis involves flexing at a point some way along the upper beak - either upwards, in which case the upper beak and lower beak or gnathotheca diverge, resembling a yawn, or downwards, in which case the tips of the beaks remain together while a gap opens up between them at their midpoint.
Unlike prokinesis, which is widespread in birds, rhynchokinesis is only known in cranes, shorebirds, swifts and hummingbirds. The adaptive significance of rhynchokinesis in certain non-probing birds is not yet known. It is hypothesized that the schizorhinal skull in proximally rhynchokinetic birds reflects ancestry, but has no adaptive explanation, in many living species.
Species in which this has been recorded photographically include the following species: short-billed dowitcher, marbled godwit, least sandpiper, common snipe, long-billed curlew, pectoral sandpiper, semipalmated sandpiper, Eurasian oystercatcher and bar-tailed godwit (see Chandler 2002 and external links).
Either prokinesis or some form of rhynchokinesis could be primitive for birds. Rhynchokinesis is not compatible with the presence of teeth in the bending zone of the ventral bar of the upper Jaw, and it probably evolved after their loss. Neognathous rhynchokinesis, however, probably evolved from prokinesis. The evolutionary origin of rhynchokinesis from prokinesis required selection for morphological changes that produced two hinge axes at the base of the upper jaw. Once evolved, the properties of these axes were subject to selection in relation to their effects on kinesis. The various forms of kinesis are hypothesized to have evolved by simple steps. In neognathous birds, prokinesis was probably ancestral to amphikinesis, and amphikinesis to rhynchokinesis in most cases, but prokinesis has also evolved secondarily.
In hares or "jackrabbits" (but not in their ancestors), there is a suture between regions in the fetal braincase that remains open in the adult, forming what is thought to be an intracranial joint, permitting relative motion between the anterior and posterior part of the braincase. It is thought that this helps absorb the force of impact as the hare strikes the ground.
Photographs of birds performing rhynchokinesis can be found here:
A very clear animation of pleurokinesis in Hadrosaurs can be found here:Aetiocetus
Aetiocetus is a genus of extinct basal mysticete, or baleen whale that lived 33.9 to 23.03 million years ago, in the late Oligocene in the North Pacific ocean, around Japan, Mexico, and Oregon, U.S. It was first described by Douglas Emlong in 1966 and currently contains known four species, A. cotylalveus, A. polydentatus, A. tomitai, and A. weltoni. These whales are remarkable for their retention of teeth and presence of nutrient foramina, indicating that they possessed baleen. Thus, Aetiocetus represents the transition from teeth to baleen in Oligocene mysticetes. Baleen is a highly derived character, or synapomorphy, of mysticetes, and is a keratinous structure that grows from the palate, or roof of the mouth, of the whale. The presence of baleen is inferred from the fossil record in the skull of Aetiocetus.
Aetiocetus is known from both sides of the Pacific Ocean: it was first documented in Oregon, United States, but it is also known from Japan and Mexico. The genus is currently constrained to the Northern hemisphere and has little value in biostratigraphic studies of the Oligocene due to its limited occurrences across the Pacific.Allosauridae
Allosauridae is a family of medium to large bipedal, carnivorous allosauroid neotheropod dinosaurs from the Late Jurassic. Allosauridae is a fairly old taxonomic group, having been first named by the American paleontologist Othniel Charles March in 1878. Allosaurids are characterized by an astragalus with a restriction of the ascending process to the lateral part of the bone, a larger medial than lateral condyle, and a horizontal groove across the face of the condyles.Batrachomorpha
Batrachomorpha ("frog form") is a name traditionally given to recent and extinct amphibians that are more closely related to modern amphibians than they are to reptiles. It most often includes the extinct groups Temnospondyli and Lepospondyli. The first tetrapods were all amphibians in the physiological sense that they laid their eggs in water, and are colloquially sometimes refereed to as labyrinthodonts or stegocephalians. In this scheme, batrachomorphs composed one branch of these early amphibians, while the reptiliomorphs composed the other. While the actual phylogeny of the modern amphibians is not well understood, their ancestors are descended from one line of batrachomorphs. All other living tetrapods (reptiles, birds and mammals) are descended from one branch of reptiliomorphs, the amniotes. Amniotes achieved dominance, while all other reptiliomorphs and most batrachomorphs have gone extinct.Beak
The beak, bill, or rostrum is an external anatomical structure of birds that is used for eating and for preening, manipulating objects, killing prey, fighting, probing for food, courtship and feeding young. The terms beak and rostrum are also used to refer to a similar mouth part in some ornithischians, pterosaurs, turtles, cetaceans, dicynodonts, anuran tadpoles, sirens, pufferfishes, billfishes and cephalopods.
Although beaks vary significantly in size, shape, color and texture, they share a similar underlying structure. Two bony projections—the upper and lower mandibles—are covered with a thin keratinized layer of epidermis known as the rhamphotheca. In most species, two holes known as nares lead to the respiratory system.Bird anatomy
Bird anatomy, or the physiological structure of birds' bodies, shows many unique adaptations, mostly aiding flight. Birds have a light skeletal system and light but powerful musculature which, along with circulatory and respiratory systems capable of very high metabolic rates and oxygen supply, permit the bird to fly. The development of a beak has led to evolution of a specially adapted digestive system. These anatomical specializations have earned birds their own class in the vertebrate phylum.Ceratosaurus
Ceratosaurus (from Greek κέρας/κέρατος, keras/keratos meaning "horn" and σαῦρος/sauros meaning "lizard") was a carnivorous theropod dinosaur in the Late Jurassic period (Kimmeridgian to Tithonian). This genus was first described in 1884 by American paleontologist Othniel Charles Marsh based on a nearly complete skeleton discovered in Garden Park, Colorado, in rocks belonging to the Morrison Formation. The type species is Ceratosaurus nasicornis.
The Garden Park specimen remains the most complete skeleton known from the genus, and only a handful of additional specimens have been described since. Two additional species, Ceratosaurus dentisulcatus and Ceratosaurus magnicornis, have been described in 2000 from two fragmentary skeletons from the Cleveland-Lloyd Quarry of Utah and from the vicinity of Fruita, Colorado. The validity of these additional species has been questioned, however, and all three skeletons possibly represent different growth stages of the same species. In 1999, the discovery of the first juvenile specimen was reported. Since 2000, a partial specimen was excavated and described from the Lourinhã Formation of Portugal, providing evidence for the presence of the genus outside of North America. Fragmentary remains have also been reported from Tanzania, Uruguay, and Switzerland, although their assignment to Ceratosaurus is currently not accepted by most paleontologists.
Ceratosaurus was a medium-sized theropod. The original specimen is estimated to be 5.3 m (17 ft) or 5.69 m (18.7 ft) long, while the specimen described as C. dentisulcatus was larger, at around 7 m (23 ft) long. Ceratosaurus was characterized by deep jaws that supported proportionally very long, blade-like teeth, a prominent, ridge-like horn on the midline of the snout, and a pair of horns over the eyes. The forelimbs were very short, but remained fully functional; the hand had four fingers. The tail was deep from top to bottom. A row of small osteoderms (skin bones) was present down the middle of the neck, back, and tail. Additional osteoderms were present at unknown positions on the animal's body.
Ceratosaurus gives its name to the Ceratosauria, a clade of theropod dinosaurs that diverged early from the evolutionary lineage leading to modern birds. Within the Ceratosauria, some paleontologists proposed it to be most closely related to Genyodectes from Argentina, which shares the strongly elongated teeth. The geologically older genus Proceratosaurus from England, although originally described as a presumed antecedent of Ceratosaurus, was later found to be unrelated. Ceratosaurus shared its habitat with other large theropod genera including Torvosaurus and Allosaurus, and it has been suggested that these theropods occupied different ecological niches to reduce competition. Ceratosaurus may have preyed upon plant-eating dinosaurs, although some paleontologists suggested that it hunted aquatic prey such as fish. The nasal horn was probably not used as a weapon as was originally suggested by Marsh, but more likely was used solely for display.Cosesaurus
Cosesaurus is a genus of archosauromorph reptiles likely belonging to the family Tanystropheidae. It is known from fossil imprints of a single small skeleton, MGB V1, which was found in Muschelkalk outcrops near the municipalities of Mont-ral and Alcover in Spain. These outcrops are dated to the Ladinian age of the middle Triassic about 242 to 237 million years ago. The specimen is stored at the Museu Martorell (a.k.a. the Museu Geologia de Barcelona), which is now part of the Museu de Ciències Naturals de Barcelona. The poor preservation and likely juvenile nature of the specimen has led to the anatomy of Cosesaurus being misidentified by several different sources. For example, Paul Ellenberger claimed that it was an ancestor to birds in the 1970s, while Dave Peters claimed that it was a pterosaur ancestor in 2000. Both of these claims contrast with mainstream scientific theories on the origins of either group, and other paleontologists who study the specimen are unable to find the features which Ellenberger or Peters reported to be present. The Ellenberger and Peters hypotheses are thus considered fringe theories with questionable scientific soundness due to their low reproducibility. Mainstream hypotheses for the relations of Cosesaurus generally agree that it is a "protorosaur", specifically a tanystropheid closely related to long-necked reptiles such as Macrocnemus, Tanytrachelos, Tanystropheus, or Langobardisaurus.Elpistostegalia
Elpistostegalia or Panderichthyida is an order of prehistoric lobe-finned fishes which lived during the Late Devonian period (about 385 to 374 million years ago). They represent the advanced tetrapodomorph stock, the fishes more closely related to tetrapods than the osteolepiform fishes. The earliest elpistostegalians, combining fishlike and tetrapod-like characters, are sometimes called fishapods, a phrase coined for the advanced elpistostegalian Tiktaalik.Enantiornithes
Enantiornithes is a group of extinct avialans ("birds" in the broad sense), the most abundant and diverse group known from the Mesozoic era. Almost all retained teeth and clawed fingers on each wing, but otherwise looked much like modern birds externally. Over 80 species of enantiornitheans have been named, but some names represent only single bones, so it is likely that not all are valid. Enantiornitheans became extinct at the Cretaceous–Paleogene boundary, along with hesperornithids and all other non-avian dinosaurs. Enantiornitheans are thought to have left no living descendants.Gerrothorax
Gerrothorax ("wicker chest") is an extinct genus of temnospondyl amphibian from the Triassic period of Greenland, Germany, Sweden, and possibly Thailand. It is known from a single species, G. pulcherrimus, although several other species such as G. pustuloglomeratus have been named in the past.
Gerrothorax was about 1 metre (3.3 ft) long, and had a remarkably flattened body. It probably hid under sand or mud on river and lake bottoms, scanning for prey with its large, upward-facing eyes. Gerrothorax had an unusually shaped skull with angular protrusions on the sides. This looked vaguely similar to the skull of the earlier, unrelated, amphibian Diplocaulus, but was not so developed.Some Gerrothorax fossils preserved hypobranchials and ceratobranchials (bony gill arches) near the neck. This shows that Gerrothorax was pedomorphic, retaining its larval gills as an adult. When originally described in 1946, these bones were considered to correspond to feather-like external gills similar to those of modern-day neotenic salamanders, such as the mudpuppy, the axolotl, and the olm.However, a 2011 paper found that it was more likely that plagiosaurids such as Gerrothorax had internal gills, like those of fish, rather than salamander-like external gills. The authors of that study noted that plagiosaurids and other ancient amphibians which retained gills as adults had grooves on their ceratobranchials. Grooved ceratobranchials are present in both modern and ancient fish, but unknown in modern amphibians. Therefore, they were indicative of internal gills. This would have also been advantageous for survival in large animals, as internal gills would have been protected by a large skin fold and were less likely to have been damaged by the environment.A 2008 study showed that Gerrothorax lifted its head rather than dropping its jaw when catching prey, which has been compared to how a toilet seat opens. In 2011 the skull of Gerrothorax was scanned using microtomography, revealing that the braincase and palatoquadrate regions are highly ossified. A 2013 study argued that Gerrothorax consumed prey using suction feeding. Gerrothorax had strong muscles capable of both raising the cranium and lowering the jaw rapidly. The robust internal gill apparatus would have expelled water through the gills during this motion, creating intense pressure in the throat that would suck in small prey items. The gill arches were also covered in small denticles, prohibiting any prey from escaping once devoured. Although suction feeding is common in fish and modern larval amphibians, Gerrothorax differs from these animals by its lack of cranial kinesis, meaning that its cranial bones could not flex against each other to envelop prey.The fossil record of Gerrothorax pulcherrimus extends 35 million years from the Ladinian stage of the Middle Triassic to the Rhaetian stage of the Late Triassic. Throughout this time span, specimens of the species show few morphologic differences, making G. pulcherrimus an extreme example of evolutionary stasis. G. pulcherrimus may have remained unchanged for so long because it could tolerate a wide range of ecological conditions. Although it always needed to live in an aquatic habitat, G. pulcherrimus may have been able to live in a variety of different water bodies with a wide range of salinity.Glossary of dinosaur anatomy
This glossary explains technical terms commonly employed in the description of dinosaur body fossils. Besides dinosaur-specific terms, it covers terms with wider usage, when these are of central importance in the study of dinosaurs or when their discussion in the context of dinosaurs is beneficial. The glossary does not cover ichnological and bone histological terms, nor does it cover measurements.Gobivenator
Gobivenator is an extinct genus of troodontid theropod dinosaur known from the late Campanian Djadokhta Formation of central Gobi Desert, Mongolia. It contains a single species, Gobivenator mongoliensis. G. mongoliensis is known from a single individual, which represents the most complete specimen of a Late Cretaceous troodontid currently known.Heterochrony
In evolutionary developmental biology, heterochrony is a developmental change in the timing or rate of events, leading to changes in size and shape. It is contrasted with heterotopy, a change in spatial positioning of some process in the embryo, which can also create morphological innovation. Following the framework of Reilly et al, heterochrony can be divided into intraspecific heterochrony that explains variation within a species, and interspecific heterochrony that explains developmental variation phylogenetically, in the timing or rate of events of a descendent species with respect to an ancestral species.
These changes all affect the start, end, rate, or timespan of a particular developmental process. The concept of heterochrony was introduced by Ernst Haeckel in 1875, and given its modern sense by Gavin de Beer in 1930.Jaw
The jaw is any opposable articulated structure at the entrance of the mouth, typically used for grasping and manipulating food. The term jaws is also broadly applied to the whole of the structures constituting the vault of the mouth and serving to open and close it and is part of the body plan of most animals.Owlet-nightjar
Owlet-nightjars are small crepuscular birds related to the nightjars and frogmouths. Most are native to New Guinea, but some species extend to Australia, the Moluccas, and New Caledonia. A flightless species from New Zealand is extinct. There is a single monotypic family Aegothelidae with the genus Aegotheles.
Owlet-nightjars are insectivores which hunt mostly in the air but sometimes on the ground; their soft plumage is a cryptic mixture of browns and paler shades, they have fairly small, weak feet (but larger and stronger than those of a frogmouth or a nightjar), a tiny bill that opens extraordinarily wide, surrounded by prominent whiskers. The wings are short, with 10 primaries and about 11 secondaries; the tail long and rounded.Prolacerta
Prolacerta is a genus of archosauromorph from the lower Triassic of South Africa and Antarctica. The only known species is Prolacerta broomi. The generic name Prolacerta is derived from Latin meaning “before lizard” and its species name broomi is in commemoration of the famous paleontologist Robert Broom, who discovered and studied many of the fossils found in rocks of the Karoo Supergroup. When first discovered, Prolacerta was considered to be ancestral to modern lizards, scientifically known as lacertilians. However, a study by Gow (1975) instead found that it shared more similarities with the lineage that would lead to archosaurs such as crocodilians and dinosaurs (including birds). Prolacerta is considered by modern paleontologists to be among the closest relatives of the Archosauriformes.Sanajeh
Sanajeh (meaning "ancient gape" in Sanskrit) is a genus of late Cretaceous madtsoiid snake from western India. A fossil described in 2010 from the Lameta Formation was found coiled around an egg and an adjacent skeleton of a 50 cm (19 in) long sauropod dinosaur hatchling. This suggests that the snake preyed on hatchling sauropods at nesting sites.Sandpiper
Sandpipers are a large family, Scolopacidae, of waders or shorebirds. They include many species called sandpipers, as well as those called by names such as curlew and snipe. The majority of these species eat small invertebrates picked out of the mud or soil. Different lengths of bills enable different species to feed in the same habitat, particularly on the coast, without direct competition for food.
Sandpipers have long bodies and legs, and narrow wings. Most species have a narrow bill, but otherwise the form and length are quite variable. They are small to medium-sized birds, measuring 12 to 66 cm (4.7–26.0 in) cm in length. The bills are sensitive, allowing the birds to feel the mud and sand as they probe for food. They generally have dull plumage, with cryptic brown, grey, or streaked patterns, although some display brighter colours during the breeding season.Most species nest in open areas, and defend their territories with aerial displays. The nest itself is a simple scrape in the ground, in which the bird typically lays three or four eggs. The young of most species are precocial.Zephyrosaurus
Zephyrosaurus (meaning "westward wind lizard") is a genus of orodromine ornithischian dinosaur. It is based on a partial skull and postcranial fragments discovered in the Aptian-Albian-age Lower Cretaceous Cloverly Formation of Carbon County, Montana, USA. New remains are under description, and tracks from Maryland and Virginia, also in the USA, have been attributed to animals similar to Zephyrosarus.